Phytogeography of seaweeds of the Azores

189 species of seaweeds have been recorded for the Atlantic archipelago of the Azores (114 Rhodophyceae, 41 Phaeophyceae and 34 Chlorophyceae). Ten of these have been described as endemic algae. The taxonomic status of these “endemics” is far from clear, however. Studies on the relatedness of this seaweed flora to seaweed floras of surrounding areas using clustering methods indicate its intermediary position between the seaweed floras of the subtropical Macaronesian Islands (Canaries Madeira and Salvages) on the one side and those of the warm temperate Eurafrican coasts on the other side. The geographic position of the Azores is NW of the other Macaronesian Archipelagos and thus nearer to the American coast. The Gulf stream may easily transport seaweeds from the Carribean to the Azores. Nevertheless, the number of species of these islands in common with the American Atlantic coast is much lower than with, for example, the W. Mediterranean. Of one group of seaweeds, viz. algae with a strictly warm temperature distribution, not a single species occurs likewise on American coasts. This absence is probably explicable because of the narrow temperature range for survival of these seaweeds. Temperatures of surface waters around the Azores during pleistocene glaciations were not much lower than they are at present. Nevertheless, it can be supposed that a number of seaweeds now occurring in the Azores have become extinct during the last glaciation. Paper presented at the XIV International Botanical Congress (Berlin, 24 July–1 August, 1987), Symposium 6-15, “Biogeography of marine benthic algae”. CANCAP-project. Contributions to the zoology, botany and paleontology of the Canarian-Cape Verdean region of the North Atlantic Ocean. No. 67.     

Subantarctic flowering plants: pre‐glacial survivors or post‐glacial immigrants?

Aim The aim here was to assess whether the present‐day assemblage of subantarctic flowering plants is the result of a rapid post‐Last Glacial Maximum (LGM) colonization or whether subantarctic flowering plants survived on the islands in glacial refugia throughout the LGM. Location The circumpolar subantarctic region, comprising six remote islands and island groups between latitudes 46° and 55° S, including South Georgia in the South Atlantic Ocean, the Prince Edward Islands, Îles Crozet, Îles Kerguelen, the Heard Island group in the South Indian Ocean and Macquarie Island in the South Pacific Ocean. Methods Floristic affinities between the subantarctic islands were assessed by cluster analysis applied to an up‐to‐date dataset of the phanerogamic flora in order to test for the existence of provincialism within the subantarctic. A review of the primary literature on the palaeobotany, geology and glacial history of the subantarctic islands was carried out and supplemented with additional palaeobotanical data and new field observations from South Georgia, Île de la Possession (Îles Crozet) and Îles Kerguelen. Results First, a strong regionalism was observed, with different floras characterizing the islands in each of the ocean basins, and endemic species being present in the South Indian Ocean and South Pacific Ocean provinces. Second, the majority of the plant species were present at the onset of accumulation of post‐glacial organic sediment and there is no evidence for the natural arrival of new immigrants during the subsequent period. Third, a review of geomorphological data suggested that the ice cover was incomplete during the LGM on the majority of the islands, and ice‐free biological refugia were probably present even on the most glaciated islands. Main conclusions Several independent lines of evidence favour the survival of a native subantarctic phanerogamic flora in local refugia during the LGM rather than a post‐LGM colonization from more distant temperate landmasses in the Southern Hemisphere.     

Phylogeny and colonization history of Pringlea antiscorbutica (Brassicaceae), an emblematic endemic from the South Indian Ocean Province

The origins and evolution of sub-Antarctic island floras are not well understood. In particular there is uncertainty about the ages of the contemporary floras and the ultimate origins of the lineages they contain. Pringlea R. Br. (Brassicaceae) is a monotypic genus endemic to four sub-Antarctic island groups in the southern Indian Ocean. Here we used sequences from both the chloroplast and nuclear genomes to examine the phylogenetic position of this enigmatic genus. Our analyses confirm that Pringlea falls within the tribe Thelypodieae and provide a preliminary view of its relationships within the group. Divergence time estimates and ancestral area reconstructions imply Pringlea diverged from a South American ancestor ～5Myr ago. It remains unclear whether the ancestor of Pringlea dispersed directly to the South Indian Ocean Province (SIOP) or used Antarctica as a stepping-stone; what is clear, however, is that following arrival in the SIOP several additional long-distance dispersal events must be inferred to explain the current distribution of this species. Our analyses also suggest that although Pringlea is likely to have inherited cold tolerance from its closest relatives, the distinctive morphology of this species evolved only after it split from the South American lineage. More generally, our results lend support to the hypothesis that angiosperms persisted on the sub-Antarctic islands throughout the Pliocene and Pleistocene. Taken together with evidence from other sub-Antarctic island plant groups, they suggest the extant flora of sub-Antarctic is likely to have been assembled over a broad time period and from lineages with distinctive biogeographic histories.     

Floristic turnover in Iceland from 15 to 6 Ma – extracting biogeographical signals from fossil floral assemblages

Aim This study aims to document the floristic changes that occurred in Iceland between 15 and 6 Ma and to establish the dispersal mechanisms for the plant taxa encountered. Using changing patterns of dispersal, two factors controlling floristic changes are tested. Possible factors are (1) climate change, and (2) the changing biogeography of Iceland over the time interval studied; that is, the presence or absence of a Miocene North Atlantic Land Bridge. Location The North Atlantic. Methods Species lists of fossil plants from Iceland in the time period 15 to 6 Ma were compiled using published data and new data. Closest living analogues were used to establish dispersal properties for the fossil taxa. Dispersal mechanisms of fossil plants were then used to reconstruct how Iceland was colonized during various periods. Results Miocene floras of Iceland (15–6 Ma) show relatively high floristic turnover from the oldest floras towards the youngest; and few taxa from the oldest floras persist in the younger floras. The frequencies of the various dispersal mechanisms seen in the 15‐Ma floras are quite different from those recorded in the 6‐Ma floras, and there is a gradual change in the prevailing mode of dispersal from short‐distance anemochory and dyschory to long‐distance anemochory. Two mechanisms can be used to explain changing floral composition: (1) climate change, and (2) the interaction between the dispersal mechanisms of plants and the increasing isolation of proto‐Iceland during the Miocene. Main conclusions Dispersal mechanisms can be used to extract palaeogeographic signals from fossil floras. The composition of floras and dispersal mechanisms indicate that Iceland was connected both to Greenland and to Europe in the early Middle Miocene, allowing transcontinental migration. The change in prevalence of dispersal modes from 15 to 6 Ma appears to reflect the break‐up of a land bridge and the increasing isolation of Iceland after 12 Ma. Concurrent gradual cooling and isolation caused changes in species composition. Specifically, the widening of the North Atlantic Ocean prevented taxa with limited dispersal capability from colonizing Iceland, while climate cooling led to the extinction of thermophilous taxa.     

THE BIOGEOGRAPHIC ORIGIN OF ARCTIC ENDEMIC SEAWEEDS: A THERMOGEOGRAPHIC VIEW1

The Arctic is geologically and biogeographically young, and the origin of its seaweed flora has been widely debated. The Arctic littoral biogeographic region dates from the latest Tertiary and Pleistocene. Following the opening of Bering Strait, about 3.5 mya, the “Great Trans‐Arctic Biotic Interchange” populated the Arctic with a fauna strongly dominated by species of North Pacific origin. The Thermogeographic Model (TM) demonstrates why climate and geography continued to support this pattern in the Pleistocene. Thus, Arctic and Atlantic subarctic species of seaweeds are likely to be evolutionarily “based” in the North Pacific, subarctic species are likely to be widespread in the warmer Arctic, and species of Atlantic Boreal or warmer origin are unlikely in the Arctic and Subarctic. Although Arctic seaweeds have been thought to have a greater affinity with the North Atlantic, we have reanalyzed the Arctic endemic algal flora, using the Thermogeographic Model and evolutionary trees based on molecular data, to demonstrate otherwise. There are 35 congeneric species of the six, abundant Arctic Rhodophyta that we treat in this paper; 32 of these species (91%) occur in the North Pacific, two species (6%) occur in the Boreal or warmer Atlantic Ocean, and a single species is panoceanic, but restricted to the Subarctic. Laminaria solidungula J. Agardh, a kelp Arctic “endemic” species, has 18 sister species. While only eleven (61%) occur in the North Pacific, this rapidly dispersing and evolving genus is a terminal member of a diverse family and order (Laminariales) widely accepted to have evolved in the North Pacific. Thus, both the physical/time‐based TM and the dominant biogeographic pattern of relatives of Arctic macrophytes suggest strong compliance with the evidence of zoology, geology, and paleoclimatology that the Arctic marine flora is largely of Pacific origin.     

Hawaiian angiosperm radiations of North American origin

Background

Putative phytogeographical links between America (especially North America) and the Hawaiian Islands have figured prominently in disagreement and debate about the origin of Pacific floras and the efficacy of long-distance (oversea) plant dispersal, given the obstacles to explaining such major disjunctions by vicariance.

Scope

Review of past efforts, and of progress over the last 20 years, toward understanding relationships of Hawaiian angiosperms allows for a historically informed re-evaluation of the American (New World) contribution to Hawaiian diversity and evolutionary activity of American lineages in an insular setting.

Conclusions

Temperate and boreal North America is a much more important source of Hawaiian flora than suggested by most 20th century authorities on Pacific plant life, such as Fosberg and Skottsberg. Early views of evolution as too slow to account for divergence of highly distinctive endemics within the Hawaiian geological time frame evidently impeded biogeographical understanding, as did lack of appreciation for the importance of rare, often biotically mediated dispersal events and ecological opportunity in island ecosystems. Molecular phylogenetic evidence for North American ancestry of Hawaiian plant radiations, such as the silversword alliance, mints, sanicles, violets, schiedeas and spurges, underlines the potential of long-distance dispersal to shape floras, in accordance with hypotheses championed by Carlquist. Characteristics important to colonization of the islands, such as dispersibility by birds and ancestral hybridization or polyploidy, and ecological opportunities associated with ‘sky islands’ of temperate or boreal climate in the tropical Hawaiian archipelago may have been key to extensive diversification of endemic lineages of North American origin that are among the most species-rich clades of Hawaiian plants. Evident youth of flowering-plant lineages from North America is highly consistent with recent geological evidence for lack of high-elevation settings in the Hawaiian chain immediately prior to formation of the oldest, modern high-elevation island, Kaua‘i.     

Additive partitioning as a tool for investigating the flora diversity in oceanic archipelagos

This paper introduces the integration of additive partitioning with species—area relationships to island biogeography in order to address the question “How are the pteridophyte and spermatophyte native and endemic flora of different oceanic archipelagos partitioned across islands?”.Species richness data of all endemic species and all native species of pteridophytes and spermatophytes were obtained for the Azores, Canaries and Cape Verde in the Atlantic Ocean and Galápagos, Hawaii and Marquesas in the Pacific Ocean. Additive partitioning of species diversity was used to quantify how much of the total diversity of an oceanic archipelago flora (γ-diversity) is due to (i) the mean species richness of the flora of each island (α-diversity), (ii) the variability in species richness of the floras across islands (β_Nestedness) and (iii) the complementarity in species composition of the floras of different islands (β_Replacement). The analysis was separately performed for the native and endemic pteridophyte and spermatophyte floras.The diversity partitioning of the six archipelagos showed large differences in how the flora of each archipelago is partitioned among the α, β_Nestedness and β_Replacement components, for pteridophytes and spermatophytes and for all endemic species and all native species. The α-diversity was more important for all native species than for endemic species and more important for pteridophytes than for spermatophytes, with the Azores showing outstanding high values of α-diversity. The β_Nestedness was higher for pteridophytes than for spermatophytes and higher for endemic species than for all native species in both pteridophytes and spermatophytes. The values of β_Replacement suggested that: (i) the spermatophyte native flora is more differentiated across islands than the pteridophyte native flora and (ii) the pteridophyte endemic flora and, especially, the spermatophyte endemic flora are more differentiated across islands than the corresponding native flora. An outstanding value of β_Replacement for endemic and all native spermatophytes was found in Hawaii, confirming the biogeographical island differentiation in this archipelago.     

The Bering Strait connection: dispersal and speciation in boreal macroalgae

A large number of boreal seaweeds have either sibling species or conspecific populations of a single species in the North Pacific and North Atlantic Oceans. This pattern is thought to have arisen from the dispersal between the two oceans through the Arctic Ocean after the opening of the Bering Strait in the mid-to-late Miocene or earliest Pliocene and from subsequent vicariant speciation as the Arctic Ocean froze and Bering Strait closed intermittently during glacial periods. Recent molecular studies of species in all three major seaweed phyla reveal patterns of vicariance. However, a number of lines of evidence point to differences in origins of these clades; some appear to be Pacific in origin whereas others appear to be derived from Atlantic stock. Different origins can be explained by recent stratigraphic finds that push the first Cenozoic opening of the Bering Strait back from 3.1–4.1 to 4.8–7.4 Ma (million years ago). Northern hemisphere ocean circulation models suggest that water flow would have been from the North Atlantic–Arctic south through the Bering Strait prior to the closure of the Panamanian Isthmus c. 3.5 Ma in contrast to the northward flow from the Pacific into the Arctic and North Atlantic, which developed after the closing of the Isthmus. Despite these differences in timing of the two invasions, there are no significant differences in levels of relationships among species with a North Atlantic origin compared with species with a North Pacific origin based on currently available data. More work is required to understand vicariance in seaweeds, especially in deciphering when a speciation event has occurred.     

Not across the North Pole: plant migration in the Arctic

? The vascular plant flora of 66 arctic islands was studied to determine whether the islands have been occupied by random long-distance dispersal (LDD) or in a highly structured northward migration pattern via intervening islands as stepping-stones. ? A maximum parsimonious migration model minimizing dispersal distances of 1256 vascular plant taxa was calculated in the framework of network analysis. ? Plant dispersal is not stochastic in the Arctic at the global scale. Inferred mean dispersal distances of the plants occurring on arctic islands are c. 580 km (median 460 km). A LDD across the North Pole could not be inferred in the model and species may be recruited mainly from the nearest mainland or islands. At smaller scales, among adjacent islands, dispersal of vascular plants may be incomplete. Arctic islands do not yet appearto be saturated with species. ? The results suggest that changes in biodiversity in Arctic islands can be more easily predicted at the global scale than at the local scale. Because islands are not yet saturated with species, new colonizations may not necessarily be linked to climate change.     

Trans-oceanic dispersal and evolution of early composites (Asteraceae)

How did Asteraceae (the daisy family) expand from its area of origin and become so widespread? This question has challenged generations of evolutionary botanists. Molecular phylogenetic and biogeographic analyses indicate a South American origin of Asteraceae, a view supported by the recent discovery of the earliest fossils of the family in Middle Eocene (ca. 50 Ma) deposits in southern South America. The early-branching lineages in the phylogenetic tree of Asteraceae are South American and African, suggesting that the earliest successful colonization of areas outside South America may have involved long-distance dispersal to Africa. However, one particularly challenging unanswered question is how early members of Asteraceae reached Africa at a time when the Atlantic Ocean constituted a barrier between the two continents. Morphological, phylogenetic, geographic, paleogeographic, and paleontologic data have been combined to propose scenarios on possible geographical and dispersal routes and vectors of dispersion of early-branching lineages of Asteraceae from South America to Africa. Of the different scenarios proposed here, two concern alternative geographical routes: (1) via the Rio Grande Rise-Walvis Ridge axis in the South Atlantic; or (2) via Antarctica, possibly including the Subantarctic islands. Three scenarios consider different dispersal routes: (1) stepping-stones; (2) single-step; and (3) sweepstakes. Finally, three vectors of dispersion are considered: (1) birds; (2) wind; and (3) floating islands. Evaluation of these scenarios suggests that early-branching lineages of Asteraceae probably dispersed from South America to Africa along an island chain formed by the Rio Grande Rise and the Walvis Ridge, transported by birds, possibly combined with rafting and/or sweepstakes. Morphological changes typically associated with evolution on islands characterize many African carduoid descendants, providing indirect evidence for step-wise dispersal along the island chain.     

World-wide latitudinal and longitudinal seaweed distribution patterns and their possible causes,as illustrated by the distribution of Rhodophytan genera

The degree of similarity between red algal generic floras in each pair of 22 climatically defined biogeographic regions was established on a world-wide scale by Jaccard's similarity index and by an hierarchical clustering with an agglomerative centroid method. Two clusterings were carried out, the first one on the basis of all 637 genera, and the second one on the basis of genera not occurring in the tropics and non-endemic to any one of the 22 regioms (145 genera). This latter clustering served to detect better the relationships among non-tropical floras. The results indicate the following division of the earth's rhodophytan seaweed floras: (1) A rich tropical-warm temperate "Tethyan" group including the rich tropical Indo W Pacific and W Atlantic floras, and the rich warm temperate NW Pacific and NE Atlantic floras; (2) the depauperate extensions of the above group (the tropical E Pacific and E Atlantic floras, and the warm temperate NW and SW Atlantic floras); (3) a cold temperate and a warm temperate N Pacific group; (4) an Arctic-cold temperate N Atlantic group and a NE Atlantic warm temperate flora; (5) an Antarctic-cold temperate southern hemisphere group including the cold temperate SE Pacific, SW Atlantic, SE Atlantic floras, and the Antarctic flora; (6) the two highly individual, but slightly related warm temperate SE Atlantic flora (S. Africa) and SW Pacific flora (Southern Australia and Northern New Zealand); (7) the depauperate warm temperate SE Pacific flora. Although the northern and southern hemisphere temperate and polar floras are quite unrelated (on the basis of genera lacking in the tropics), they share nonetheless a number of cool water genera which apparently have succeeded in passing the adverse tropical belt. The rich tropical-warm temperate group is thought to consist of vicariant portions of a formerly continuous Tethyan flora. The N Pacific and N Atlantic temperate floras are thought to have developed independently since the Oligocene (~ 40.10⁶ y) deterioration of the climate and to have partially mixed their cool water genera only after the Pliocene inundation (2.10⁶ y) of the Bering Land Bridge. The warm-temperate floras of S Africa and southern Australia probably owe their richness and individuality to a very long isolation (already at the start of the Cenozoicum) and a continued residence in warm temperate conditions with small seasonal fluctuations.Paper presented at the Seaweed Biogeography Workshop of the International Working Group on Seaweed Biogeography, held from 3–7 April 1984, at the Department of Marine Biology, University of Groningen (The Netherlands). Convenor: C. van den Hoek.     

Origin of Macquarie Island echinoderms

The origin of echinoderms from Macquarie Island in the Southern Ocean is analysed through a novel application of multivariate statistics. Ordinations are produced from a combination of species distribution, bathymetric, habitat and life history data in order to assess patterns of migration. The analyses distinguish groups of species derived from the Kerguelen Plateau, New Zealand and eastern Antarctica. These groups correlate with attributes expected for epiplanktonic dispersal and range expansion along the North and South Macquarie Ridges respectively. There is no convincing evidence for long-distance pelagic dispersal, migration from the abyssal plain or for human translocation of species. The results indicate that taxonomic groups differ in their ability to disperse, and emphasize the importance of depth in biogeographical analyses. Dispersal by range expansion appears to have been more significant than epiplanktonic dispersal and vicariant rather than long-distance dispersal mechanisms are the preferred explanation for some disjunct distribution patterns. Received: 4 December 1997 / Accepted: 11 April 1998     

The biogeography of seaweeds in Southeast Alaska

Aim This article reviews the history of seaweed collections in Southeast Alaska from the early Russian explorers to contemporary efforts. It summarizes other studies of Southeast Alaskan seaweeds from a biogeographical perspective, and compares the known seaweed flora near three population centres (Ketchikan, Sitka and Juneau) with those of other regions within Alaska, and with nearby regions. Location For this article, Southeast Alaska includes all inside and outside waters of the Alexander Archipelago from Dixon Entrance (54°40′ N, 133°00′ W) to Icy Point (58°23′10″ N, 137°04′20″ W). Methods The literature on seaweeds occurring in Southeast Alaska is reviewed from a biogeographical perspective, and herbarium records for Southeast Alaska from the Alaska Seaweed Database project are used to provide an overview of the biogeography of the area. Records for the population centres of Ketchikan, Sitka and Juneau are compared with records from other areas within Alaska and with nearby regions to determine floristic similarities. Results Southeast Alaska has the most diverse seaweed flora of any region of Alaska. A list of species known to occur in Southeast Alaska is appended (in Supplementary Material) and includes their reported occurrences in three population centres (Juneau, Ketchikan and Sitka). Recognition of at least three distinct biogeographical areas associated with these three centres is supported by a comparison of their floras with those of other regions in the North Pacific. A close relationship of some species with conspecifics in the north‐west Atlantic is also noted. In contrast, ecological, physiological and genetic differentiation of Southeast Alaskan seaweeds from conspecifics in Washington State or even from different areas of Southeast Alaska are documented. A ShoreZone coastal habitat system, which is being implemented to inventory and map the entire shoreline of Southeast Alaska, is defining new biogeographical units called ‘bioareas’ on the basis of the distribution of canopy kelps and lower intertidal algal assemblages. Main conclusions Southeast Alaska has the most diverse seaweed flora of any region of Alaska. This is a reflection of its extensive coastline, with varied past and present environmental conditions. Different parts of Southeast Alaska show similarities to different areas outside Southeast Alaska. Despite this, much remains to be learned about the biogeography of seaweeds in Southeast Alaska, and many questions remain to be answered.     

Incidence of dioecy in relation to growth form,pollination and dispersal

Summary Recent theories predict the evolution of dioecy among higher plants, in association with certain pollination and fruit dispersal traits. However, reported associations of dioecy with pollination, dispersal and growth form traits have not distinguished the effects of each trait separately, controlling for the others. Because these traits are associated among themselves, existing analyses may involve spurious or indirect correlations. This paper reports the incidence of dioecy in a subarctic and an arctic flora, and analyzes the occurrence of dioecy among vascular plants classified jointly by growth form, floral (pollination) syndrome, and fruit (dispersal) syndrome. Dioecism is no more frequent in the arctic flora as a whole, but its incidence increases northward among woody plants. This increase is associated with an increase in the proportion of woody species having small, inconspicuous flowers, and not with the syndrome of fleshy or animal dispersed fruits. Within the floras of Alaska, California, and the Northeastern US, dioecy is markedly more frequent among woody plants and among plants having small, inconspicuous flowers, and that is the only strong statistical association of dioecy for the species of these floras. When genera and families are analyzed similarly, dioecy is also associated significantly with dispersal syndrome. Thus, among angiosperms, evidence currently does not support either an uniquely strong or exclusive association of dioecy with dispersal traits, as it does for gymnosperms (Givnish 1980). It is extremely desirable to analyze the occurrence of dioecy among taxa classified jointly by all relevant ecological traits, rather than analyzing marginal distributions.     

New Zealand's pteridophyte flora—Plants of ancient lineage but recent arrival?

A hypothesis is presented that most pteridophytes arrived in New Zealand relatively recently, by long-distance dispersal. The flora comprises 194 native species, of which 89 (46%) are endemic and 105 (54%) are widespread. Of the latter, 90% are shared with temperate Australasia, 53% with tropical regions, 14% with temperate southern Africa and 13% with the circum-Antarctic islands and South America. New Zealand has undergone such dramatic changes in location, land area, and topography since initial separation from Gondwana 85 Ma that it seems improbable that the 95 species shared with temperate Australasia could have remained conspecific throughout that time. Modern fossil and molecular evidence strongly suggest that many families of ferns had not even evolved prior to separation, and palynological evidence from New Zealand indicates that 78% of pteridophyte genera first appeared there only after separation from Gondwana. Present-day distributions in New Zealand suggest that ferns have greater dispersal potential than flowering plants, and that pteridophyte distributions are more heavily influenced by temperature, rainfall, and geothermal activity than by geological history. Most endemic pteridophyte species have a predominantly southern distribution pattern and are characteristic of cool, lowland to montane forest. Pteridophytes in the northern part of New Zealand show a lower level of endemism than elsewhere and tend to be widespread species that have arrived from temperate Australasian and tropical regions. There is also evidence that at least some pteridophytes have migrated from New Zealand to Australia. It is suggested that the hypothesis of long-distance dispersal of pteridophytes across the Tasman Sea could be tested by molecular techniques.     

Biogeography of the benthic marine algae of the North Atlantic Ocean—an overview

An overview of the biogeography of the benthic marine algae of the North Atlantic Ocean is presented. General and specific distribution patterns are discussed in the light of current knowledge of extant species, and of known events in the evolution of the North Atlantic Ocean. The close relationships between the Arctic, NW and NE Atlantic floras suggest their possible origin as a single flora in the early Oligocene Arctic Ocean, when it was isolated by the Bering Land Bridge and the Greenland-Scotland Ridge. Migration of the flora into the North Atlantic Ocean could have occurred with the subsidence of the Greenland-Scotland Ridge. The present day distribution patterns are the main clue to unravelling the past, and study of vicariant amphi-Atlantic taxa using a variety of experimental techniques will yield the most valuable information in attempts to interpret major biogeographical events in the North Atlantic Ocean.     

A molecular test of Huxley's line: Cyrtandra (Gesneriaceae) in Borneo and the Philippines

A biogeographic and phylogenetic study of Cyrtandra (Gesneriaceae) in the Sundaland region (Borneo and Peninsular Malaysia) and the Philippines using nuclear ribosomal (ITS) DNA sequence data reveals a major division between the Cyrtandra floras of Sundaland and the Philippines. Palawan, the most westerly of the Philippine islands, emerges as an area of mixing between these two. The Bornean element in the Cyrtandra flora of Palawan (two species in our sample) appears to result from recent (i.e. Pleistocene) dispersal from Borneo. The remaining seven species sampled from Palawan are most closely related to those from elsewhere in the Philippines. However, the Palawan clade is sister to the other Philippine taxa, suggesting an ancient (possibly Pliocene) vicariance event. Huxley's line–a zoogeographic boundary placing Palawan and Borneo together–receives some support from this study as there is evidence of recent dispersal of Bornean flora into Palawan. However, in terms of more ancient biogeographic patterning of the region, Palawan has stronger links with the other Philippine islands.     

Strong indication of an extinction‐based saturation of the flora on the Pacific Robinson Crusoe Islands

Oceanic islands are vulnerable ecosystems and their flora has been under pressure since the arrival of the first humans. Human activities and both deliberately and inadvertently introduced biota have had and continue to have a severe impact on island endemic plants. The number of alien plants has increased nearly linearly on many islands, perhaps resulting in extinction‐based saturation of island floras. Here, we provide evidence for such a scenario in Alejandro Selkirk, Robinson Crusoe Islands (Archipelago Juan Fernández, Chile). We compared species richness and species composition of historical vegetation samples from 1917 with recent ones from 2011. Changes in species’ relative occurrence frequency were related to their taxonomic affiliation, dispersal mode, distribution status, and humidity and temperature preferences. While total species richness of vascular plants remained relatively similar, species composition changed significantly. Plants endemic to the Robinson Crusoe Islands declined, exotic species increased substantially within the period of ca. 100 years. Further, the relative occurrence frequency of plants with preferences for very warm and humid climate decreased, while the opposite was found for plants preferring drier and colder environments. Potential drivers responsible for this dramatic shift in the vegetation within only one century might have been the large goat population affecting especially small populations of endemic plants and climatic changes. Taking into account a substantial extinction debt, we expect further shifts in the vegetation of this small oceanic island toward alien plants. This would have significant negative consequences on global biodiversity, considering that island floras contribute substantially to global plant species richness due to their high proportion of endemics.     

On the Limited Potential of Azorean Fleshy Fruits for Oceanic Dispersal

How plants arrived to originally sterile oceanic islands has puzzled naturalists for centuries. Dispersal syndromes (i.e., diaspore traits that promote dispersal by long-distance dispersal vectors), are generally considered to play a determinant role in assisting island colonization. However, the association between diaspore traits and the potential vectors by which diaspores are dispersed is not always obvious. Fleshy fruits, in particular, are considered to have evolved to promote the internal dispersal of seeds by frugivores (endozoochory), however some fleshy fruits can also float in saltwater, and thus be potentially transported by oceanic current (thalassochory). We performed saltwater floatation and viability experiments with fruits of the 14 European fleshy-fruited species that naturally colonized the Azores archipelago (North Atlantic Ocean). We show that only Corema album (a berry) and Juniperus oxycedrus (a fleshy cone) floated for as long as 60 days, the estimated minimum time needed to reach the Azores by oceanic currents. Regardless the floatation potential, exposure to saltwater largely reduced the viability of most seeds of the 14 species (46% of viability decline within 15 days and 77% within 60 days of immersion), including those of Corema album (61%) and Juniperus oxycedrus (83%). Floatability and viability trials suggest that while some fleshy-fruited species might have arrived to the Azores by oceanic currents, such would have required extreme meteorological events that could largely reduce the duration of the trip. Thus, the alternative hypothesis that fleshy-fruited species were mostly dependent on animal dispersers (endozoochory) to colonize these remote islands is reinforced.     

The distribution of seaweed floras in the tropical and subtropical Atlantic Ocean: a quantitative approach

Records of nearly 1500 species of marine algae present in over 80 countries, states, and islands of the tropical and subtropical Atlantic were analyzed to ascertain the relationships between their floras. Association analysis was unsatisfactory, cluster analysis was better, but, of the classificatory methods, indicator species analysis was best. However, the most meaningful results were obtained by ordination using reciprocal averaging. This method demonstrates that after the separation of the marine flora of South-West Africa (Namibia), two clearly defined groupings representing the eastern Atlantic and the western Atlantic are apparent. On the western side, the floras, apart from that of Uruguay, are a relatively close-knit group, except that the northern coast of the Gulf of Mexico together with the southern Atlantic States of the U.S.A. are distinct from the southern Gulf of Mexico and Caribbean groups. On the eastern side of the Atlantic the floras are less closely related. A well-marked tropical group extends from Gambia to Cameroun but does not include the Gulf of Guinea islands which form another group with Angola. This latter group is transitional to the colder water floras lying northwards of Senegal and south of Angola.