The phytogeography ofCladophora (Chlorophyceae) in the northern Atlantic Ocean,in comparison to that of other benthic algal species

About 43Cladophora species inhabit the coasts of the northern Atlantic Ocean. These can be subdivided into seven distribution groups: (a) the tropical western Atlantic group (16 species); (b) the warm temperate Mediterranean-Atlantic group (9 species); (c) the warm temperate North American group (1 species); (d) the Arctic group (1 species); (e) the amphiatlantic tropical to warm temperature group (7 species); (f) the amphiatlantic tropical to temperate group (4 species), and (g) the amphiatlantic temperate group (5 species). These groups agree with general phytogeographic patterns. Thus, the high numbers of species restricted to the tropical western Atlantic region and the warm temperate Mediteranean-Atlantic region are in agreement with the richness and high degree of endemism of these regions. The fact that all species occurring in northeast America also occur in Europe agrees with the high floristic similarity of the boreal areas in America and Europe. The sediment coasts of the Carolinas are an efficient barrier to the south-north dispersal of benthic algae. The temperature bound phytogeographic limits are set in most cases by the species ability to survive adverse temperatures; for “northern” species to survive a high summer temperature in the south, and for “southern” species to survive a low winter temperature in the north. The limits in the Arctic region are all set by the species ability for sufficient growth and reproduction in summer. Conversely, only few northern species have a southern limit which is set by a winter temperature that is not too high for sufficient growth and reproduction. Most species of this group are winter-annuals at their southern limit, and summer-annuals at their northern limit. A comparatively small number of species with a tropical-to-warm temperate distribution have a northern limit at temperate latitudes which is set by a sufficiently high summer temperature for growth and reproduction. A high proportion of this group are lagoonal or quiet water species, which profit by higher summer temperatures in sheltered waters.C. vagabunda is an example.C. rupestris andC. sericea have an amphiboreal distribution and also occur in the southern temperate belt. They probably used a Pleistocene temperature drop to disperse, through the Atlantic along the African coast, from one hemisphere to the other. In the Pacific temperatures were not sufficiently low for this dispersal; and hence these two species reached the Pacific probably by way of the Bering Strait.     

The role of E. maritimum (L.) in the dune pollination network of the Balearic Islands

Eryngium maritimum L. (Apiaceae) is a geophyte that inhabits in the dunes of the Mediterranean and Atlantic. Although it is a highly entomophilous species, there is little literature on its pollinator assemblage. The aim of this study is to analyze the role played by E. maritimum in the dune pollination network of the Balearic Islands, where there is an intense anthropogenic impact in its habitat. For this purpose, two populations located in the North and South of Mallorca were chosen, in which diurnal transects were carried out to observe and capture pollinators on 15 plant species during the anthesis period of E. maritimum. The flowering period of 10 plant species flowering at the same period than E. maritimum was analyzed to identify periods of competition. A total of 71 pollinator species were found, belonging to 30 different families. Eryngium maritimum is a strongly generalist species, with a total of 45 pollinator species. Two new species, Odice blandula and Leucospis gigas, were found for the first time in Mallorca. In terms of pollinators, Teucrium dunense and Helichrysum stoechas are the most similar species to E. maritimum. However, analysis of phenology suggests that these three species have been able to decouple their blooms to avoid competition. The present study shows that E. maritimum plays an important role in the dune pollination network, being its anthesis located at the end of the dune flowering season, when there are no functionally similar species in flower.     

Global seagrass distribution and diversity: A bioregional model

Seagrasses, marine flowering plants, are widely distributed along temperate and tropical coastlines of the world. Seagrasses have key ecological roles in coastal ecosystems and can form extensive meadows supporting high biodiversity. The global species diversity of seagrasses is low (< 60 species), but species can have ranges that extend for thousands of kilometers of coastline. Seagrass bioregions are defined here, based on species assemblages, species distributional ranges, and tropical and temperate influences. Six global bioregions are presented: four temperate and two tropical. The temperate bioregions include the Temperate North Atlantic, the Temperate North Pacific, the Mediterranean, and the Temperate Southern Oceans. The Temperate North Atlantic has low seagrass diversity, the major species being Zostera marina, typically occurring in estuaries and lagoons. The Temperate North Pacific has high seagrass diversity with Zostera spp. in estuaries and lagoons as well as Phyllospadix spp. in the surf zone. The Mediterranean region has clear water with vast meadows of moderate diversity of both temperate and tropical seagrasses, dominated by deep-growing Posidonia oceanica. The Temperate Southern Oceans bioregion includes the temperate southern coastlines of Australia, Africa and South America. Extensive meadows of low-to-high diversity temperate seagrasses are found in this bioregion, dominated by various species of Posidonia and Zostera. The tropical bioregions are the Tropical Atlantic and the Tropical Indo-Pacific, both supporting mega-herbivore grazers, including sea turtles and sirenia. The Tropical Atlantic bioregion has clear water with a high diversity of seagrasses on reefs and shallow banks, dominated by Thalassia testudinum. The vast Tropical Indo-Pacific has the highest seagrass diversity in the world, with as many as 14 species growing together on reef flats although seagrasses also occur in very deep waters. The global distribution of seagrass genera is remarkably consistent north and south of the equator; the northern and southern hemispheres share ten seagrass genera and only have one unique genus each. Some genera are much more speciose than others, with the genus Halophila having the most seagrass species. There are roughly the same number of temperate and tropical seagrass genera as well as species. The most widely distributed seagrass is Ruppia maritima, which occurs in tropical and temperate zones in a wide variety of habitats. Seagrass bioregions at the scale of ocean basins are identified based on species distributions which are supported by genetic patterns of diversity. Seagrass bioregions provide a useful framework for interpreting ecological, physiological and genetic results collected in specific locations or from particular species.     

Within and between population variation in plant traits predicts ecosystem functions associated with a dominant plant species

Linking intraspecific variation in plant traits to ecosystem carbon uptake may allow us to better predict how shift in populations shape ecosystem function. We investigated whether plant populations of a dominant old-field plant species (Solidago altissima) differed in carbon dynamics and if variation in plant traits among genotypes and between populations predicted carbon dynamics. We established a common garden experiment with 35 genotypes from three populations of S. altissima from either Tennessee (southern populations) or Connecticut (northern populations) to ask whether: (1) southern and northern Solidago populations will differ in aboveground productivity, leaf area, flowering time and duration, and whole ecosystem carbon uptake, (2) intraspecific trait variation (growth and reproduction) will be related to intraspecific variation in gross ecosystem CO(2) exchange (GEE) and net ecosystem CO(2) exchange (NEE) within and between northern and southern populations. GEE and NEE were 4.8× and 2× greater in southern relative to northern populations. Moreover, southern populations produced 13× more aboveground biomass and 1.4× more inflorescence mass than did northern populations. Flowering dynamics (first- and last-day flowering and flowering duration) varied significantly among genotypes in both the southern and northern populations, but plant performance and ecosystem function did not. Both productivity and inflorescence mass predicted NEE and GEE between S. altissima southern and northern populations. Taken together, our data demonstrate that variation between S. altissima populations in performance and flowering traits are strong predictors of ecosystem function in a dominant old-field species and suggest that populations of the same species might differ substantially in their response to environmental perturbations.     

Marine longitudinal biodiversity: causes and conservation

The horizontal temperature zones of the earth tend to restrict the latitudinal ranges of species but allow the possibility of exceedingly broad longitudinal dispersals. In the Tropical Zone, biodiversity on the continental shelves is not homogeneous but is concentrated in two conspicuous peaks, one in the Indo‐Pacific Ocean and the other in the Atlantic. The Indo‐Pacific biodiversity peak is located within a relatively small area called the East Indies Triangle. The Atlantic peak is located in the southern Caribbean Sea. Evidence that has been accumulated over the years indicates that each area functions as a centre of origin and evolutionary radiation. What are the causes of these concentrations and their present functions? A newly published model indicates a positive relationship between environmental temperature and the rate of speciation. While this helps to explain the generally high tropical diversity, and the negative relationship between diversity and latitude, it does not provide a reason for the longitudinal concentrations. But, other new research serves to substantiate previous indications of a positive relationship between speciation rate and species diversity. The existence of this positive feedback, together with some contributory factors, provides the reason why concentrations occur. The evolutionary radiation probably begins when the build‐up of species diversity reaches a critical level. The warm‐temperate biotas are derived from the tropics. Their northern longitudinal relationships tend to be minor but, in the southern hemisphere, the West Wind Drift is an important dispersal mechanism for both warm‐temperate and cold‐temperate species. The cold‐temperate biotas peaked in two areas, the North Pacific and the Antarctic; each has developed into a centre of origin. The continuous dispersal of well‐adapted species from the centres helps peripheral communities maintain diversity.     

Flowering,fruiting and seed germination in Chilean rain forest myrtaceae: ecological and phylogenetic constraints

Phenological studies in plant communities have generally focused on taxonomically heterogeneous species assemblages, and have only occasionally examined the evolutionary and ecological constraints on the phenological patterns of species within a single family or a genus. Here, we determine relative importance of phylogenetic versus other constraints on the flowering and fruiting periods of 12 species and the germination ecology of 10 species of Myrtaceae sympatric to the temperate rainforest of Isla Grande de Chiloé (42 °Cs 30 S), in southern Chile.We found that, for most species in the family Myrtaceae, flowering was strongly aggregated in January and February. Although this pattern is consistent with the expectation of the 'facilitation' hypothesis (i.e., interspecific overlaps are maximized to attract pollinators), available evidence suggests that pollinators, mainly hymenopterans and dipterans, do not limit fruit production in these species of Myrtaceae in the temperate rainforest. In contrast to flowering, fruiting occurred all-year-round, showing greater segregation in time among the species. According to the their temporal patterns of fruit ripening, two functional groups were defined within the Myrtaceae: those that ripen their fruits immediately after flowering (species in the subtribe Myrtinae) and those in which green fruit develops slowly for several months before ripening (subtribe Myrciinae). Seed germination in the field occurred mainly between August and October. Lab assays showed that the species of Myrtaceae, subtribe Myrtinae, exhibited a long seed dormancy (>40 days), while the seeds of species in the subtribe Myrciinae often germinated within one week after harvesting. The analysis of the phenology of reproductive events in the species of Myrtaceae in this rainforest suggests that: (1) flowering periods patterns are constrained mainly by phylogenetic inertia at the family level, and (2) differences in fruiting patterns and dormancy periods are determined mainly by fruit and seed size, which in turn are associated primarily with phylogenetic closeness within the family, and secondarily with the activity of vertebrate seed dispersers.     

Reproductive interactions mediated by flowering overlap in a temperate hummingbird–plant assemblage

Pollinator‐mediated competition through shared pollinators can lead to segregated flowering phenologies, but empirical evidence for the process responsible for this flowering pattern is sparse. During two flowering seasons, we examined whether increasing overlap in flowering phenology decreased conspecific pollination, increased heterospecific pollination, and depressed seed output in the seven species composing a hummingbird–plant assemblage from the temperate forest of southern South America. Overall trends were summarized using meta‐analysis. Despite prevailing negative associations, relations between phenological overlap and conspecific pollen receipt varied extensively among species and between years. Heterospecific pollen receipt was low and presumably of limited biological significance. However, our results supported the hypothesis that concurrent flowering promotes interspecific pollen transfer, after accounting for changes in the abundance of conspecific flowers. Seed output was consistently reduced during maximum phenological overlap during the first flowering season because of limited fruit set. Responses varied more during the second year, despite an overall negative trend among species. Relations between estimated effects of phenological overlap on pollination and seed output, however, provided mixed evidence that conspecific pollen loss during pollinator visits to foreign flowers increases pollen limitation. By flowering together, different plant species might benefit each other's pollination by increasing hummingbird recruitment at the landscape level. Nevertheless, our results are mostly consistent with the hypothesis of pollinator‐mediated competition shaping the segregated flowering pattern reported previously for this temperate plant assemblage. The mechanisms likely involve effects on male function, whereby pollen‐transport loss during heterospecific flower visits limit pollen export, and more variable effects on female function through pollen limitation.     

The distribution of benthic marine algae in relation to the temperature regulation of their life histories

Mainly on the basis of the distribution patterns of 42 species of the recently revised genus Cladopkora (Chlorophyceae) in the north Atlantic Ocean, it appeared possible to distinguish 10 phytogeographic distribution groups of wide applicability. Experimentally determined critical temperatures limiting essential events in the life histories of 17 benthic algal species were used to infer possible phytogeographic boundaries; these appeared to fit closely the phytogeographic boundaries derived from field-distribution data. For a temperate species, at least six different boundaries can be postulated and should be checked in the northern hemisphere: (1) the ‘northern lethal boundary’ (corresponding to the lowest winter temperature which a species can survive); (2) the ‘northern growth boundary’ (corresponding to the lowest summer temperature which, over a period of several months, permits sufficient growth); (3) the ‘northern reproductive boundary’ (corresponding to the lowest summer temperature permitting reproduction over a period of several months); (4–6) the corresponding southern boundaries. Photoperiodic responses may influence the temperature responses. Many phytogeographic boundaries appear to be of a composite nature. For instance, the southern boundary of Laminaria digitata follows the European 10°C February isotherm (which corresponds to the highest winter temperature permitting fertility in the female gametophyte, i.e. to the ‘southern reproductive boundary’), and the American 19°C summer isotherm (corresponding to the ‘southern lethal boundary’). Thus, experimental evidence supports the validity of eight of the following 10 distribution groups (for distribution groups 2 and 6, such evidence could not be found): (1) the amphiatlantic tropical-to-warm temperate group, with a north-eastern extension (examples: Gracilaria foliifera and Centroceras clavulalum); (2) the amphiatlantic tropical-to-warm temperate group, with a north-western extension (example: Hypnea musciformis); (3) the amphiatlantic tropical-to-temperate group (example: Sphacelaria rigidula =furcigera); (4) the amphiatlantic temperate group: the Cladophora rupestris type (examples: Callithamnion hookeri, Dumontia contorta; Laminaria saccharina is transitional to type 10, I., digitata to types 5 and 10); (5) the amphiatlantic temperate group: the Cl. albida type (examples: Scytosiphon lomentaria, Petalonia fascia); (6) the tropical western Atlantic group; (7) the north-east American tropical-to-temperate group (example: Gracilaria tikvahiae); (8) the north-east American temperate group and the corresponding Japanese temperate group (examples: Campylaephora hypneoides and Sargassum muticum); (9) the warm-temperate Mediterranean-Atlantic group, and the corresponding warm-temperate Californian group (examples: Saccorhiza polyschides, Laminaria hyperborea, I., ockroleuca, Macrocystis pyrifera, Hedophyllum sessile); (10) the Arctic group (examples: Saccorhiza dermatodea and Sphacelaria arctica). Distribution groups 6, 9 and 10 have comparatively narrow temperature ranges with a span of 18 22°C between their lethal boundaries and of 5 12°C between their reproductive or growth boundaries. These narrow temperature ranges limit the species in these groups to the tropics; the temperate coasts on the eastern sides of the north Pacific and north Atlantic Oceans and in the southern hemisphere; and to the Arctic, respectively. The narrow temperature ranges of group 9 make the species in this group unfit for life on the western temperate coasts of the north Pacific and north Atlantic Oceans, where algae must cope with annual temperature fluctuations of more than 20°C. Conversely, algae in group 8 (containing the numerous Japanese endemic species) are characterized by wide temperature spans (e.g. 29°C between ‘lethal boundaries’, 12–19°C between ‘growth and/or reproductive boundaries’) and must be potentially capable of occupying wide latitudinal belts on temperate coasts along the east sides of the north Pacific and north Atlantic Oceans. Algae ‘escaped’ from Japan, such as Sargassum muticum, conform to this picture. Apparently Japanese algae do not have the capacity for long distance dispersal. The corresponding east American coasts (30–45 N) harbour very few endemic species, probably as a result of the adverse nature of these sediment coasts for benthic macroalgae and their functioning as a barrier to latitudinal displacements of the flora during glaciations. The remaining distribution groups (1,2,3,4,5,7) are characterized by wide temperature spans and wide distributions, often in both the Atlantic and Pacific Oceans and in both hemispheres. Six temperate species (in distribution groups 4, 5 and 9) with an amphiaequatorial distribution have similar winter-temperature maxima permitting reproduction and corresponding with winter isotherms of 15–17°C; their upper lethal temperatures are more dissimilar and correspond with summer isotherms of 20–30°C. Their amphiaequatorial distribution can be explained by assuming glacial temperature drops along east Pacific and east Atlantic equatorial coasts in narrow belts of intensified upwelling during the presumably intensified glacial circulation of the ocean gyres.     

Coastal dune vegetation and pollen representation in south Buenos Aires Province, Argentina

Aim To establish the relationship between coastal dune vegetation and its pollen representation as an aid to interpret Holocene vegetation dynamics and environmental changes from pollen assemblages. Location The study area is situated on the temperate Atlantic coast of south Buenos Aires Province, Argentina (c. 39° S and 61°20′ W). Methods The vegetation of the active dune area adjacent to the beach was described on the basis of its floristic composition from 25 plots. Classification of the vegetation into distinct zones was carried out by cluster analysis. Surface samples were collected from each vegetation stand and analysed for their pollen composition. Pollen percentage data were analysed using principal components analysis in order to investigate the degree to which the different vegetation units can be distinguished by their pollen spectra. Pollen–vegetation relationships for selected taxa were explored using simple scatter plots and indices of association, under‐ and over‐representation. Indices of floristic diversity and palynological richness were used to assess the representation of the vegetation in the pollen spectra. Results and conclusions Five vegetation zones are defined on the basis of species composition and their quantitative variation: back shore, mobile dunes, slacks, semi‐fixed and fixed dunes. Pollen assemblages from back shore, mobile dunes and slacks are clearly differentiated from semi‐fixed and fixed dunes. Pollen assemblages differ considerably from the associated vegetation composition. Major discrepancies are caused by large differences in pollen and vegetation proportion of Hyalis argentea and Discaria americana. There is a considerable proportion of non‐local pollen in every spectrum. Pollen representation in the coastal dunes at Monte Hermoso is influenced by differences in pollen production, dispersal and preservation of individual taxa as well as by the spatial distribution of the vegetation, the topography of the dune system and the wind pattern. The pollen–vegetation relationship established in this study has important implication for understanding and interpreting fossil pollen records from coastal dune environments.     

Phylogeographic analyses suggest that a deciduous species (Ostryopsis davidiana Decne., Betulaceae) survived in northern China during the Last Glacial Maximum

Aim Palaeontologial data suggest that all temperate forest species in northern China migrated southwards during the Last Glacial Maximum (LGM) and recolonized post‐glacially within the Holocene. We tested this assumption using phylogeographical studies of a temperate deciduous shrub species (Ostryopsis davidiana Decne., Betulaceae), which has a wide distribution in northern China. Location Northern China. Methods We sequenced two chloroplast DNA (cpDNA) fragments (trnL–trnF and psbA–trnH, together about 1300 bp in length) of 294 plants from 21 populations across the total distribution range of this species. We used maximum parsimony and haplotype network methods to construct phylogenetic relationships among haplotypes. Results The analysis of cpDNA variation identified eight haplotypes. A single haplotype was fixed in all populations except for one population that was polymorphic, having two haplotypes. The population subdivisions were extremely high (G_ST = 0.972 and N_ST = 0.974), suggesting very low gene flow between populations. Haplotypes clustered into two tentative clades, both of which occur in the southern region of the species’ range but only one of which occurs in the northern region. Across the sampled populations, the haplotype distributions were differentiated geographically. Main conclusions Our analyses suggest that multiple refugia were maintained across the range of O. davidiana in both northern (north of the Qing Mountains) and southern (south of the Qing Mountains) regions during the LGM rather than that the species survived only in the south and subsequently colonized northwards. The extremely low within‐population diversity of this species suggests strong bottleneck or founder effects within each fragmented region during the Quaternary climatic oscillations. These findings provide important clues for understanding range shifts and changes in within‐ and/or between‐population genetic diversity of temperate forests in response to past climatic oscillations in northern China.     

Genetic connectivity between north and south Mid-Atlantic Ridge chemosynthetic bivalves and their symbionts

Transform faults are geological structures that interrupt the continuity of mid-ocean ridges and can act as dispersal barriers for hydrothermal vent organisms. In the equatorial Atlantic Ocean, it has been hypothesized that long transform faults impede gene flow between the northern and the southern Mid-Atlantic Ridge (MAR) and disconnect a northern from a southern biogeographic province. To test if there is a barrier effect in the equatorial Atlantic, we examined phylogenetic relationships of chemosynthetic bivalves and their bacterial symbionts from the recently discovered southern MAR hydrothermal vents at 5°S and 9°S. We examined Bathymodiolus spp. mussels and Abyssogena southwardae clams using the mitochondrial cytochrome c oxidase subunit I (COI) gene as a phylogenetic marker for the hosts and the bacterial 16S rRNA gene as a marker for the symbionts. Bathymodiolus spp. from the two southern sites were genetically divergent from the northern MAR species B. azoricus and B. puteoserpentis but all four host lineages form a monophyletic group indicating that they radiated after divergence from their northern Atlantic sister group, the B. boomerang species complex. This suggests dispersal of Bathymodiolus species from north to south across the equatorial belt. 16S rRNA genealogies of chemoautotrophic and methanotrophic symbionts of Bathymodiolus spp. were inconsistent and did not match the host COI genealogy indicating disconnected biogeography patterns. The vesicomyid clam Abyssogena southwardae from 5°S shared an identical COI haplotype with A. southwardae from the Logatchev vent field on the northern MAR and their symbionts shared identical 16S phylotypes, suggesting gene flow across the Equator. Our results indicate genetic connectivity between the northern and southern MAR and suggest that a strict dispersal barrier does not exist.     

The potential key seed-dispersing role of the arboreal marsupial Dromiciops gliroides

Marsupial seed dispersal is a rare phenomenon, although it may be ecologically significant in southern South America. The marsupial Dromiciops gliroides is endemic to the northern part of the temperate forest of South America. Here we describe the food habits and examine the potential role of D. gliroides as a seed disperser. We evaluated the diet of this marsupial in its natural habitat and in captivity. Dromiciops gliroides is omnivorous showing high consumption of a diversity of fruits. In captivity, D. gliroides consumed fruits from 80% of 22 native plant species we examined. Experiments conducted with fruits from two common understory shrubs show that seed passage through the digestive tract of D. gliroides enhances germination. Our results suggest that this species may have an important role as a seed disperser in the temperate forest of South America, which might offset a scarcity of frugivorous bird species.     

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.     

Tooth row counts, vicariance, and the distribution of the sand tiger shark Carcharias taurus

Geographic variation in tooth row counts among sand tiger sharks Carcharias taurus (Chondrichthyes), from the SW Atlantic, NW Atlantic and the East China Sea is analyzed in this paper. We found significant differences between sand tigers from the SW Atlantic (Southern Hemisphere population) and each of the other two (Northern Hemisphere) regions in the number of upper lateral tooth rows, and between individuals from the SW Atlantic and the East China Sea in the total number of upper tooth rows. Sand tiger sharks from the two Northern Hemisphere populations did not differ in any of the studied variables. Our results agree with comparisons of vertebral counts between sand tiger sharks from Southern and Northern Hemispheres. Both lines of evidence suggest that Southern and Northern Hemisphere populations of C. taurus were isolated to a larger extent than populations of the Northern Hemisphere. The fossil record of the genus Carcharias begins in the Early Cretaceous and C. taurus is certainly known since the Late Miocene. During the Miocene, the Tethys Sea separating northern and southern land masses was still present and it provided a continuous temperate shallow sea that could allow dispersal of sand tiger sharks along Northern Hemisphere seas. Independent observations on the distribution and evolutionary history of the genera Myripristis , Neoniphon , Sargocentron and Aphanius , and genetic studies on the temperate shark genus Mustelus that indicate a close relationship between the Indo-Pacific M. manazo and the Mediterranean M. asterias suggest that this hypothesis is plausible and deserves to be tested.     

Relative importance of temperature and other factors in determining geographic boundaries of seaweeds: Experimental and phenological evidence

Experimentally determined ranges of thermal tolerance and requirements for completion of the life history of some 60 seaweed species from the North Atlantic Ocean were compared with annual temperature regimes at their geographic boundaries. In all but a few species, thermal responses accounted for the location of boundaries. Distribution was restricted by: (a) lethal effects of high or low temperatures preventing survival of the hardiest life history stage (often microthalli), (b) temperature requirements for completion of the life history operating on any one process (i.e. [sexual] reproduction, formation of macrothalli or blades), (c) temperature requirements for the increase of population size (through growth or the formation of asexual propagules). Optimum growth/reproduction temperatures or lethal limits of the non-hardiest stage (often macrothalli) were irrelevant in explaining distribution. In some species, ecotypic differentiation in thermal responses over the distribution range influenced the location of geographic boundaries, but in many other species no such ecotypic differences were evident. Specific daylength requirements affected the location of boundaries only when interacting with temperature. The following types of thermal responses could be recognised, resulting in characteristic distribution patterns: (A) Species endemic to the (warm) temperate eastern Atlantic had narrow survival ranges (between ca 5 and ca 25°C) preventing occurrence in NE America. In species with isomorphic life histories without very specific temperature requirements for reproduction, northern and southern boundaries in Eur/Africa are set by lethal limits. Species with heteromorphic life histories often required high and/or low temperatures to induce reproduction in one or both life history phases which further restricted distribution. (B) Species endemic to the tropical western Atlantic also had narrow survival ranges (between ca 10 and ca 35°C). Northern boundaries are set by low, lethal winter temperatures. Thermal properties would potentially allow occurrence in the (sub) tropical eastern Atlantic, but the ocean must have formed a barrier to dispersal. No experimental evidence is so far available for tropical species with an amphi-Atlantic distribution. (C) Tropical to temperate species endemic to the western Atlantic had broad survival ranges (<0 to ca 35°C). Northern boundaries are set by low summer temperatures preventing (growth and) reproduction. Thermal properties would permit occurrence in the (sub)tropical eastern Atlantic, but along potential “stepping stones” for dispersal in the northern Atlantic (Greenland, Iceland, NW Europe) summer temperatures would be too low for growth. (D) In most amphi-Atlantic (tropical-) temperate species, northern boundaries are set by low summer temperatures preventing reproduction or the increase of population size. On European shores, species generally extended into regions with slightly lower summer temperatures than in America, probably because milder winters allow survival of a larger part of the population. (E) Amphi-Atlantic (Arctic-) temperate species survived at subzero temperatures. In species with isomorphic life histories not specifically requiring low temperatures for reproduction, southern boundaries are set by lethally high summer temperatures on both sides of the Atlantic. None of the species survived temperatures over 30°C which prevents tropical occurrence. Species with these thermal responses are characterized by distribution patterns in which southern boundaries in Eur/Africa lie further south than those in eastern N America because of cooler summers. In most species with heteromorphic life histories (or crustose and erect growth forms), low temperatures were required for formation of the macrothalli (either directly or through the induction of sexual reproduction). These species have composite southern boundaries in the north Atlantic Ocean. On American coasts, boundaries are set by lethally high summer temperatures, on European coasts by winter temperatures too high for the induction of macrothalli. Species with this type of thermal responses are characterized by distribution patterns in which the boundaries in Eur/Africa lie further north than those in eastern N America because of warmer winters. Paper presented at the XIV International Botanical Congress (Berlin, 24 July–1 August, 1987), Symposium 6-15, “Biogeography of marine benthic algae”.     

Flowering and fruiting phenology in the coastal shrublands of Doñana,south Spain

Flowering and fruiting phenological patterns at the individual-, population-, and community level were studied in a southern Spanish scrub community composed of 30 shrub species. Few individuals of any species produced a high number of flowers. Intrapopulation deviation in the peak time of flowering showed a strong and positive skewness. Relative flowering duration, however, displayed a virtually normal distribution. Generally, species flowering in spring have a short flowering time, while species flowering earlier or later in the year show significantly longer flowering periods. Species were in bloom throughout the year, but there was a major peak during spring and two lesser ones in autumn and early summer. Shallow rooting taxa in typically mediterranean genera displayed a strategy of spring flowering and summer fruiting. Summer and autumn flowering occurred among heath-like shrubs of relatively wet places, and forest-associated, vertebrate-dispersed species which commonly have underground storage organs. Species with ripe fruits presented two peaks, the major one during the summer including the majority of taxa with seeds dispersed by non-vertebrate agents. There was a minor fruiting peak in autumn dominated by taxa that rely on vertebrates for dispersal. The complex seasonal patterns observed are interpreted in relation to environmental conditions and physiological constraints on species living in a highly seasonal climate.     

Spatial distributions of copepod genera along the Atlantic Meridional Transect

The distribution of copepod taxa at a basin scale was analysed using three Atlantic transects (U.K. – Malvinas 1997, Malvinas – U.K. 1997, and South Africa – U.K. 1998). Integrated 200 m to surface zooplankton samples were taken daily, using WP2 nets (200-m mesh). The zooplankton were size-fractionated and sub-samples taken for carbon analysis. The remainder of the samples was preserved for taxonomic analysis of copepod genera. Multidimensional scaling (MDS) was used to identify zoogeographic regions from the copepod genera. Seven regions were identified: northern temperate, northern subtropical, equatorial, southern tropical, southern sub-tropical, southern temperate and Benguela upwelling. Analysis of similarity showed that most regions were significantly different from each other except: northern temperate and southern temperate, northern temperate and southern subtropical, and northern subtropical and southern subtropical. The genera significant in determining the regions were identified. These regions were compared to other schemes of biological and hydrographic areas. The MDS also showed that the copepod composition in the tropical and subtropical regions was less variable than the temperate and Benguela stations. Latitudinal trends in diversity and size were also investigated. Copepod genera showed a reduction in richness at higher latitudes. Copepod size did not show any substantial or consistent change with latitude along these transects, as demonstrated by both the numerical abundances in each size category, and the carbon biomass per individual. The proportion in each size fraction was quite uniform over the transect.     

From Amazonia to the Atlantic forest: Molecular phylogeny of Phyzelaphryninae frogs reveals unexpected diversity and a striking biogeographic pattern emphasizing conservation challenges

Documenting the Neotropical amphibian diversity has become a major challenge facing the threat of global climate change and the pace of environmental alteration. Recent molecular phylogenetic studies have revealed that the actual number of species in South American tropical forests is largely underestimated, but also that many lineages are millions of years old. The genera Phyzelaphryne (1 sp.) and Adelophryne (6 spp.), which compose the subfamily Phyzelaphryninae, include poorly documented, secretive, and minute frogs with an unusual distribution pattern that encompasses the biotic disjunction between Amazonia and the Atlantic forest. We generated >5.8kb sequence data from six markers for all seven nominal species of the subfamily as well as for newly discovered populations in order to (1) test the monophyly of Phyzelaphryninae, Adelophryne and Phyzelaphryne, (2) estimate species diversity within the subfamily, and (3) investigate their historical biogeography and diversification. Phylogenetic reconstruction confirmed the monophyly of each group and revealed deep subdivisions within Adelophryne and Phyzelaphryne, with three major clades in Adelophryne located in northern Amazonia, northern Atlantic forest and southern Atlantic forest. Our results suggest that the actual number of species in Phyzelaphryninae is, at least, twice the currently recognized species diversity, with almost every geographically isolated population representing an anciently divergent candidate species. Such results highlight the challenges for conservation, especially in the northern Atlantic forest where it is still degraded at a fast pace. Molecular dating revealed that Phyzelaphryninae originated in Amazonia and dispersed during early Miocene to the Atlantic forest. The two Atlantic forest clades of Adelophryne started to diversify some 7Ma minimum, while the northern Amazonian Adelophryne diversified much earlier, some 13Ma minimum. This striking biogeographic pattern coincides with major events that have shaped the face of the South American continent, as we know it today.     

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.