Patterns of endemism in south-eastern Pacific benthic polychaetes of the Chilean coast

Aim In this study we evaluate patterns of endemism for benthic polychaete species along the southeastern Pacific coast of Chile. Our goals were (1) to describe latitudinal gradients of endemism and identify areas of high endemism, (2) to evaluate the effect of biogeographical limits on endemism patterns, and (3) to evaluate indirectly the role played by evolutionary dynamics on patterns of endemism. Location South‐eastern Pacific coast of Chile, ranging from Arica (18° S) to Cape Horn (56° S). Methods We used a list of 178 species of endemic, shallow benthic polychaetes to evaluate patterns of endemism. Parsimony analysis of endemicity (PAE) and the endemism index (EI) were used to evaluate hierarchical relationships of endemism between different latitudinal bands, and to identify areas with high degrees of endemism and differences in endemism. We evaluated the effect of biogeographical limits on endemic polychaete fauna by testing for the existence of geometric constraints (mid‐domain effect). The role of evolutionary dynamics on latitudinal patterns of endemism was evaluated with nestedness analysis (NA) using the temperature index. Results The PAE analysis indicated two large, separate areas of endemism: (1) the northern area between 18° S and 38° S, and (2) the southern area between 39° S and 56° S. The endemism index showed a maximum value (32 species) around 39°–41° S. Species‐richness curves of each 3° band of latitude showed a clear mid‐domain effect (69%), but the two maximum points of species richness at mid‐latitudes (36° S to 38° S and 39° S to 41° S) did not correspond to the mid‐domain peak in species richness, presenting a greater number of species than expected by the mid‐domain effect. The nestedness analysis showed that the number of genera reaches a maximum of 70 at mid‐latitudes (36°–41° S), decreasing towards both the northern and southern areas. The spatial distribution of the entire data set of endemic species showed a nested pattern (T° = 24.5°, P < 0.0001). Main conclusions Our results strongly support the existence of a latitudinal gradient of endemism for benthic polychaete species along the Chilean coast. The shape of this gradient is clearly non‐linear, with a marked peak of endemism occurring at mid‐latitudes (36°–41° S, endemism hotspot), which also corresponds to a peak in species richness. Furthermore, this hotspot is the midpoint separating two distinct areas of endemism to the north and south. We suggest that the observed pattern of endemism for benthic polychaete taxa of the Chilean coast can be explained by a combination of geometric constraints and historical mechanisms, such as the processes that affected the Chilean coast during the Neogene (e.g. ENSO, oxygen minimum zone, glaciations).     

Stomach contents of long-finned pilot whales (Globicephala melas) from southern Chile

The widely distributed long-finned pilot whale (Globicephala melas) has been reported off the Chilean coast, from Iquique (20°12′S) south to Navarino Island (55°15′S; 67°30′W), but little is known about its biology or ecology in the region. Here, we report on the prey of this species, identified by stomach content analyses from animals stranded on Holget Islets, Beagle Channel, southern Chile in August 2006. The stomachs of seven individuals (six females and one male) contained cephalopod remains. The prey composition found in these southern Chilean pilot whales was similar to that described in other parts of the world and the Southern Ocean. This is the first report on the feeding habits of this species from Chile.     

Complex patterns of expression of heat-shock protein 70 across the southern biogeographical ranges of the intertidal mussel Mytilus californianus and snail Nucella ostrina

Aim It is often assumed that species reach their highest densities in the centre of their ranges and decline in abundance toward the edges of the range. Implicit in this notion, which we call the abundant centre hypothesis, is the assumption that the edges of the range are more stressful to organisms and are more likely to show responses to climate change. However, an earlier review and empirical study of patterns of abundance across the range of intertidal invertebrates show little support for the abundant centre hypothesis and further demonstrated that few studies have examined patterns in either abundance or stress across species ranges. In part this gap is due to the logistical difficulties of sampling species across large geographical ranges. Here we use intertidal invertebrates, which have relatively simple linear latitudinal ranges, and heat‐shock proteins, which have been shown to be an integrative measure of organismal stress, to test the hypothesis that species are more stressed at the edges of their range. We use complementary data on population density to test the relationship between stress proteins and overall species density across the species’ range. Location Our sampling programme covered the southern half of the large geographical ranges of two intertidal invertebrates on the Pacific Coast of North America. Sites were spread between northern Baja California, Mexico and Vancouver Island, Canada, a range of c. 22 degrees of latitude. Method We sampled levels of heat‐shock protein 70 (Hsp70) in eight to 12 individuals from each of 20 sites for the intertidal mussel Mytilus californianus and 11 sites for the intertidal snail Nucella ostrina, spread throughout the southern half of their geographical ranges. The relationships between levels of Hsp70 in individuals from a site and (1) latitude of the site, (2) the site's position in the species’ range and (3) average population density were determined. Results No significant relationship was found in either species between levels of Hsp70 and latitude, position in the range or population density. Complex patterns that did emerge may be explained by nonlinear gradients in environmental conditions along the Pacific coast. Specifically, we observed peak values of Hsp70 for both species in northern Oregon, where intertidal zones are disproportionately exposed to daytime emersion (exposure to air) in the summer months of collections. A second peak for M. californianus was found south of Point Conception, California, which is marked by dramatic shifts toward warmer sea temperatures and decreased wave exposure. Main conclusions Patterns that emerged were not predicted by simple models based on the abundant centre hypothesis. However, they are consistent with more complex pictures of heat stress, organismal condition and abundance along a latitudinal gradient that have been demonstrated in recent studies. We suggest that latitudinal complexity, species‐specific differences and local effects must be considered before generalizing the relationship between environmental stress, abundance, range limits and responses of ranges to climate change.     

The zoogeography of algae-associated peracarids along the Pacific coast of Chile

Abstract Aim To describe the zoogeography of the algae‐associated peracarid crustaceans from exposed rocky shores along the SE‐Pacific. Location Chile, 18° S to 42° S. Methods A standardized sampling programme was used at all sites. Samples of macroalgae were taken at twenty sites distributed along the entire study area. Quantitative samples (n = 6 replicates of 8 cm^?2 surface area each) of calcareous and non‐calcareous red algae were taken in the low intertidal, preserved immediately in 4%‐formalin and washed over a 0.2‐mm mesh before sorting. All peracarid individuals were sorted, identified to the species level and then categorized in separate functional groups according to their feeding habits. Graphical representations of species replacement within each functional group along the latitudinal gradient are provided. A classification analysis employing the unweighted paired group method using arithmetic average (UPGMA) was conducted in order to reveal the main zoogeographical zones. Results A total of forty epifaunal peracarid species was found. A gradual replacement of species within different functional groups (grazing and suspension‐feeding species) was observed in the central region (c. 26° S?37° S). In this central region, species with northern and those with southern distribution overlapped, while other species were only found here, resulting in high species richness. The number of species/site/algal species in the northern (north of c. 25.5° S) and southern region (south of c. 38.5° S) was considerably lower than in the central region. The distribution of most grazing peracarids showed a more continuous pattern than that of suspension‐feeding amphipods. The distribution of the remaining species (predators, scavengers, deposit‐feeders, unknowns) was scattered along the examined sites. The cluster analysis for the epifaunal peracarid assemblage confirmed the separation of a northern and southern zone connected by a central (transitional) zone between c. 26° S and c. 37° S. Similar zonation patterns have been found by most other studies on the zoogeography of the Chilean coast, although little agreement exists about the exact limits of this transitional zone. It is discussed that the distribution limits of algae‐associated peracarids (and other macroinvertebrates) – particularly in the transitional zone – may show interannual variations as a result of varying oceanographic conditions. The large affinity of the algae‐associated peracarid fauna from the central and southern Chilean coast to those of other regions indicates that dispersal may be facilitated by rafting with floating algae transported in the Antarctic Circumpolar Current. Main conclusions The zoogeographical analysis of algae‐associated peracarids confirms the existence of a northern and a southern zone connected by an extensive transitional zone. General biology, habitat use and the abundant presence of dispersal vectors such as floating macroalgae may affect the zoogeography of species living in transitional zones with strong interannual variations in current regimes. In these areas, species associated with substrata of high dispersal potential may show different distribution patterns than species inhabiting other substrata.     

Faunal changes and geographic crypticism indicate the occurrence of a biogeographic transition zone along the southern East Pacific Rise

Aim Deep‐sea hydrothermal vents have now been reported along all active mid‐ocean ridges and back‐arc basins, but the boundaries of biogeographic entities remain questionable owing to methodological issues. Here we examine biogeographic patterns of the vent fauna along the East Pacific Rise (EPR) and determine the relative roles of regional and local factors on the distribution of biodiversity associated with mussel beds along a poorly explored zone, the southern EPR (SEPR). Location East Pacific Rise. Methods A species list of macrobenthic invertebrates along the EPR was compiled from the literature and supplemented with data recovered during the French research cruise BIOSPEEDO carried out in 2004 along the SEPR. Biogeographic patterns were assessed by combining the identification of morphological species with a molecular barcoding approach. A multivariate regression tree (MRT) analysis was performed to identify any geographic breaks, and an empirical distribution of species richness was compared with predictions provided by a mid‐domain effect model. Macrofaunal community structure associated with mussel beds along the SEPR was analysed in relation to environmental factors using cluster and canonical redundancy analyses. Results Sequencing of the cytochrome c oxidase subunit I gene revealed the occurrence of several cryptic species complexes along the EPR, with the equator separating the southern and northern clades. Furthermore, during the BIOSPEEDO cruise at least 10 still unnamed species were collected between 7°25′ S and 21°33′ S. The shift in community structure identified by MRT analysis was located south of 17°34′ S or south of 13°59′ S, depending on the data used, suggesting that the southern part of the SEPR (17°25′–21°33′ S) constitutes a biogeographic transition zone in the vent fauna along the EPR. At a regional scale, latitude combined with the type of venting was significantly correlated with the community structure associated with mussel beds. Main conclusions Together, the molecular data, in situ observations, and the distribution of species suggest that the high diversity of vent fauna species presently observed between 17°25′ S and 21°33′ S is probably a result of the overlap of several distinct biogeographic provinces. We argue that this area thus constitutes a biogeographic vent fauna transition zone along the EPR.     

Range limits and geographic patterns of abundance of the rocky intertidal owl limpet,Lottia gigantea

Aim We evaluate the stability of the range limits of the rocky intertidal limpet, Lottia gigantea, over the last c. 140 years, test the validity of the abundant centre hypothesis, and test indirectly the roles played by recruitment limitation and habitat availability in controlling the range limits. Because this species is size‐selectively harvested, our results also allow us to assess conservation implications. Location The Pacific coast of North America, from northern California to southern Baja California (41.74° N–23.37° N), encompassing the entire range of L. gigantea. Methods The historical and modern distributions of L. gigantea were established using museum data and field observations. Overall and juvenile abundances of local populations were estimated at 25 field sites. The spatial distribution of abundance was evaluated statistically against the predictions of five hypothetical models. The availability of habitat was estimated by measuring the percentage of unavailable sandy beach within cumulative bins of coast across the range of L. gigantea. Results The northern limit of L. gigantea has contracted by c. 2.4° of latitude over recent decades (after 1963), while the southern limit has remained stable. The highest abundances of L. gigantea occur in the centre of its geographic range. Habitat availability is ample in the centre and northern portions of its range, but is generally lacking in the southern range. The northern range is only sparsely populated by adults, with sharp declines occurring north of Monterey Bay (36.80° N). In the southern range, abundance drops precipitously south of Punta Eugenia (27.82° N), coinciding with the region where suitable habitat becomes sparse. Main conclusions Support for the abundant centre hypothesis was found for L. gigantea. Northern populations are characterized as being recruitment‐limited, demographically unstable and prone to local extinctions, while southern populations are suggested to be habitat‐limited. The abundant centre is suggested to result partly from a combination of the indirect effects of human harvesting, generating denser populations of smaller individuals, and larval recruitment from well‐protected offshore rocky islands primarily found in the range centre.     

The Indochinese–Sundaic zoogeographic transition: a description and analysis of terrestrial mammal species distributions

Aim We describe the distributions of mammal species between the Indochinese and Sundaic subregions and examine the traditional view that the two faunas show a transition near the Isthmus of Kra on the Thai–Malay peninsula. Location Species distributions are described along a 2000‐km transect from 20° N (northernmost Thailand) to 1° N (Singapore). Methods For the 325 species of native non‐marine mammals occurring along the transect we used published records to provide a database of their distributional records by degree of latitude. Results Along the transect we found 128 Indochinese species with southern range limits, 121 Sundaic species with northern range limits, four un‐assignable endemics and 72 widespread species. In total, 152 southern and 147 northern range limits were identified, and their distribution provides no evidence for a narrow faunal transition near the Isthmus of Kra (10°30′ N) or elsewhere. Range limits of both bats and non‐volant mammals cluster in northernmost peninsular Malaysia (5° N) and 800 km further north, where the peninsula joins the continent proper (14° N). The clusters of northern and southern range limits are not concordant but overlap by 100–200 km. Similarly, the range limits of bats and non‐volant mammals cluster at slightly different latitudes. There are 30% fewer species and range limits in the central and northern peninsula (between 6 and 13° N), and 35 more widely distributed species have range gaps in this region. In addition, we found 70 fewer species at the southern tip of the peninsula (1° N) than at 3–4° N. Main conclusions The deficiencies of both species and species range limits in the central and northern peninsula are attributed to an area effect caused by repeated sea‐level changes. Using a new global glacioeustatic curve developed by Miller and associates we show that there were > 58 rapid sea‐level rises of > 40 m in the last 5 Myr that would have resulted in significant faunal compression and local population extirpation in the narrow central and northern parts of the peninsula. This new global sea‐level curve appears to account for the observed patterns of the latitudinal diversity of mammal species, the concentration of species range limits north and south of this area, the nature and position of the transition between biogeographical subregions, and possibly the divergence of the faunas themselves during the Neogene. The decline of species diversity at the southern end of the transect is attributed to a peninsula effect similar to that described elsewhere.     

Two oceans, two taxa and one mode of development: latitudinal diversity patterns of South American crabs and test for possible causal processes

The latitudinal gradient of species diversity is a widely recognized but poorly understood phenomenon. In marine systems, differences in dispersal abilities among species may pose an additional problem in identifying the processes that affect diversity. We compared latitudinal diversity gradients along two parallel continental coasts, the east and west coasts of South America, of two groups of Crustacea (Brachyura and Anomura), which exclusively exhibit planktonic development. We also evaluated the species‐area and the energy‐input hypotheses. Diversity decreased with increasing latitude for both groups in both oceans. Results suggest that the spatial structure of sea surface temperature (SST) explains diversity of both groups at large, but not small (< 5°), scales. Range size and latitude were not correlated. We hypothesize that SST differentially affects taxa with contrasting modes of development, influencing patterns of diversity. We suggest that developmental modes of marine organisms should be considered in future diversity analyses.     

Patterns of distribution for southern Australian marine echinoderms and decapods

Aim To relate patterns of distribution of marine echinoderms and decapods around southern Australia to major ecological and historical factors. Location Shallow‐water (0–100 m) marine waters off southern Australia, south of 30° S. Methods (1) Record the presence/absence of known echinoderm and decapod species in cells of c. 1° latitude and longitude, along the coast of southern mainland Australia and Tasmania. (2) Describe patterns in species composition, species richness and endemism through gradient analysis, ordination and cluster analysis. (3) Relate these patterns to distance and temperature gradients, the area of continental shelf, the average size of species range, and known historical factors. Results Species composition varied with both latitude and longitude. Species richness was relatively constant from east to west but graded with latitude from high in the warm‐temperate regions around Perth and Sydney to low in cool‐temperate southern Tasmania. Species richness was not related to the area of continental shelf or average species range size. Species turnover was not correlated with rates of temperature change. It was problematic to separate distance from temperature gradients, but there was evidence that the southern distribution limits of some species are related to minimum sea surface temperature. Within the taxonomic groups surveyed, evolutionary radiation has been largely limited to a few cosmopolitan species‐rich genera. Main conclusions There are historical as well as ecological hypotheses explaining the latitudinal gradient of marine species richness in southern Australia: (1) the continual invasion and speciation of species of tropical origin as Australia has split from Gondwana and drifted northward; (2) progressive extinction of some Gondwanan cool‐temperate species at the limits of their range; (3) low level of immigration of additional cool‐temperate species; and (4) some in situ endemic speciation.     

Marine biogeography of southern Australia: phylogeographical structure in a temperate sea-star

Aim To test whether marine biogeographical patterns observed at the community level are also important within species. It is postulated that historical hydrogeographic barriers have driven in situ diversification. Location The intertidal and shallow subtidal zones of southern Australia, New Zealand and nearby islands. Australia's temperate marine communities are characterized by a high degree of endemism and show strong biogeographical structure along an east–west axis. Methods Phylogeographical analysis of the widespread asteriid sea‐star Coscinasterias muricata Verrill across southern Australia and New Zealand. Forty‐two samples from 27 locations were included in phylogenetic analyses of mitochondrial (CO1; control region) and nuclear (ITS2) DNA sequences. Results Analysis of mtDNA revealed a deep phylogenetic split within Australian C. muricata, strongly correlated with latitude. ‘Northern’ haplotypes (latitude ≤ 37.6° S, nine sites, 15 samples) were 7.3–9.4% divergent from ‘southern’ haplotypes (latitude ≥ 37.6° S, 19 sites, 27 samples), consistent with late Pliocene separation. Eastern and western representatives of the ‘northern’ clade were 0.5–1.0% divergent, probably reflecting Pleistocene isolation. The ‘southern’ clade of Australia is also represented in New Zealand, indicating Pleistocene oceanic dispersal. Nuclear DNA (ITS2) sequences yielded relatively little phylogenetic resolution, but were generally congruent with mtDNA‐based groupings. Main conclusions The phylogeographical pattern detected within Australian C. muricata closely resembles marine biogeographical groupings proposed on the basis of community and species distributions. Recurring evolutionary patterns may have been driven by the hydrographic history of southern Australia. Specifically, we suggest that Plio‐Pleistocene temperature change and the repeated opening and closure of Bass Strait promoted allopatric divergence and perhaps cryptic speciation in C. muricata.     

Biogeographical patterns and Rapoport's rule in southeastern Pacific benthic polychaetes of the Chilean coast

Recently three biogeographical units were identified along the Chilean coast (the Magellanic Province, an Intermediate Area, and the Peruvian Province), however few studies have focused on the factors and dynamic processes that formed these spatial units (e.g. Rapoport's rule and its causal mechanisms). In this study we used benthic polychaetes of the Chilean coast to evaluate patterns of latitudinal distribution and species richness, and the existence of the three main biogeographical provinces described for the Chilean coast. Additionally, we evaluated the latitudinal Rapoport effects and geometric constraint as a null hypothesis explaining the species richness distribution.
We found that benthic polychaete diversity increased towards southern latitudes. The cluster and ordination (non-metric MultiDimensional Scaling, nMDS) analyses of the distribution data, presented only two statistically significant (bootstrapping techniques) biogeographic provinces along the Chilean coast, with a break occurring between 41° and 42°S. While, our results did not support a latitudinal Rapoport effect, they did support the view that latitudinal Rapoport effects are a local phenomenon, occurring only for the Northeastern Pacific marine taxa. The relationship between latitudinal range extent and mean latitude indicated the existence of two hard boundaries at either extreme of the Chilean coast, limiting the geographical ranges of the species. However, geometric constraints tested using a Monte Carlo simulation approach showed a weak level of mid-domain effect on species richness. Finally, we propose that geometric constraint together with the geomorphology and historical characteristics of the Chilean coast explain the biogeographical patterns of benthic polychaete taxa in Chile.     

SEX RATIO VARIATION IN THE LESSONIA NIGRESCENS COMPLEX (LAMINARIALES,PHAEOPHYCEAE): EFFECT OF LATITUDE,TEMPERATURE, AND MARGINALITY1

Little is known about variation of sex ratio, the proportion of males to females, in natural populations of seaweed, though it is a major determinant of the mating system. The observation of sexual chromosomes in kelps suggested that sex is partly genetically determined. However, it is probably not purely genetic since the sex ratio can be modified by environmental factors such as salinity or temperature. In this paper, sex ratio variation was studied in the kelp Lessonia nigrescens Bory complex, recently identified as two cryptic species occurring along the Chilean coast: one located north and the other south of the biogeographic boundary at latitude 29°–30° S. The life cycle of L. nigrescens is characterized by an alternation of microscopic haploid gametophytic individuals and large macroscopic fronds of diploid sporophytes. The sex ratio was recorded in progenies from 241 sporophytic individuals collected from 13 populations distributed along the Chilean coast in order (i) to examine the effect of an environmental gradient coupled with latitude, and (ii) to compare marginal populations to central populations of the two species. In addition, we tested the hypothesis that the sex ratios of the two cryptic species would be affected differently by temperature. First, our results demonstrate that sex ratio seems to be mainly genetically determined and temperature can significantly modify it. Populations of the northern species showed a lower frequency of males at 14°C than at 10°C, whereas populations of the southern species showed the opposite pattern. Second, both species displayed an increased variation in sex ratio at the range limits. This greater variation at the margins could be due either to differential mortality between sexes or to geographic parthenogenesis (asexual reproduction).     

Individualistic species limitations of climate‐induced range expansions generated by meso‐scale dispersal barriers

Aim Evidence indicates that species are responding to climate change through distributional range shifts that track suitable climatic conditions. We aim to elucidate the role of meso‐scale dispersal barriers in climate‐tracking responses. Location South coast of England (the English Channel). Methods Historical distributional data of four intertidal invertebrate species were logistically regressed against sea surface temperature (SST) to determine a climate envelope. This envelope was used to estimate the expected climate‐tracking response since 1990 along the coast, which was compared with observed range expansions. A hydrodynamic modelling approach was used to identify dispersal barriers and explore disparities between expected and observed climate tracking. Results Range shifts detected by field survey over the past 20 years were less than those predicted by the changes that have occurred in SST. Hydrodynamic model simulations indicated that physical barriers produced by complex tidal currents have variably restricted dispersal of pelagic larvae amongst the four species. Main conclusions We provide the first evidence that meso‐scale hydrodynamic barriers have limited climate‐induced range shifts and demonstrate that life history traits affect the ability of species to overcome such barriers. This suggests that current forecasts may be flawed, both by overestimating range shifts and by underestimating climatic tolerances of species. This has implications for our understanding of climate change impacts on global biodiversity.     

The distribution of the ectoparasite fauna of Sebastes capensis from the southern hemisphere does not correspond with zoogeographical provinces of free-living marine animals

Aim We examined the ectoparasite fauna of Sebastes capensis over almost all its geographical distribution range (Chilean, Argentinean and South African coasts) to determine (1) whether the ectoparasites of this host show a zoogeographical pattern and, if so, (2) how this pattern is related to known zoogeographical patterns for free‐living organisms. Location Fish were captured from 20, 24, 30, 33, 36, 40, 45 and 52° S along the Chilean coast; 11° S on the Peruvian coast; 43° S on the Argentina coast; and 34° S on the South African coast. Methods From April to September 2003 and from April to August 2004, 626 fish were captured. The parasites were collected using standard parasitological techniques. At the component community level, zoogeographical distribution patterns were evaluated using cluster analysis. At the infra‐community level, patterns of similarity in parasite composition among localities were investigated with multivariate discriminant analyses. Results The ectoparasite fauna of S. capensis consists of six species distributed along the whole of the Chilean coast. Four other species are distributed only within the transitional zone between the northern warm temperate region (Peruvian faunistic province), extending from Peru to the northern Chilean coast up to c. 30° S, and the cold temperate region (Magellanic faunistic province). The component communities from latitudes 30 to 40° S showed higher ectoparasite species richness, while localities on the margins of the geographical range showed lower species richness. Cluster analysis indicated a grouping of localities consistent with the transitional zone. Argentina and South Africa always emerged as separate localities. Main conclusions The ectoparasite communities of S. capensis do not follow a distributional pattern concordant with the known biogeographical zones for invertebrates and/or fish along the south‐eastern Pacific. Therefore their ectoparasite fauna is not useful as a zoogeographical indicator, although it does allow us to distinguish the transitional zone of the south‐eastern Pacific. On a more extended geographical scale, it is possible to distinguish the ectoparasite communities of S. capensis in the south‐eastern Pacific (as a whole) from those of Argentina and South Africa.     

Terrestrial biodiversity along the Ross Sea coastline,Antarctica: lack of a latitudinal gradient and potential limits of bioclimatic modeling

Antarctica has several apparent advantages for the study of biodiversity change along latitudinal gradients including a relatively pristine environment and simple community structures. Published analyses for lichens and mosses show no apparent gradient in biodiversity along the western Ross Sea coast line, the longest ice-free area in Antarctica spanning 14° latitude. One suggestion is that the area remains poorly surveyed. Here, we combine available species lists from four sites along the coast with new own data from two additional sites [Taylor Valley (77°30′S) and Diamond Hill (79°S)]. We show a decline in total terrestrial biodiversity with latitude from Cape Hallett (72°S) to Diamond Hill. However, the southernmost site, the Queen Maud Mountains (84°S), is exceptional with almost the same diversity as Cape Hallett. A categorization of lichens according to their proposed ecology shows the proportion of tolerant species remains relatively constant. However, the absolute number of conformant species declines with latitude, again with a minimum at Diamond Hill. Similarity indices are low and not very different between sites with Diamond Hill being the exception with very few species. We suggest that terrestrial biodiversity best reflects microhabitat water availability rather than macroclimatic temperature changes and use climate data from Taylor Valley and Diamond Hill to support this suggestion. We propose that the importance of microhabitats and landscape location is one of several possible limitations to the application of bioclimatic modeling along the Ross sea coastline. In the absence of a definitive link between macroclimate and the biota, predicting the effects of climate changes will be more challenging.     

Does low temperature cause the dominance of Acacia on the central Australian mountains? Evidence from a latitudinal gradient from 11° to 26° South in the Northern Territory,Australia

A latitudinal gradient, from the central north coast of the Northern Territory (11°S) to the South Australian State border (26°S), was defined to subsample a large 20 m×20 m quadrat data set (N>2000 quadrats) collected during the course of the Northern Territory 1:10⁶ vegetation mapping programme. The mean and standard error of a range of environmental data, and structural and floristic variables pertaining to woody species were calculated for fifteen cells (3.5°E wide and 1°S long) on the transect using a total of 1050 quadrats. It was found that the interrelated measures of mean canopy height, mean canopy cover and mean total basal area steadily declined from the north coast to reach their minimum levels at 18.5°S. There was little variation in these variables south of this latitude. This pattern is probably controlled by precipitation given that there is a highly significant (r²=98%) negative exponential decay of mean annual rainfall with latitude for ten meteorological stations on or near the transect, and that the southern limit of the summer monsoon rains is at about 18°S. The mean percentage of the woody species quadrat richness attributable to Acacia species was found to increase at around 18.5°S. However, the mean Acacia basal area and the percentage of the total basal area composed of Acacia basal area increased at 21.5°S. At this latitude the mean Eucalyptus basal area, the percentage of the total basal area made up of Eucalyptus species, and the mean percentage of woody species quadrat richness composed of Eucalyptus species all decreased to minimum levels. South of 21.5°S mean landscape elevation ranges between 400 and 700 m above sea level while north of this latitude mean landscape elevation ranges 10–300 m above sea level. The combined effects of continentality and environmental lapse rates result in a highly significant (r²=82%) negative exponential decay of mean July (winter) minimum temperature with latitude for the five climate stations on the transect for which data are available. Mean slope angle, rock cover, surface soil gravel content and surface soil clay content were unrelated to any of the above patterns. It is suggested that the sharp change-over in Eucalyptus to Acacia dominance at 21.5°S is related to minimum temperature, but this hypothesis requires testing with detailed ecophysiological studies. None the less, the local dominance of Acacia shirleyi at 16.5°S suggests that environmental history also may have a strong influence on the contemporary latitudinal distribution of Acacia and Eucalyptus in the northern half of the Australian continent. Regrettably, few data are available to evaluate critically the importance of long-term environmental change on current distributional patterns of Acacia and Eucalyptus.     

Implications of acute temperature and salinity tolerance thresholds for the persistence of intertidal invertebrate populations experiencing climate change

To predict whether populations of marine animals will persist in the face of changing climate conditions, it is informative to understand how past climate conditions have shaped present‐day tolerance thresholds. We examined 4 species of intertidal invertebrates (Nucella lamellosa, Littorina scutulata, Littorina sitkana, and Balanus glandula) inhabiting the coasts of Vancouver Island, Canada, where the east coast experiences historically warmer sea surface temperature (SST), warmer low tide (i.e., emersion) rock surface temperature (RST), and lower sea surface salinity (SSS) than the west coast. To determine if east coast populations have higher tolerance thresholds to acute stress than west coast populations, animals from 3 sites per coast were exposed to stressful temperatures and salinities in common garden experiments. Emersion temperature tolerance differed between populations only in N. lamellosa and B. glandula, tolerance thresholds being 1.4–1.5°C higher on the east coast. Water temperature tolerance differed between populations only in B. glandula and L. scutulata but was highest on the west coast. No differences in salinity tolerance were observed within any species. Thus, there is limited evidence of divergence among east and west coast populations in tolerance of acute stress despite the substantial historical differences in extreme temperature and salinity conditions between coasts. However, based on present‐day summertime SST and RST and known rates of change in these parameters, we predict present‐day tolerance thresholds would be sufficient to allow adults of these populations to tolerate extreme temperatures predicted for the next several hundred years, and that even a slow rate of change in acute tolerance thresholds might suffice to keep up with future temperature extremes.     

Intraregional variation among Alexandrium catenella (Dinophyceae) strains from southern Chile: Morphological,toxicological and genetic diversity

Morphological, toxicological and phylogenetic analyses, using the partial LSU gene and internal spacer (ITS) regions of the rDNA gene, were combined to evaluate the intraregional diversity of Alexandrium catenella occurring along the southern coast of Chile. Twenty-two strains isolated from different localities along the wide area of distribution of the species (from 42°S to 55°S) were examined by these three approaches. Morphologically, although the strains showed diagnostic characters according to the species definition, variations in these traits within and between strains were also observed. The absence of an apical or posterior attachment pore, for instance, was observed mainly in old isolates. Indirect connection between the apical and 1′ plates, traits normally seen in other species of the same genus, was also noted in some strains. However, the lack of a ventral pore on the 1′ plate was one of the most distinctive characteristics present in all the Chilean strains. Toxicologically, the Chilean strains were characterized by the dominance of N-sulfocarbamate (C1,2) and gonyautoxins (GTX1–4), but also by the scarcity or absence of saxitoxin. Considering the dominance of these toxins in each strain, at least two distinctive toxin patterns were distinguished. Through rDNA sequence analysis, the Chilean strains were segregated as part of Clade I (North American) of the Alexandrium tamarense species complex. Nevertheless, significant genetic diversity was also observed among the Chilean strains, especially using ITS sequences. Through these three approaches, Chilean strains of A. catenella showed significant intraregional variability, which is appropriate for a native species. However, the distribution of its genetic diversity seems to be inconsistent with the apparent northward expansion observed along the west coast of South America.     

BROAD THERMAL TOLERANCE OF THE SYMBIOTIC DINOFLAGELLATE SYMBIODINIUM MUSCATINEI (DINOPHYTA) IN THE SEA ANEMONE ANTHOPLEURA ELEGANTISSIMA (CNIDARIA) FROM NORTHERN LATITUDES1

The sea anemone Anthopleura elegantissima (Brandt) hosts two species of symbiotic dinoflagellates, known as zooxanthellae, which coexist within the host at southern latitudes only. One of these species, Symbiodinium muscatinei LaJeunesse et Trench, has a broad latitudinal distribution, occurring in intertidal anemones from Washington state to Southern California. To investigate whether high thermal tolerance contributes to the ability of S. muscatinei to inhabit anemones from northern and southern regions, the upper thermal tolerance limit for photosynthesis of symbionts in northern (48°24′ N) populations of A. elegantissima was determined by subjecting anemones to a gradual increase in temperature from 12°C to 30°C over a 10‐week period. Light‐saturated photosynthetic rates of isolated zooxanthellae were the same over the range of 12°C–24°C and declined significantly at 26°C, which is 14°C and 5°C above average summertime seawater temperatures in northern Puget Sound and Southern California, respectively. At 28°C, zooxanthellae isolated from the anemones, and those expelled by their hosts, exhibited extremely low rates of photosynthesis and highly reduced chl content. The photosynthetic rates and chl content of expelled zooxanthellae were lower than those of retained zooxanthellae. The high thermal tolerance of S. muscatinei isolated from northern populations of anemones supports the broad latitudinal distribution of this symbiont, allowing it to coexist with S. californium (#383, Banaszak et al. 1993 ) in southern populations of anemones.     

Are Chilean coastal forests pre‐Pleistocene relicts? Evidence from foliar physiognomy,palaeoclimate, and phytogeography

Aim We ask whether contemporary forests of the Chilean Coastal Range can be considered to be direct and conservative descendants of pre‐Pleistocene palaeofloras that occurred in southern South America from the Palaeogene to early Neogene periods (65–10 Ma), maintaining foliar physiognomies that do not match their present‐day climate. We also identify the most likely ancestors of present‐day coastal forests. Location Coastal Range of south‐central Chile (33–40° S). Methods We compared leaf morphology between five representative modern floras from mid‐latitude forests of the Chilean Coastal Range, and 14 Palaeogene–early Neogene palaeofloras from southern South America. We also compared the composition of biogeographical elements (defined by the modern distribution of plant genera) between fossil and present‐day assemblages. Palaeoclimatic reconstructions were based on a canonical correspondence analysis between leaf morphology of modern assemblages and eight climatic variables, and tested by a Monte Carlo permutation test. We compared the relative positions of fossil and modern floras on the environmental vector space defined by Canoco, and on axes defined by instrumental and estimated temperature and precipitation data. Results According to foliar characters, Palaeogene palaeofloras were strikingly divergent from present‐day coastal forests of central Chile. In contrast, two extant forest floras of the Chilean Coastal Range have a foliar morphology that resembles some late Eocene to early Miocene mixed palaeofloras, at least 23 Myr older. These two modern sites are representative of an area of the Coastal Range (36–37° S) that has been highlighted for its relictual character. None of the 14 fossil floras corresponded exactly to the modern composition of phytogeographic elements, although correspondence analyses showed that mixed and Neogene subtropical fossil floras were compositionally close to the extant woody floras of coastal forests in central Chile. Main Conclusions Contemporary forests of the Chilean Coastal Range exhibit strong physiognomic resemblance to the mixed palaeofloras from 33°57′ to 41°15′ S, which may be the closest ancestor of the deciduous and endemic‐rich Maulino forest, presently restricted to coastal areas between 36° and 38° S. In turn, the Neogene subtropical palaeoflora that occurred in the Proto‐Andean foothills of central Chile is the likely predecessor of Mediterranean‐type sclerophyllous forests of central Chile (32–33° S). Despite foliar resemblance between the late pre‐Pleistocene and extant forest floras, our palaeoclimatic reconstructions suggest that modern assemblages exist under climatic conditions that do not match their foliar physiognomy. We attribute this convergence in foliar morphology to the ‘evolutionary inertia’ of surviving lineages, favoured by the buffering effect of the coastal environment on climatic variability.