Spatial patterns of growth in the mussel, Mytilus californianus, across a major oceanographic and biogeographic boundary at Point Conception, California, USA

The Point Conception, California, USA region (hereafter PC) is one of the most important biogeographic and oceanographic discontinuities on the US west coast. Here we address how mesoscale oceanographic variability in the region around PC affects the growth of the competitively dominant species in the rocky intertidal assemblage: the mussel Mytilus californianus. Strong upwelling and high wave exposure dominate the California coast north of PC, and weak, seasonal upwelling and warmer water temperatures are characteristic of the region south/east of PC. We hypothesized that the oceanographic gradients in temperature, upwelling, wave exposure and productivity around PC would exert strong bottom–up influences on growth rates of mussels, potentially underlying large-scale differences in community structure around the PC region. We evaluated these predictions by measuring mussel growth rates across the PC region both in the intertidal and offshore on moorings. Intertidal mussels grew at much higher rates at sites south relative to north of PC and growth rates decreased in a gradient from south to north. The gradient in intertidal mussel growth around PC was uncorrelated with inshore concentrations of chlorophyll-a, and was most strongly correlated with the alongshore gradient in wave exposure and intertidal temperature. Mussels on moorings offshore from the intertidal sites grew at much higher rates than those in the corresponding intertidal areas, and mussel growth rates did not differ significantly among moored locations around PC. The gradient of increasing temperature from north to south among mooring sites was correlated with a decreasing gradient in productivity in the same direction, potentially contributing to equal and opposite effects on mussel growth at offshore moorings. This study suggests that environmental factors such as cold temperatures and high wave exposure contribute to the spatial pattern of decreasing mussel growth rates from south to north around PC, underlying large-scale patterns of community structure in this region.     

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.     

Voltinism flexibility of a riverine dragonfly along thermal gradients

Potential effects of future warming should be reflected in life history patterns of aquatic organisms observed in warmer climates or in habitats that are different in ambient temperature. In the special case of the dragonfly Gomphus vulgatissimus (L.) (Odonata: Gomphidae) previous research suggests that voltinism decreases from south to north. We analysed data on voltinism from 11 sample sites along a latitudinal gradient from about 44°N to 53°N, comprising small streams to medium-sized rivers. Furthermore, to simulate different conditions and to allow projections for future climate change scenarios, we developed a population dynamic model based on a projection matrix approach. The parameters of the model are dependent on temperature and day length. Our field results indicate a decrease in voltinism along the latitudinal gradient from southern to northern Europe and a corresponding increase of voltinism with higher temperatures. An increase in voltinism with width of the running water implies an effect of varying habitat temperature. Under the impact of global warming, our model predicts an increased development speed, particularly in the northern part of the latitudinal gradient, an extension of the northern range limit and changes in phenology of G. vulgatissimus , leading to an extension of the flight season in certain regions along the gradient.     

Arthropod community structure along a latitudinal gradient: Implications for future impacts of climate change

Abstract The structure of free‐living arthropod communities on the foliage of Acacia falcata was assessed along an extensive latitudinal gradient in eastern Australia. We hypothesized that abundance and biomass of arthropods within feeding groups would increase from temperate latitudes towards the tropics. We also hypothesized that the ratio of carnivores to herbivores would be consistent along the latitudinal gradient. Three sites at each of four latitudes, spanning 9° and 1150 km (Batemans Bay, Sydney, Grafton, Gympie in Australia), were sampled every season for 2 years, using pyrethrum knockdown. Abundance and biomass (based on dry weight) of arthropods within eight feeding groups were measured. The relative size of the feeding groups, and the ratio of carnivores to herbivores were then compared among latitudes and seasons. We found no consistent north to south (tropical to temperate) change in feeding group structure in terms of abundance. A weak latitudinal trend was evident for predator biomass, consisting of a reduction from north to south, but no significant trends in biomass for other feeding groups were found. Relative abundance and relative biomass of both carnivores and herbivores, as well as the ratio of carnivores to herbivores were consistent among latitudes. Finally, we compared a subset of these data to arthropod communities found on congeneric host species at individual sites along the latitudinal gradient. Overall, 68% of comparisons showed no significant differences in abundance or biomass within different feeding groups between host plants and among latitudes. We conclude that arthropod communities show consistencies among latitudes and between congeneric host species, in terms of feeding group and trophic structure. These results have implications for predicting the impacts of future climate change on arthropod communities.     

Latitudinal patterns and regionalization of plant diversity along a 4270‐km gradient in continental Chile

According to the global latitudinal diversity gradient, a decrease in animal and plant species richness exists from the tropics towards higher latitudes. The aim of this study was to describe the latitudinal distribution patterns of Chilean continental flora and delineate biogeographic regions along a 4270‐km north–south gradient. We reviewed plant lists for each of the 39 parallels of continental Chile to build a database of the geographical distribution of vascular plant species comprising 184 families, 957 genera and 3787 species, which corresponded to 100%, 94.9% and 74.2% of the richness previously defined for Chile, respectively. Using this latitudinal presence–absence species matrix, we identified areas with high plant richness and endemism and performed a Cluster analysis using Jaccard index to delineate biogeographic regions. This study found that richness at family, genus and species levels follow a unimodal 4270‐km latitudinal distribution curve, with a concentration of richness in central Chile (31–42°S). The 37th parallel south (central Chile) presented the highest richness for all taxonomic levels and in specific zones the endemism (22–37°S) was especially high. This unimodal pattern contrasts the global latitudinal diversity gradient shown by other studies in the Northern hemisphere. Seven floristic regions were identified in this latitudinal gradient: tropical (18–22°S), north Mediterranean (23–28°S), central Mediterranean (29–32°S), south Mediterranean (33–37°S), north temperate (38–42°S), south temperate (43–52°S) and Austral (53–56°S). This regionalization coincides with previous bioclimatic classifications and illustrates the high heterogeneity of the biodiversity in Chile and the need for a reconsideration of governmental conservation strategies to protect this diversity throughout Chile.     

Latitudinal pattern in plant composition along the Peruvian and Chilean fog oases

Fog oases in western South America (locally named lomas) are distributed in a kind of fragmented or patchy way into the coastal desert. Their origin, as well as their current ecological connections in terms of species' dispersal capability, remains an open question. We analyzed the latitudinal pattern in plant species and phylogenetic similarities of 13 lomas, which cover the latitudinal extent of these habitats, from 7°58′ to 26°15′ S. A data-set of 1004 species from available checklists was considered. Plant species composition and phylogenetic relationships among lomas were analyzed by non-metric multidimensional scaling. Our results show three main groups of lomas (northern Peruvian, southern Peruvian, and North-Central Chilean lomas) that are aligned along a complex, nonlinear north–south gradient in ordination space. The weak species overlap between Peruvian and Chilean lomas, together with the higher content in endemic species of the Chilean communities, supports the hypothesis that, at least recently, species composition of the three main groups of lomas has been shaped by desert barriers limiting plant dispersal.     

Fire persistence traits of plants along a productivity and disturbance gradient in mediterranean shrublands of south-east Australia

Aim To understand changes in fire persistence traits of plants along a latitudinal gradient, considering the interactions between productivity, community (fuel) structure and fire regime. Location A gradient in the south of Australia (latitude 33–37° S; longitude 140–143° E), including: Little Desert National Park (VIC), Big Desert Wilderness Park (VIC), Murray‐Sunset National Park (VIC), Danggali Conservation Park (SA) and Tarawi Nature Reserve (NSW). Methods We selected four areas along a latitudinal gradient for which information on fire history and vegetation was available. Then, we tested to what extent the four selected areas have different climate and different fire regimes. Plant cover values of different life forms provided an indication of the plant community structure and flammability, and the proportion of species with different fire persistence traits (resprouting, seedbank persistence) informed us on the trait selection. Results Precipitation decreases and temperature increases from south to north. Thus the selected sites represent a gradient from high productivity (low aridity) in the south to low productivity (high aridity) in the north. Fire statistics suggest that fire frequency parallels productivity. There is a tendency for life form dominance and community structure to shift in such a way that fuel connectivity is reduced towards the north. Resprouting species increase and obligate seeders decrease along the fire–productivity gradient. Main conclusions Changes in plant traits are difficult to understand without simultaneous consideration of both the disturbance and the productivity gradients. In our study area, fire regime and productivity interact in such a way that decreases in productivity imply changes in fuel structure that produce a reduction in fire frequency. Resprouting species are better represented at the high fire–productivity part of the gradient, while obligate seeders are better represented at the opposite end of the gradient. The results also emphasize the importance of considering not only climate changes but also changes in fuel structure to predict future fire regimes.     

Distribution, abundance, size and recruitment of the mussel, Mytilus californianus, across a major oceanographic and biogeographic boundary at Point Conception, California, USA

The Point Conception, California, USA region (hereafter PC) is one of the most important biogeographic and oceanographic discontinuities on the US west coast. Here we address how mesoscale oceanographic and environmental variability in the region around PC, CA may influence the distribution, abundance and size of the mussel Mytilus californianus, a competitively dominant species in rocky intertidal assemblages along the northeast Pacific. Strong upwelling and high wave exposure dominate the California coast north of PC, and weak, seasonal upwelling and warmer water temperatures are characteristic of the region south/east of PC. We hypothesized that the gradient in temperature, upwelling, and wave exposure around PC would greatly influence patterns of recruitment and abundance of mussels, potentially underlying large-scale differences in community structure. We evaluated these predictions by surveying intertidal community structure, mussel distribution, size, abundance and recruitment at a range of intertidal sites around PC. We found that intertidal communities north of PC were dominated mainly by macrophytes, while mussels and barnacles were relatively scarce. Intertidal communities south of PC were dominated by mussels and barnacles, with a low abundance of macrophytes. Mussels were larger and mussel beds were more expansive and extended lower in elevation at sites ranging from north to south around PC. At northern sites, high abundances of sea star predators and elevated wave exposure effectively displaced the entire mussel zone upwards. We found no differences in the numbers of mussel recruits to sites around PC, suggesting that spatial patterns of mussel abundance were not driven by differential recruitment. These results suggest that unlike other well-studied systems, supply of benthic larvae does not underly the large-scale gradient in community structure around PC. We suggest that environmental conditions favorable to macroalgal growth north of PC, and conditions favorable to filter-feeder growth south of PC may underly mesoscale patterns of intertidal community structure in this region.     

Climate change alters plant biogeography in Mediterranean prairies along the West Coast,USA

Projected changes in climate are expected to have widespread effects on plant community composition and diversity in coming decades. However, multisite, multifactor climate manipulation studies that have examined whether observed responses are regionally consistent and whether multiple climate perturbations are interdependent are rare. Using such an experiment, we quantified how warming and increased precipitation intensity affect the relative dominance of plant functional groups and diversity across a broad climate gradient of Mediterranean prairies. We implemented a fully factorial climate manipulation of warming (+2.5–3.0 °C) and increased wet‐season precipitation (+20%) at three sites across a 520‐km latitudinal gradient in the Pacific Northwest, USA. After seeding with a nearly identical mix of native species at all sites, we measured plant community composition (i.e., cover, richness, and diversity), temperature, and soil moisture for 3 years. Warming and the resultant drying of soils altered plant community composition, decreased native diversity, and increased total cover, with warmed northern communities becoming more similar to communities further south. In particular, after two full years of warming, annual cover increased and forb cover decreased at all sites mirroring the natural biogeographic pattern. This suggests that the extant climate gradient of increasing heat and drought severity is responsible for a large part of the observed biogeographic pattern of increasing annual invasion in US West Coast prairies as one moves further south. Additional precipitation during the rainy season did little to relieve drought stress and had minimal effects on plant community composition. Our results suggest that the projected increase in drought severity (i.e., hotter, drier summers) in Pacific Northwest prairies may lead to increased invasion by annuals and a loss of forbs, similar to what has been observed in central and southern California, resulting in novel species assemblages and shifts in functional composition, which in turn may alter ecosystem functions.     

Shallow subtidal macroalgae in the North-eastern Atlantic archipelagos (Macaronesian region): a spatial approach to community structure

Shallow subtidal macroalgal communities in the North-eastern Atlantic archipelagos (Azores, Madeira, Canaries and Cape Verde) were studied in order to identify their spatial organization patterns and the main drivers of change. Fifteen islands and 145 sites across 15º of latitude and 2850 km were sampled. We found high spatial variability across the scales considered (archipelago, island and site). The structure of macroalgal communities differed among archipelagos, except between Madeira and the Canaries, which were similar. Across a latitudinal gradient, macroalgal communities in the Azores were clearly separated from the other archipelagos; communities in Madeira and the Canaries occupied an intermediate position, while those in Cape Verde appeared at the opposite end of the gradient. In the Azores, species with warm-temperate affinities dominated communities. Cape Verde communities were, in contrast, dominated by tropical taxa, whereas in the subtropical Canaries and Madeira there was a mixture of species with colder and warmer affinities. Apart from crustose coralline algae, the Dictyotales were the group with greatest cover; larger and longer-lived species were progressively replaced by short-lived species along a latitudinal gradient from north to south. The perennial species Zonaria tournefortii dominated the sea-bottom in the Azores, the semi-perennial Lophophora variegata in the Canaries, the filamentous algae in Madeira and the ephemeral Dictyota dichotoma in Cape Verde. We hypothesized that the differences among archipelagos could be explained by synergies between temperature and herbivory, which increased in diversity southwards, especially in Cape Verde. This was supported by the predominance of non-crustose macroalgae in the Azores and of crustose macroalgae in Cape Verde, as would be predicted from the greater herbivore activity. At the scale of islands and sites, the same set of environmental variables drove differences in macroalgal community structure across all the Macaronesian archipelagos.     

Phylogeography of the tidewater goby, Eucyclogobius newberryi (Teleostei, Gobiidae), in coastal California

The tidewater goby, Eucyclogobius newberryi, inhabits discrete, seasonally closed estuaries and lagoons along approximately 1500 km of California coastline. This species is euryhaline but has no explicit marine stage, yet population extirpation and recolonization data suggest tidewater gobies disperse intermittently via the sea. Analyses of mitochondrial control region and cytochrome b sequences demonstrate a deep evolutionary bifurcation in the vicinity of Los Angeles that separates southern California populations from all more northerly populations. Shallower phylogeographic breaks, in the vicinities of Seacliff, Point Buchon, Big Sur, and Point Arena segregate the northerly populations into five groups in three geographic clusters: the Point Conception and Ventura groups between Los Angeles and Point Buchon, a lone Estero Bay group from central California, and San Francisco and Cape Mendocino groups from northern California. The phylogenetic relationships between and patterns of molecular diversity within the six groups are consistent with repeated, and sometimes rapid, northward and southward range expansions out of central California caused by Quaternary climate change. Plio-Pleistocene tectonism, Quaternary coastal geography and hydrography, and historical human activities probably also influenced the modern geographic and genetic structure of E. newberryi. The phylogeography of E. newberryi is concordant with phylogeographic patterns in several other coastal California taxa, suggesting common extrinsic factors have had similar effects on different species. However, there is no evidence of a phylogeographic break coincident with a biogeographic boundary at Point Conception.     

Pigment variability in larval Sebastes jordani off central California

Atypical pigment was observed in pre‐flexion, flexion and post‐flexion larval Sebastes collected off central California in 1991 and 1992 that otherwise resembled Sebastes jordani . Atypical pigment occurred on the axillary region at the base of the pectoral fin (most prominent on the inner edge of the pectoral fin base adjacent to the gut), on the median and distal regions of the pectoral fin and on the median and distal regions of the pelvic fin. In addition, lower and upper jaw pigment was observed at a much smaller size in these specimens than previously described in the literature. Identifications of these atypical specimens as S. jordani were confirmed using meristic and otolith characters as well as mitochondrial DNA sequence data. The ontogenetic variability of S. jordani is described. Specimens collected north of Point Conception were more pigmented than larval S. jordani described in the literature (collected predominately south of Point Conception), suggesting geographic variation in pigment development during the larval stage.     

Cryptic speciation in the vermilion rockfish (Sebastes miniatus) and the role of bathymetry in the speciation process

A recent phylogenetic review of the genus Sebastes suggested the existence of a cryptic species of vermilion rockfish ( Sebastes miniatus ). To evaluate the geographical and bathymetric range of the Type 1 and Type 2 forms reported in that study, cytochrome b sequences were examined from 548 fish. Type 1 fish were found primarily south of Point Conception on reefs deeper than 100 m. Type 2 fish were common range-wide at sites shallower than 100 m. Reproductive isolation between the two types was tested using nine microsatellite loci. Estimates of genetic divergence were made using the fixation index ( F _ST) and correspondence between haplotype and genotype was tested by Bayesian population assignment and multivariate plotting of individual genotypes. Microsatellite analyses gave strong support for the presence of two distinct groups of genotypes. All fish with Type 1 haplotypes and fish with Type 2 haplotypes from < 100 m depth had genotypes unique to their haplotype group. However, most (68%) fish with Type 2 haplotypes from > 100 m depth assigned strongly to the Type 1 genotype group. Morphometric comparisons between the two genotypic groups revealed significant differences at three of the six examined measurements. Differences in both genetics, depth of occurrence, and morphology suggest these are separate species. This observation along with evidence of depth segregation in many recent species pairs led us to hypothesize a speciation model for Sebastes spp. by which the loss or truncation of a depth-related ontogenetic migration can lead to the creation of reproductively isolated populations.     

Latitudinal influence on gametogenesis and host–parasite ecology in a marine bivalve model

Reproduction and parasites have significant impacts on marine animal populations globally. This study aimed to investigate the associative effects of host reproduction and a host–parasite interplay on a marine bivalve, along a geographic gradient of latitude. Cockles Cerastoderma edule were sampled from five European sites (54°N to 40°N), between April 2018 and October 2019. A histological survey provided data on trematode (metacercaria and sporocyst life stages), prevalence, and cockle stage of gametogenesis to assess the influence of a latitudinal gradient on both interplays. Sex ratios at the northernmost sites were skewed toward females, and spawning size was reduced at the lower latitudes. Trematode infection did not follow a latitudinal gradient. Localized site‐related drivers, namely seawater temperature, varied spatially, having an impact on cockle–trematode interactions. Spawning was related to elevated temperatures at all sites. Prolonged spawning occurred at southern latitudes, where seawater temperatures were warmer. Trematode prevalence and the impact of trematodes on gametogenesis were found to be spatially variable, but not latitudinally. Therefore, it is not possible to determine the likelihood of boom and bust events in cockles, based on the latitudinal location of a population. In terms of sublethal impacts, it appeared that energy was allocated to reproduction rather than somatic growth in southern populations, with less energy allocated to reproduction in the larger, northern cockles. The demonstrated spatial trend of energy allocation indicates the potential of a temporal trend of reduced cockle growth at northern sites, as a result of warming sea temperatures. This awareness of the spatially varying drivers of populations is crucial considering the potential for these drivers/inhibitors to be exacerbated in a changing marine environment.     

Testing latitudinal patterns of tidepool fish assemblages: local substrate characteristics affect regional-scale trends

We tested the regional-scale latitudinal patterns of occurrence and community structure of rocky tidepool fishes in relation to local environmental factors, particularly substrate characteristics. Data were derived from intensive field observations conducted on 36 shore sites spread across ca. 1,000 km north–south (24°03′N–32°45′N) in the south-western Japan. While numbers of families, genera, species and individuals per unit area decreased with latitude, these were dependent on substrate types: sites with non-limestone rock substrates tended to harbour larger numbers than limestone sites at the same latitude. Relative abundances of two dominant families (Blenniidae and Gobiidae) varied among sites with weak latitudinal gradients. Species-specific trends of latitudinal distribution were observed in most of the common intertidal fishes, with over half of the species demonstrating substrate-dependent variation. Species composition was clearly different between the Kyushu Island and the Ryukyus and also between limestone and non-limestone sites. Thus, our results clearly demonstrated that the regional-scale latitudinal trends of tidepool fish assemblages were partially dependent on local environmental characteristics (substrate types). Consideration was given to the influences of the Kuroshio Current and other factors including species interactions that might have helped modify observed latitudinal patterns.     

Arctic plant diversity in the Early Eocene greenhouse

For the majority of the Early Caenozoic, a remarkable expanse of humid, mesothermal to temperate forests spread across Northern Polar regions that now contain specialized plant and animal communities adapted to life in extreme environments. Little is known on the taxonomic diversity of Arctic floras during greenhouse periods of the Caenozoic. We show for the first time that plant richness in the globally warm Early Eocene (approx. 55-52 Myr) in the Canadian High Arctic (76° N) is comparable with that approximately 3500 km further south at mid-latitudes in the US western interior (44-47° N). Arctic Eocene pollen floras are most comparable in richness with today's forests in the southeastern United States, some 5000 km further south of the Arctic. Nearly half of the Eocene, Arctic plant taxa are endemic and the richness of pollen floras implies significant patchiness to the vegetation type and clear regional richness of angiosperms. The reduced latitudinal diversity gradient in Early Eocene North American plant species demonstrates that extreme photoperiod in the Arctic did not limit taxonomic diversity of plants.     

On the importance of habitat continuity for delimiting biogeographic regions and shaping richness gradients

The formation and maintenance of biogeographic regions and the latitudinal gradient of species richness are thought to be influenced, in part, by the spatial distribution of physical habitat (habitat continuity). But the importance of habitat continuity in relation to other variables for shaping richness gradients and delimiting biogeographic regions has not been well established. Here, we show that habitat continuity is a top predictor of biogeographic structure and the richness gradient of eastern Pacific rocky shore gastropods (spanning c. 23 000 km, from 43°S to 48°N). Rocky shore habitat continuity is generally low within tropical/subtropical regions (compared to extratropical regions), but particularly at biogeographic boundaries where steep richness gradients occur. Regions of high rocky shore habitat continuity are located towards the centres of biogeographic regions where species turnover tends to be relatively low. Our study highlights the importance of habitat continuity to help explain patterns and processes shaping the biogeographic organisation of species.