Benthic foraminifera associated with the invasive ascidian, Didemnum sp. A

The invasive ascidian, Didemnum sp. A, first appeared in New England bays and harbors in the early 1990s, and in the waters around Cape Cod in 1993. While ship traffic was the likely vector introducing the species, its origin and precise date and location of its introduction are presently unknown. Colony surfaces of Didemnum sp. A appear very clean and not favorable substrates for epibiota settlement, but closer inspection revealed the presence of benthic foraminifera. During 2003 and 2004, 52 samples of Didemnum sp. A and other ascidians were collected to determine whether or not the foraminiferal assemblages might also be non-native and thus provide a potential clue to the place of origin of Didemnum sp. A. Sample locations included the New England coast from Connecticut to Maine (with a concentration in the Cape Cod area), northern California, Zeeland, The Netherlands, and Shakespeare Bay, New Zealand. From New England samples, 18 species of benthic foraminifera were identified. The most common species represented were Cornuspira involvens, C. planorbis, Elphidium galvestonense, E. margaritaceum, Glabratellina lauriei, Miliolinella subrotunda, Quinqueloculina bicornis, and Rosalina floridana. Foraminiferal assemblages on Didemnum sp. A from other regions sampled were composed of the same cosmopolitan species found in New England, plus other species which were indigenous to each region. Because no exotic foraminifera species were found it is concluded that Didemnum sp. A likely did not introduce non-native foraminifera originating from their native habitats into the New England region.     

Ecological observations on the colonial ascidian Didemnum sp. in a New England tide pool habitat

The colonial ascidian Didemnum sp. has colonized northwestern Atlantic coastal habitats from southern Long Island, New York, to Eastport, Maine. It is also present in offshore habitats of the Georges Bank fishing grounds. It threatens to alter fisheries habitats and shellfish aquacultures.Observations in a tide pool at Sandwich, MA from December 2003 to February 2006 show that Didemnum sp. tolerates water temperatures ranging from ≤ 1 to > 24 °C, with daily changes of up to 11 °C. It attaches to pebbles, cobbles, and boulders, and it overgrows other tunicates, seaweeds, sponges, and bivalves. From May to mid July, colonies appear as small patches on the bottoms of rocks. Colonies grow rapidly from July to September, with some growth into December, and they range in color from pink to pale yellow to pale orange. Colony health declines from October through April, presumably in response to changes in water temperatures, and this degenerative process is manifested by color changes, by the appearance of small dark brown spots that represent clumps of fecal pellets in the colony, by scavenging by periwinkles, and by a peeling-away of colonies from the sides of cobbles and boulders. At Sandwich, colonies died that were exposed to air at low tide. The species does not exhibit this seasonal cycle of growth and decline in subtidal habitats (40-65 m) on the Georges Bank fishing grounds where the daily climate is relatively stable and annual water temperatures range from 4 to 15 °C. Experiments in the tide pool with small colony fragments (5 to 9 cm²) show they re-attach and grow rapidly by asexual budding, increasing in size 6- to 11-fold in the first 15 days. Didemnum sp. at Sandwich has no known predators except for common periwinkles (Littorina littorea) that graze on degenerating colonies in the October to April time period and whenever colonies are stressed by desiccation.The tendencies of the ascidian (1) to attach to firm substrates, (2) to rapidly overgrow other species, (3) to tolerate a wide temperature range, (4) to be free from predation, and (5) to spread by colony fragmentation combine to make it a potential threat to benthic marine habitats and aquacultures. Didemnum sp. is known to overgrow mussels, oysters, and sea scallops, and it likely envelops other bivalves too.     

Variation in the ability of Didemnum sp. to invade established communities

Over the past 30 years southern New England, USA has been invaded by several species of ascidians, including Botrylloides violaceus, Diplosoma listerianum, Styela clava, and Ascidiella aspersa. These species have become dominate in coastal embayments and marinas but are usually absent from more open water coastal areas. A colonial ascidian, Didemnum sp. has invaded southern New England during the past 10 years and we first observed this species in eastern Long Island Sound in 2000. It has become the dominant at several field sites while remaining in low abundance at others. We conducted an experiment at two places, a protected marina and an open coast site, to examine its ability to compete with the established fouling community. Small colonies of Didemnum were transplanted onto panels with communities that varied in age from one to four weeks old and these treatments along with controls with only Didemnum were exposed at both sites. In most treatments Didemnum became a dominant species in the communities at both sites but it reached higher abundances at the open coast site. Potential causes of the observed differences are predation on other species of ascidians at the open coast site reducing recruitment of these species and competition, lower tolerance for elevated temperatures at the marina site, or other environmental parameters that might affect growth rates.     

Introductions and developments of oysters in the North Sea area: a review

To replenish the exploited native stocks ofOstrea edulis, imports from almost all European coasts have arrived in the North Sea, particularly in the Dutch Oosterschelde estuary. The American oysterCrassostrea virginica and the Portuguese oysterC. angulata have also been imported into the North Sea several times. However, only the introductions of various genetic strains of the Pacific oysterC. gigas have been of lasting success. Spat from British Columbia (Canada) was first imported to the Oosterschelde in 1964. First spatfalls in this area took place in the warm summers of 1975 and 1976. Further larval outbursts occurred in 1982 and 1989, and good settlements took place on culture plots as well as along the dikes of the Oosterschelde. Since 1977 no more cupped oysters have been imported from overseas. The population maintained itself and was able to spread in a northern direction along the Dutch North Sea coast. In Britain, hatchery-producedC. gigas were transferred to several sites, in the 1970s including the British North Sea coast. Here, occurrence in the wild seems to be rather limited up to now. Into the German Wadden Sea,C. gigas larvae and spat from a Scottish hatchery have been introduced since 1971, as were medium-sized oysters from a variety of European sources in the 1980s. Strong spatfalls on intertidal mussel beds in the northern German Wadden Sea occured in 1991 and 1994. For the introductions ofC. gigas along the west European coasts, precautionary measures to minimize unintentional transfers of associated organisms, as recommended by the ICES Code of Practice in 1994, came too late.     

The contribution of marine snow to the particle food supply of the benthic suspension feeder, Mytilus edulis

In order to examine the importance of the settling of large particles to the food supply and feeding behavior of a benthic culture of the blue mussel, Mytilus edulis, we investigated the tidal dynamics of large (>0.5 mm diameter) marine aggregates, commonly known as marine snow, during three tidal cycles in July 1998 at a shallow, subtidal, low current flow regime site along the coast of Maine (Shorey Cove, Roque Island, Englishman's Bay, Maine).In situ, optically measured marine snow showed a distinct tidal signal displaying an increase in size and abundance through high tide with a peak on the early ebb tide as it settled to the bottom. Marine snow volume ranged an order of magnitude through the tidal cycle, from under 8 to over 80 mm³ l⁻¹. An increase in the in situ marine snow volume corresponded with an increase in benthic mussel feeding activity (from 20% to 60% of maximum exhalant siphon area, which is an estimate of pumping rate) and maximum rates of pseudofeces production by the mussels during periods of low tidal current speeds. In contrast, mussels from the same population feeding on surface waters in shipboard chambers produced no pseudofeces and had high pumping rates (80-100% maximum exhalant siphon area) over the whole tidal period. A second peak in benthic mussel pumping rates also occurred during flood tide.Food quality was lower in the bottom waters due to significantly higher particulate inorganic matter (PIM, >2 mg l⁻¹) when compared with the surface waters. PIM accounted for 95% of the total settled mass flux of 3.4 g m⁻² day⁻¹ measured in sediment traps deployed 1 m off the bottom, with organic carbon representing only 2.5% of the mass flux during the mid-summer conditions. At low-current sites such as Shorey Cove, Roque Island, Maine, the settling of marine snow provides an important additional source of food, albeit of low quality, to benthic populations of blue mussels.     

Distribution and long-term temporal patterns of four invasive colonial ascidians in the Gulf of Maine

Invasive ascidians are a growing concern for ecologists and natural resource managers, yet few studies have documented their short- and long-term temporal patterns of abundance. This study focuses on the invasion of the Gulf of Maine by the colonial ascidians Botryllus schlosseri, Botrylloides violaceus, Diplosoma listerianum and Didemnum sp. A. We examined the time of arrival and potential vectors for these four invasive ascidians using survey data (collected from 1969 onwards) and literature documentation. We also compared the dominance and seasonal patterns of abundance of these species using data from two identical panel studies; one conducted from 1979 to 1980, the other from 2003 to 2005. Didemnum and Botrylloides were most likely first introduced into the Damariscotta River, Maine in the early 1970's through oyster aquaculture while Botryllus and Diplosoma were probably transported by commercial and recreational vessels. The overall abundance of colonial ascidians has increased since 1979 and 1980. Botryllus was the only invasive colonial ascidian present during the 1979 to 1980 study and accounted for an average of 6.16% cover on panels. From 2003 to 2005, the more recently arrived colonial ascidians Botrylloides and Didemnum accounted for 7.38% and 2.08% cover respectively, while Botryllus covered only 1.16%. Our results reveal a shift in seasonal abundance between 1979 to 1980 and 2003 to 2004. In 1979 and 1980, colonial ascidians had the highest percent cover in fall and winter while in 2003 and 2005 they had highest percent cover in summer and fall. Seasonal patterns of space occupation by colonial ascidians were correlated with seasonal changes in seawater temperature.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

European green crabs (Carcinus maenas) in the northeastern Pacific: genetic evidence for high population connectivity and current-mediated expansion from a single introduced source population

Aim The European green crab (Carcinus maenas) expanded dramatically after its introduction to the west coast of North America, spreading over 1000 km in < 10 years. We use samples of Carcinus maenas collected over time and space to investigate the genetic patterns underlying the species’ initial establishment and spread, and discuss our findings in the context of the species’ life history characteristics and demography. Location The central west coast of North America, encompassing California, Oregon, and Washington (USA) and British Columbia (Canada). Methods We collected 1040 total samples from 21 sites representing the major episodes of population establishment and expansion along the west coast of North America. Microsatellite markers were used to assess genetic diversity and structure at different time points in the species’ spread, to investigate connectivity between embayments and to estimate both short‐term effective population sizes and the number of original founders. Assignment testing was performed to determine the likely source of the introduction. Results Carcinus maenas in western North America likely derived from a single introduction of a small number of founders to San Francisco Bay, CA from the east coast of North America. Throughout its western North American range, the species experiences periodic migration between embayments, resulting in a minor loss of genetic diversity in more recently established populations versus the populations in the area of initial establishment. Main conclusions Low genetic diversity has not precluded the ability of C. maenas to successfully establish and spread on the west coast of North America. An efficient oceanographic transport mechanism combined with highly conducive life history traits are likely the major drivers of C. maenas spread. Evidence for a single introduction underscores the potential utility of early detection and eradication of high‐risk invasive species.     

Ecological interaction between sympatric Mytilus species on the west coast of Canada investigated using PCR markers

M. californianus is the dominant marine mussel species on exposed rocky shores, while M. trossulus is usually the dominant mussel species in more sheltered waters on the west coast of North America. Since these species are physically indistinguishable when small (< 10.0 mm), we developed two polymerase chain reaction (PCR) -based markers to discriminate between them. Using these markers, we identified mussels taken from an exposed coast ( n = 114), a sheltered harbour ( n = 80), and an upper-intertidal pool ( n = 42) on the west coast of Vancouver Island, British Columbia, Canada. M. californianus were found only on the open coast. Small M. trossulus (< 20.0 mm) were common to all three sample sites, but were extremely rare at larger sizes (> 20.0 mm) on the open coast. Our results indicate that M. californianus are excluded from sheltered waters via early life factors, while M. trossulus are excluded from the open coast due to mortality later in life.     

Monarch butterflies cross the Appalachians from the west to recolonize the east coast of North America

Each spring, millions of monarch butterflies (Danaus plexippus) migrate from overwintering sites in Mexico to recolonize eastern North America. However, few monarchs are found along the east coast of the USA until mid-summer. Brower (Brower, L. P. 1996 J. Exp. Biol. 199, 93–103.) proposed that east coast recolonization is accomplished by individuals migrating from the west over the Appalachians, but to date no evidence exists to support this hypothesis. We used hydrogen (δD) and carbon (δ¹³C) stable isotope measurements to estimate natal origins of 90 monarchs sampled from 17 sites along the eastern United States coast. We found the majority of monarchs (88%) originated in the mid-west and Great Lakes regions, providing, to our knowledge, the first direct evidence that second generation monarchs born in June complete a (trans-) longitudinal migration across the Appalachian mountains. The remaining individuals (12%) originated from parents that migrated directly from the Gulf coast during early spring. Our results provide evidence of a west to east longitudinal migration and provide additional rationale for conserving east coast populations by identifying breeding sources.     

Mitochondrial ATPase 6/8 genes reveal genetic divergence in the Coilia dussumieri (Valenciennes, 1848) populations of north east and northwest coasts of India

The golden anchovy, Coilia dussumieri, though possessing discontinuous distribution along northeast and northwest coasts of India, it is being managed as unit stock for fishery assessment purposes. By considering the need for stock specific management of the species, mitochondrial ATP synthase 6 and 8 (ATPase 6/8) genes were analyzed for delineating genetic stock structure of the species. Sequence analysis revealed a total of 34 haplotypes across four populations from both the east and west coasts of India. Haplotype diversity (h) was found in the range of 0.7421–0.9368. Similarly, nucleotide diversity (π) varied from 0.0012 to 0.0025. AMOVA results indicated a high total variance of 72.66 % between east and west coast populations and less (1.34 %) among populations within the respective coast. Phylogenetic tree constructed using pair wise F_ST also indicated the genetic divergence of populations of east and west coasts of India. The findings of the present study will be helpful in developing stock specific management measures for conservation and sustainable utilization of the species.     

Patterns of larval release in the Florida stone crab, Menippe mercenaria

Larval release patterns in brachyuran crabs are often synchronized with environmental cycles. While previous studies have focused extensively on supratidal and intertidal taxa, there have been relatively few investigations of subtidal species. This study examined patterns of larval release by the Florida stone crab, Menippe mercenaria, from three different tidal regimes. Ovigerous stone crabs were collected from Sebastian Inlet on the east coast of Florida, Tampa Bay on the west coast of Florida, and the Florida Keys. Patterns of larval release were monitored in the laboratory in relation to local tidal and diel cycles. Results showed a significant diel pattern in initiation of hatching by crabs from each of three study areas. Larval release consistently occurred during the diurnal phase despite the maintenance of females in constant laboratory conditions for up to 96 h prior to hatching. This implies that release may be controlled by a circadian clock. Patterns of release by stone crabs in relation to tidal cycle were more variable. Larval release by females from populations near Tampa Bay and Sebastian Inlet were not synchronized with the tides, whereas females collected from the Florida Keys exhibited a pattern that was strongly related to tidal cycle. These results may be explained by differences in tidal amplitude at the three sampling locations.     

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.     

Will the invasive mussel Mytilus galloprovincialis Lamarck replace the indigenous Perna perna L. on the south coast of South Africa?

The mussel Mytilus galloprovincialis is invasive worldwide, has displaced indigenous species on the west coast of South Africa and now threatens Perna perna on the south coast. We tested the hypothesis that Mytilus will replace Perna by examining changes in their distribution on shores where they co-exist. Total cover, adult density, recruit density, recruit/adult correlations and mean maximum lengths of both species were measured in 2001 at two contrasting sites (Plettenberg Bay and Tsitsikamma) 70 km apart, each including two locations 100 m apart. Cover and density were measured again in 2004. Total mussel abundance was significantly lower in Tsitsikamma, and recruit density was only 17% that of Plettenberg Bay. Abundance and cover increased upshore for Mytilus, but decreased for Perna, giving Mytilus higher adult and recruit density and total cover than Perna in the upper zones. Low shore densities of recruits and adults were similar between species but cover was lower for Mytilus, reflecting its smaller size, and presumably slower growth or higher mortality there. Thus, mechanisms excluding species differed among zones. Recruitment limitation delays invasion at Tsitsikamma and excludes Perna from the high shore, while Mytilus is excluded from the low shore by post-recruitment effects. Recruitment limitation also shapes population structure. Recruit/adult correlations were significant only where adult densities were low, and this effect was species-specific. Thus, at low densities, larvae settle or survive better near adult conspecifics. After 3 years, these patterns remained strongly evident, suggesting Mytilus will not eliminate Perna and that co-existence is possible through partial habitat segregation driven by recruitment limitation of Perna on the high shore and post-settlement effects on Mytilus on the low shore.     

Comparative phylogeography reveals host generalists, specialists and cryptic diversity: Hexabothriid, microbothriid and monocotylid monogeneans from rhinobatid rays in southern Australia

The phylogeography and host specificity of three monogenean species infecting different sites on the southern fiddler ray, Trygonorrhina fasciata (Rhinobatidae) in South Australia (SA) were studied: Branchotenthes octohamatus (Hexabothriidae: gills), Calicotyle australis (Monocotylidae: cloaca) and Pseudoleptobothrium aptychotremae (Microbothriidae: skin). Five rhinobatid species (Aptychotrema vincentiana, T. fasciata, Trygonorrhina sp. A, Aptychotrema rostrata and Rhinobatos typus) with distributions spanning west, south and east Australian coastal waters, were surveyed for monogeneans resembling the three species documented from T. fasciata in SA. The identities of hosts and parasites collected were investigated using the mitochondrial genes ND4 and Cytochrome b (cytb), respectively, in addition to the nuclear marker, Elongation factor 1-alpha (EF1a) for Pseudoleptobothrium. Genetic analyses confirmed that B. octohamatus is geographically widespread and displays little genetic structure, suggesting high levels of gene flow. It was collected from four rhinobatid species throughout its distribution and is not, therefore, host specific. For C. australis, genetic analyses revealed two discrete populations with a genetic divergence of ∼4%, one population occurring west of Bass Strait on two sympatric host species and the other population on the east coast, also occurring on sympatric host species. Similarly, for Pseudoleptobothrium, specimens collected west of Bass Strait were genetically distinct (∼3.5%) from those collected to the east. However, on the east coast, a third Pseudoleptobothrium population was revealed, separated by a genetic distance of >11%, indicating a morphologically cryptic species. Host preferences were indicated for each Pseudoleptobothrium lineage. These genetic discoveries are discussed in relation to life history characteristics of each monogenean species, highlighting the value of phylogeographic analyses to understand the parasite-host relationship.     

Comparative ecology of subtidal actiniarians from the coasts of California and the Gulf of Maine,USA

Shallow subtidal populations of sea anemones occupying hard substrates along the California (Pacific) and Gulf of Maine (Atlantic) coasts of the USA are compared. The diversity of anemones in the southern Gulf of Maine to 35 m is low (4 species) in contrast to similar habitats in California (at least 11 species), but parallel patterns of individual size, distribution and predation impacts occur. Water motion influences depth distribution and size, while predation affects size, population structure and habitat selection. The nudibranch Aeolidia papillosa exerts a similar pressure on population structure in some anemone species in both areas as a size selective predator. The seastar Dermasterias imbricata appears to magnify the selective pressure for large size in the eastern Pacific coastal zone. Interspecific aggression of anemones determines some distribution patterns in areas of overlap.     

The identification of a new Giardia duodenalis assemblage in marine vertebrates and a preliminary analysis of G. duodenalis population biology in marine systems

Giardia duodenalis is an intestinal parasite of many vertebrates. The presence of G. duodenalis in the marine environment due to anthropogenic and wildlife activity is well documented, including the contributions from untreated sewage and storm water, agricultural run-off and droppings from terrestrial animals. Recently, studies have detected this protistan parasite in the faeces of marine vertebrates such as whales, dolphins, seals and shore birds. To explore the population biology of G. duodenalis in marine life, we determined the prevalence of G. duodenalis in two species of seal (Halichoerus grypus, Phoca vitulina vitulina and Phoca vitulina richardsi) from the east and west coasts of the USA, sequenced two loci from G. duodenalis-positive samples to assess molecular diversity and examined G. duodenalis distribution amongst these seals and other marine vertebrates along the east coast. We found a significant difference in the presence of G. duodenalis between east and west coast seal species. Only the zoonotic lineages of G. duodenalis, Assemblages A and B and a novel lineage, which we designated as Assemblage H, were identified in marine vertebrates. Assemblages A and B are broadly distributed geographically and show a lack of host specificity. Only grey seal (Halichoerus grypus) samples and one gull sample (Larus argentatus) from a northern location of Cape Cod, Massachusetts, USA, showed the presence of Assemblage H haplotypes; only one other study of harbour seals from the Puget Sound region of Washington, USA previously recorded the presence of an Assemblage H haplotype. Assemblage H sequences form a monophyletic clade that appears as divergent from the other seven Assemblages of G. duodenalis as these assemblages are from each other. The discovery of a previously uncharacterised lineage of G. duodenalis suggests that this parasite has more genetic diversity and perhaps a larger host range than previously believed.     

Early devonian flora of the trout valley formation of northern maine

The data on the plant fossils of the Trout Valley Formation of north-central Maine, U.S.A., are summarized. Evidence from lithological correlation, from plant megafossils and paleopalynology suggest a late Early Devonian (Emsian) age for the Formation. Plant megafossils present are several species of Psilophyton, Pertica quadrifaria, Kaulangiophyton akantha, Prototaxites sp., Drepanophycus sp., Taeniocrada sp. and Thursophyton sp. The microfossil assemblage, through poorly preserved, contains Deltoidospora sp., Apiculiretusispora sp., Emphanisporites rotatus, E. annulatus, Clivosisporites cf. C. verrucata, Grandispora sp. and a chitinozoan, Sphaerochitina sp. The ancient landscape is suggested as having been an area a few square miles in extent with modest relief and comparable to a modern brackish or fresh-water marsh. Three other floras, one in Maine, a second in New Brunswick, Canada, and a third along the coast of Gaspé Bay, Quebec, are thought to have been at least partially contemporaneous with the Trout Valley flora.     

Holocene sand beaches of southern California: ENSO forcing and coastal processes on millennial scales

A proxy record of sand beach accretion for the past 10,000 years has been assembled from radiocarbon dates on the Pismo clam, Tivela stultorum, in archaeological sites along the southern California coast. When this record is compared with numerous climate proxies, it appears that El Niño-Southern Oscillation (ENSO) controls on wave climate and sediment flux have acted upon regional geomorphology at different sea levels to either accrete or erode the Holocene beaches of southern California. Tivela dates from the Santa Maria coast indicate that perennial sand beaches built by 9000 years ago in response to abundant riverine sediment contained by the natural groin at Point Sal, wave sheltering by the massive headland of Point Buchon, and Early Holocene El Niño events. On the western Santa Barbara coast, sand beaches were forming by 7000 years ago in response to high sand fluxes from the Santa Ynez Mountains to the many small littoral catchments, possibly aided by high local rates of uplift. Decline of these sand beach habitats 5000-4000 years ago coincides with increased El Niño-driven wave energy. In accord with slowing in sea-level rise ca. 6000 years ago, sand beaches were most widespread in the period 6000-5000 years ago on Estero Bay, the western Santa Barbara coast, and west of Point Dume. However, Tivela dates first appear 5000 years ago in the Oceanside and Silver Strand littoral cells of the San Diego region. This lag coincides with the Middle Holocene shift to a more variable climate and modern periodicity in El Niño events that increased sediment supply to the southern coast. The ontogeny of the littoral cells provides timelines for modeling coastal evolution with implications for sand beach ecology, prehistoric human coastal adaptations, and coastal planning for future climate change.