The settlement of intertidal barnacles
Chthamalus
spp. was measured for an entire recruitment season at 3 sites separated by 100 m in Baja California, Mexico. During a 10 d pulse that accounted for ...nearly 30% of yearly settlement, coastal wind speed and direction, nearshore water column temperature, and current velocities were also measured. During this period, strong internal tidal forcing was observed, with short episodes (1-1.5 h) of rapid fluctuations in water column temperature, stratification, and currents.
Chthamalus
spp. settlement was significantly and positively correlated with cumulative high-frequency fluctuations in temperature, thermal stratification, and surface current flows, but not with onshore winds. Furthermore, the spatial pattern in the number of settlers was correlated with the angle of propagation estimated for onshore-moving internal waves, potentially modulated by nearshore bathymetry. This relationship between
Chthamalus
spp. settlement and high-frequency changes in water column temperature was also shown for a site in La Jolla, California, USA, but not for a similar barnacle,
Balanus glandula
, suggesting that interspecific differences in larval behavior and nearshore depth distribution may translate into differences in transport. Our results suggest that both the number of settlers at shore and their spatial pattern can be modulated by the intensity and direction of internal wave events, at least during peak settlement periods. Future research should consider both the intensity and geometry of internal-wave-induced variability in the nearshore, as well the small-scale features of coastal geomorphology and bathymetry.
The Pacific coast of the Baja California peninsula (Mexico) is a unique biogeographic transition zone, where many kelp species encounter their southern subtropical limits and have experienced a range ...contraction in the northern hemisphere. Using reciprocal transplant experiments with giant kelp populations from their southern limit and a northern temperate site on the Baja California peninsula, we hypothesized that juveniles from the southern site would grow longer, have more fronds (an individual stipe with its blades), and have higher photosynthetic efficiency (α) and blade tissue nitrogen, especially under warmer nitrate-limited field conditions experienced in their habitat. At the southern-limit transplant site, local juveniles had more fronds compared with those from the north after 90 days in field conditions (warm waters). At the northern transplant site, blade tissue nitrogen of juveniles from both populations was low after 23 days in field conditions (not significantly different during a warm water period), however, after a further 56 day cooling period and shortly after a strong upwelling event, juveniles from the southern population had greater blade tissue nitrogen concentrations than those from the north, which showed little change. Respiration and a were both greater at higher temperatures in juveniles from the southern population, and this pattern was maintained under warm water field conditions. These physiological differences in conspecific juveniles from different populations may be driven by population-level adaptation to frequently occurring thermal and nutrient stress in southern Baja California, which is the southern distribution limit for giant kelp. Further exploration of population level differences in ecophysiology should be applied to kelp forest restoration efforts as oceans warm.
Inbreeding, the mating between genetically related individuals, often results in reduced survival and fecundity of offspring, relative to outcrossing. Yet, high inbreeding rates are commonly observed ...in seaweeds, suggesting compensatory reproductive traits may affect the costs and benefits of the mating system. We experimentally manipulated inbreeding levels in controlled crossing experiments, using gametophytes from 19 populations of Macrocystis pyrifera along its Eastern Pacific coastal distribution (EPC). The objective was to investigate the effects of male–female kinship on female fecundity and fertility, to estimate inbreeding depression in the F1 progeny, and to assess the variability of these effects among different regions and habitats of the EPC. Results revealed that the presence and kinship of males had a significant effect on fecundity and fertility of female gametophytes. Females left alone or in the presence of sibling males express the highest gametophyte size, number, and size of oogonia, suggesting they were able to sense the presence and the identity of their mates before gamete contact. The opposite trend was observed for the production of embryos per female gametes, indicating higher costs of selfing and parthenogenesis than outcrossing on fertility. However, the increased fecundity compensated for the reduced fertility, leading to a stable overall reproductive output. Inbreeding also affected morphological traits of juvenile sporophytes, but not their heatwave tolerance. The male–female kinship effect was stronger in high‐latitude populations, suggesting that females from low‐latitude marginal populations might have evolved to mate with any male gamete to guarantee reproductive success.
Rocky reefs and kelp forests form conspicuous habitats that promote species diversity and support livelihoods in coastal communities. However, taxonomic approaches often disregard biological identity ...and differences between species. In this study, we explore the relationship between functional diversity of fish communities and habitat complexity in temperate reefs and test if greater habitat complexity, such as rugosity or kelp three-dimensional structure, would result in higher functional diversity. We conducted fish surveys using SCUBA in four kelp forest sites and rocky reef sites. Although the rocky reef and kelp forest sites showed differences in habitat complexity, no significant differences in fish taxonomic or functional diversity were found between the two habitat types. However, we did find differences at smaller spatial scales for the kelp forest sites, where number of stipes, stipe bundle diameter, and kelp density influenced functional richness, but not species richness, highlighting the importance of functional approaches in certain ecosystems. The differences found among kelp forest sites may be linked with small-scale spatiotemporal oceanographic drivers of productivity such as upwelling exposure or nutrient availability. We recommend considering small-scale spatial drivers when aiming to understand how habitat characteristics link with functional diversity.
At small spatial and temporal scales, genetic differentiation is largely controlled by constraints on gene flow, while genetic diversity across a species' distribution is shaped on longer temporal ...and spatial scales. We assess the hypothesis that oceanographic transport and other seascape features explain different scales of genetic structure of giant kelp, Macrocystis pyrifera. We followed a hierarchical approach to perform a microsatellite‐based analysis of genetic differentiation in Macrocystis across its distribution in the northeast Pacific. We used seascape genetic approaches to identify large‐scale biogeographic population clusters and investigate whether they could be explained by oceanographic transport and other environmental drivers. We then modelled population genetic differentiation within clusters as a function of oceanographic transport and other environmental factors. Five geographic clusters were identified: Alaska/Canada, central California, continental Santa Barbara, California Channel Islands and mainland southern California/Baja California peninsula. The strongest break occurred between central and southern California, with mainland Santa Barbara sites forming a transition zone between the two. Breaks between clusters corresponded approximately to previously identified biogeographic breaks, but were not solely explained by oceanographic transport. An isolation‐by‐environment (IBE) pattern was observed where the northern and southern Channel Islands clustered together, but not with closer mainland sites, despite the greater distance between them. The strongest environmental association with this IBE pattern was observed with light extinction coefficient, which extends suitable habitat to deeper areas. Within clusters, we found support for previous results showing that oceanographic connectivity plays an important role in the population genetic structure of Macrocystis in the Northern hemisphere.
The frequency of marine heatwaves (MHWs) is increasing due to climate change. Although seaweeds are resilient to environmental changes, an increasing body of evidence shows that rising sea surface ...temperatures have deleterious effects on temperate kelp species. However, information on the vulnerability of juvenile kelp to these stressors and their population stability is limited. This study summarizes findings on the ability of juvenile sporophytes of Macrocystis pyrifera to survive and recover from simulated MHW conditions (22°C, 5 d) in combination with nitrate limitation (<1 µM) by evaluating photosynthetic capacity, nitrate uptake, tissue composition, bio-optical properties, and oxidative stress of single-blade juvenile sporophytes (<20 cm). Temperature, nitrate availability, and their interaction had significant effects on the physiological status of juvenile sporophytes after the exposure and recovery periods. Overall, as expected, the photosynthetic capacity of juvenile sporophytes decreased with increased temperature and lower nitrate availability. Short-term exposure to simulated MHWs resulted in oxidative damage and reduced growth. The termination of the experimental warming allowed partial recovery to control values, indicating high physiological resilience. However, the interaction of both high temperature and nitrate scarcity induced irreversible damage to their photosynthetic capacity, with an increase in compensation irradiance, highlighting potential limitations in the carbon balance of juvenile sporophytes.
The paradigm of past climate-driven range shifts structuring the distribution of marine intraspecific biodiversity lacks replication in biological models exposed to comparable limiting conditions in ...independent regions. This may lead to confounding effects unlinked to climate drivers. We aim to fill in this gap by asking whether the global distribution of intraspecific biodiversity of giant kelp (Macrocystis pyrifera) is explained by past climate changes occurring across the two hemispheres. We compared the species' population genetic diversity and structure inferred with microsatellite markers, with range shifts and long-term refugial regions predicted with species distribution modelling (SDM) from the last glacial maximum (LGM) to the present. The broad antitropical distribution of Macrocystis pyrifera is composed by six significantly differentiated genetic groups, for which current genetic diversity levels match the expectations of past climate changes. Range shifts from the LGM to the present structured low latitude refugial regions where genetic relics with higher and unique diversity were found (particularly in the Channel Islands of California and in Peru), while post-glacial expansions following ~ 40% range contraction explained extensive regions with homogenous reduced diversity. The estimated effect of past climate-driven range shifts was comparable between hemispheres, largely demonstrating that the distribution of intraspecific marine biodiversity can be structured by comparable evolutionary forces across the global ocean. Additionally, the differentiation and endemicity of regional genetic groups, confers high conservation value to these localized intraspecific biodiversity hotspots of giant kelp forests.
Kelp beds provide significant ecosystem services and socioeconomic benefits globally, and prominently in coastal zones of the California Current. Their distributions and abundance, however, vary ...greatly over space and time. Here, we describe long-term patterns of Giant Kelp (
Macrocystis pyrifera
) sea surface canopy area off the coast of San Diego County from 1983 through 2019 along with recent patterns of water column nitrate (NO
3
-
) exposure inferred from
in situ
temperature data in 2014 and 2015 at sites spanning 30 km of the coastline near San Diego California, USA. Site-specific patterns of kelp persistence and resilience were associated with ocean and climate dynamics, with total sea surface kelp canopy area varying approximately 33-fold over the almost 4 decades (min 0.34 km
2
in 1984; max 11.25 km
2
in 2008, median 4.79 km
2
). Site-normalized canopy areas showed that recent kelp persistence since 2014 was greater at Point Loma and La Jolla, the largest kelp beds off California, than at the much smaller kelp bed off Cardiff. NO
3
-
exposure was estimated from an 11-month time series of
in situ
water column temperature collected in 2014 and 2015 at 4 kelp beds, using a relationship between temperature and NO
3
-
concentration previously established for the region. The vertical position of the 14.5°C isotherm, an indicator of the main thermocline and nutricline, varied across the entire water column at semidiurnal to seasonal frequencies. We use a novel means of quantifying estimated water column NO
3
-
exposure integrated through time (mol-days m
-2
) adapted from degree days approaches commonly used to characterize thermal exposures. Water column integrated NO
3
-
exposure binned by quarters of the time series showed strong seasonal differences with highest exposure in Mar - May 2015, lowest exposure in Sep - Dec 2014, with consistently highest exposure off Point Loma. The water column integrated NO
3
-
signal was filtered to provide estimates of the contribution to total nitrate exposure from high frequency variability (ƒ >= 1 cycle 30 hr
-1
) associated predominantly with internal waves, and low frequency variability driven predominantly by seasonal upwelling. While seasonal upwelling accounted for > 90% of NO
3
-
exposure across the full year, during warm periods when seasonal upwelling was reduced or absent and NO
3
-
exposure was low overall, the proportion due to internal waves increased markedly to 84 to 100% of the site-specific total exposure. The high frequency variability associated with internal waves may supply critical nutrient availability during anomalously warm periods. Overall, these analyses support a hypothesis that differences in NO
3
-
exposure among sites due to seasonal upwelling and higher frequency internal wave forcing contribute to spatial patterns in Giant Kelp persistence in southern California. The study period includes anomalously warm surface conditions and the marine heatwave associated with the “Pacific Warm Blob” superimposed on the seasonal thermal signal and corresponding to the onset of a multi-year decline in kelp canopy area and marked differences in kelp persistence among sites. Our analysis suggests that, particularly during periods of warm surface conditions, variation in NO
3
-
exposure associated with processes occurring at higher frequencies, including internal waves can be a significant source of NO
3
-
exposure to kelp beds in this region. The patterns described here also offer a view of the potential roles of seasonal and higher frequency nutrient dynamics for Giant Kelp persistence in southern California under continuing ocean surface warming and increasing frequency and intensity of marine heatwaves.
In the Galápagos Islands, two eulimid snails parasitize the common pencil sea urchin, Eucidaris galapagensis. Past work in the Galápagos suggests that fishing reduces lobster and fish densities and, ...due to this relaxation of predation pressure, indirectly increases urchin densities, creating the potential for complex indirect interactions between fishing and parasitic snails. To measure indirect effects of fishing on these parasitic snails, we investigated the spatial relationships among urchins, parasitic snails, commensal crabs, and large urchin predators (hogfish and lobsters). Parasitic snails had higher densities at sites where urchins were abundant, probably due to increased resource availability. Commensal crabs that shelter under urchin spines, particularly the endemic Mithrax nodosus, preyed on the parasitic snails in aquaria, and snails were less abundant at field sites where these crabs were common. In aquaria, hogfish and lobsters readily ate crabs, but crabs were protected from predation under urchin spines, leading to a facultative mutualism between commensal crabs and urchins. In the field, fishing appeared to indirectly increase the abundance of urchins and their commensal crabs by reducing predation pressure from fish and lobsters. Fished sites had fewer snails per urchin, probably due to increased predation from commensal crabs. However, because fished sites also tended to have more urchins, there was no significant net effect of fishing on the number of snails per square meter. These results suggest that fishing can have complex indirect effects on parasites by altering food webs.
The frequency of marine heatwaves (MHWs) is increasing due to climate change. Although seaweeds are resilient to environmental changes, an increasing body of evidence shows that rising sea surface ...temperatures have deleterious effects on temperate kelp species. However, information on the vulnerability of juvenile kelp to these stressors and their population stability is limited. This study summarizes findings on the ability of juvenile sporophytes of Macrocystis pyrifera to survive and recover from simulated MHW conditions (22°C, 5 d) in combination with nitrate limitation (<1 µM) by evaluating photosynthetic capacity, nitrate uptake, tissue composition, bio‐optical properties, and oxidative stress of single‐blade juvenile sporophytes (<20 cm). Temperature, nitrate availability, and their interaction had significant effects on the physiological status of juvenile sporophytes after the exposure and recovery periods. Overall, as expected, the photosynthetic capacity of juvenile sporophytes decreased with increased temperature and lower nitrate availability. Short‐term exposure to simulated MHWs resulted in oxidative damage and reduced growth. The termination of the experimental warming allowed partial recovery to control values, indicating high physiological resilience. However, the interaction of both high temperature and nitrate scarcity induced irreversible damage to their photosynthetic capacity, with an increase in compensation irradiance, highlighting potential limitations in the carbon balance of juvenile sporophytes.
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