Mesoscale fronts as foraging habitats: composite front mapping reveals oceanographic drivers of habitat use for a pelagic seabird

The oceanographic drivers of marine vertebrate habitat use are poorly understood yet fundamental to our knowledge of marine ecosystem functioning. Here, we use composite front mapping and high-resolution GPS tracking to determine the significance of mesoscale oceanographic fronts as physical drivers of foraging habitat selection in northern gannets Morus bassanus. We tracked 66 breeding gannets from a Celtic Sea colony over 2 years and used residence time to identify area-restricted search (ARS) behaviour. Composite front maps identified thermal and chlorophyll-a mesoscale fronts at two different temporal scales—(i) contemporaneous fronts and (ii) seasonally persistent frontal zones. Using generalized additive models (GAMs), with generalized estimating equations (GEE-GAMs) to account for serial autocorrelation in tracking data, we found that gannets do not adjust their behaviour in response to contemporaneous fronts. However, ARS was more likely to occur within spatially predictable, seasonally persistent frontal zones (GAMs). Our results provide proof of concept that composite front mapping is a useful tool for studying the influence of oceanographic features on animal movements. Moreover, we highlight that frontal persistence is a crucial element of the formation of pelagic foraging hotspots for mobile marine vertebrates.

Predicting future thermal habitat suitability of competing native and invasive fish species: from metabolic scope to oceanographic modelling

Global increase in sea temperatures has been suggested to facilitate the incoming and spread of tropical invaders. The increasing success of these species may be related to their higher physiological performance compared with indigenous ones. Here, we determined the effect of temperature on the aerobic metabolic scope (MS) of two herbivorous fish species that occupy a similar ecological niche in the Mediterranean Sea: the native salema (Sarpa salpa) and the invasive marbled spinefoot (Siganus rivulatus). Our results demonstrate a large difference in the optimal temperature for aerobic scope between the salema (21.8°C) and the marbled spinefoot (29.1°C), highlighting the importance of temperature in determining the energy availability and, potentially, the distribution patterns of the two species. A modelling approach based on a present-day projection and a future scenario for oceanographic conditions was used to make predictions about the thermal habitat suitability (THS, an index based on the relationship between MS and temperature) of the two species, both at the basin level (the whole Mediterranean Sea) and at the regional level (the Sicilian Channel, a key area for the inflow of invasive species from the Eastern to the Western Mediterranean Sea). For the present-day projection, our basin-scale model shows higher THS of the marbled spinefoot than the salema in the Eastern compared with the Western Mediterranean Sea. However, by 2050, the THS of the marbled spinefoot is predicted to increase throughout the whole Mediterranean Sea, causing its westward expansion. Nevertheless, the regional-scale model suggests that the future thermal conditions of Western Sicily will remain relatively unsuitable for the invasive species and could act as a barrier for its spread westward. We suggest that metabolic scope can be used as a tool to evaluate the potential invasiveness of alien species and the resilience to global warming of native species.

Effect of habitat fragmentation on the macroinvertebrate infaunal communities associated with the seagrass Zostera marina

1. The effect of habitat fragmentation was investigated in two adjacent, yet separate, intertidal Zostera marina beds in the Salcombe Estuary, Devon, UK. The seagrass bed on the west bank comprised a continuous meadow of ca. 2.3 ha, whilst the bed on the east bank of the estuary was fragmented into patches of 6–9 m2.2. Three 10 cm diameter core samples for infaunal macroinvertebrates were taken from three stations within each bed. No significant difference was found in univariate community parameters between beds, or in measured seagrass parameters. However, multivariate analysis revealed a significant difference in community composition, due mainly to small changes in species abundance rather than differences in the species present.3. The species contributing most to the dissimilarity between the two communities were polychaetes generally associated with unvegetated habitats (e.g. Magelona mirabilis) and found to be more common in the fragmented bed.4. A significant difference in median grain size and sorting coefficient was recorded between the two beds, and median grain size was found to be the variable best explaining multivariate community patterns.5. The results of the study provide evidence for the effects of habitat fragmentation on the communities associated with seagrass beds, habitats which are of high conservation importance. As the infaunal community is perhaps intuitively the component least likely to be affected by fragmentation at the scale observed, the significant difference in community composition recorded has consequences for more sensitive and high-profile parts of the biota (e.g. fish), and thus for the conservation of seagrass habitats and their associated communities.

Lamellibrachia anaximandrin. sp., a new vestimentiferan tubeworm (Annelida) from the Mediterranean, with notes on frenulate tubeworms from the same habitat

A new species of lamellibrachiid vestimentiferan, Lamellibrachia anaximandri n. sp., has been found in the Eastern Mediterranean, close to cold seeps of fluid carrying dissolved methane and sources of sulfide in superficial sediments. It occurs at about 1100 to 2100 m depth, on some of the mud volcanoes on the Anaximander Mountains, south of Turkey, on the Mediterranean Ridge, south of Crete, and on the Nile deep-sea fan. In addition, it has been obtained from rotting paper inside a sunken ship, torpedoed in 1915 and lying at 2800 m depth, southeast of Crete. Some frenulate pogonophores also occur on the mud volcanoes (including a species of Siboglinum resembling S. carpinei and tubes of other unidentified genera). The new Lamellibrachia is the first vestimentiferan species to be described from the Mediterranean. It differs from L. luymesi taken from the Gulf of Mexico population in the very weak development of collars on its tube and in having a smaller number of pairs of branchial lamellae in the branchial plume. Sequencing of the COI and the mt16S genes confirms a difference at the species level between the new species and L. luymesi, and a difference between these two species and four described species of Lamellibrachia from the Pacific Ocean. The largest individuals of L. anaximandri n. sp. may be many years old, but there are numerous young individuals at some sites, showing that favourable conditions are available for settlement and early growth. The development of the branchial plume in a series of young stages reveals that the sheath lamellae, which are characteristic of the genus Lamellibrachia, begin to form only after the establishment of several pairs of branchial lamellae. Examination of the adult trophosome by transmission electron microscopy shows Gram-negative bacteria without internal stacked membranes, indicating that the symbionts are most probably sulfide oxidizing.

Effect of habitat fragmentation on levels and patterns of genetic diversity in natural populations of the peat moss Polytrichum commune

Peat bogs represent unique ecosystems that are under particular threat from fragmentation due to peat harvesting, with only 38% of the original peatland in Europe remaining intact and unaffected by peat cutting, drainage and silviculture. In this study, we have used microsatellite markers to determine levels and patterns of genetic diversity in both cut and uncut natural populations of the peat moss Polytrichum commune. Overall diversity levels suggest that there is more genetic variation present than had previously been assumed for bryophytes. Despite this, diversity values from completely cut bogs were found to be lower than those from uncut peatlands (average 0.729 versus 0.880). In addition, the genetic diversity was more highly structured in the cut populations, further suggesting that genetic drift is already affecting genetic diversity in peat bogs subjected to fragmentation.

A species invasion mediated through habitat structure, intraguild predation and parasitism.

With field, laboratory, and modeling approaches, we examined the interplay among habitat structure, intraguild predation (IGP), and parasitism in an ongoing species invasion. Native Gammarus duebeni celticus (Crustacea: Amphipoda) are often, but not always, replaced by the invader Gammarus pulex through differential IGP. The muscle-wasting microsporidian parasite Pleistophora mulleri infects the native but not the invader. We found a highly variable prevalence of P. mulleri in uninvaded rivers, with 0–91% of hosts parasitized per sample. In addition, unparasitized natives dominated fast-flowing riffle patches of river, whereas parasitized individuals dominated slower- flowing, pooled patches. We examined the survivorship of invader and native in single and mixed-species microcosms with high, intermediate, and zero parasite prevalence. G. pulex survivorship was high in all treatments, whereas G. duebeni subsp. celticus survivorship was significantly lower in the presence of the invader. Further, parasitized G. duebeni subsp. celticus experienced near-total elimination. Models of the species replacement process implied that parasite-enhanced IGP would make invasion by G. pulex more likely, regardless of habitat and parasite spatial structure. However, where heterogeneity in parasite prevalence creates a landscape of patches with different susceptibilities to invasion, G. pulex may succeed in cases where invasion would not be possible if patches were equivalent. The different responses of parasitized and unparasitized G. duebeni subsp. celticus to environmental heterogeneity potentially link landscape patterns to the success or failure of the invasion process.

The area-independent effects of habitat complexity on biodiversity vary between regions.

Potential explanatory variables often co-vary in studies of species richness. Where topography varies within a survey it is difficult to separate area and habitat-diversity effects. Topographically complex surfaces may contain more species due to increased habitat diversity or as a result of increased area per se. Fractal geometry can be used to adjust species richness estimates to control for increases in area on complex surfaces. Application of fractal techniques to a survey of rocky shores demonstrated an unambiguous area-independent effect of topography on species richness in the Isle of Man. In contrast, variation in species richness in south-west England reflected surface availability alone. Multivariate tests and variation in limpet abundances also demonstrated regional variation in the area-independent effects of topography. Community composition did not vary with increasing surface complexity in south-west England. These results suggest large-scale gradients in the effects of heterogeneity on community processes or demography.

Challenging assumptions of future coastal habitat development around the UK

One habitat management requirement forced by 21st century relative sea-level rise (RSLR), will be the need to re-comprehend the dimensions of long-term transgressive behaviour of coastal systems being forced by such RSLR. Fresh approaches to the conceptual modelling and subsequent implementation of new coastal and peri-marine habitats will be required. There is concern that existing approaches to forecasting coastal systems development (and by implication their associated scarce coastal habitats) over the next century depend on a certain premise of orderly spatial succession of habitats. This assumption is shown to be questionable given the possible future rates of RSLR, magnitude of shoreline retreat and the lack of coastal sediment to maintain the protective morphologies to low-energy coastal habitats. Of these issues, sediment deficiency is regarded as one of the major problem for future habitat development. Examples of contemporary behaviour of UK coasts show evidence of coastal sediment starvation resulting from relatively stable RSLR, anthropogenic sealing of coastal sources, and intercepted coastal sediment pathways, which together force segmentation of coastal systems. From these examples key principles are deduced which may prejudice the existence of future habitats: accelerated future sediment demand due to RSLR may not be met by supply and, if short- to medium-term hold-the-line policies predominate, long-term strategies for managed realignment and habitat enhancement may prove impossible goals. Methods of contemporary sediment husbandry may help sustain some habitats in place but otherwise, instead of integrated coastal organization, managers may need to consider coastal breakdown, segmentation and habitat reduction as the basis of 21st century coastal evolution and planning.

Habitat-specific adaptation of immune responses of stickleback (Gasterosteus aculeatus) lake and river ecotypes

Freshwater populations of three-spined sticklebacks (Gasterosteus aculeatus) in northern Germany are found as distinct lake and river ecotypes. Adaptation to habitat-specific parasites might influence immune capabilities of stickleback ecotypes. Here, naive laboratory-bred sticklebacks from lake and river populations were exposed reciprocally to parasite environments in a lake and a river habitat. Sticklebacks exposed to lake conditions were infected with higher numbers of parasite species when compared with the river. River sticklebacks in the lake had higher parasite loads than lake sticklebacks in the same habitat. Respiratory burst, granulocyte counts and lymphocyte proliferation of head kidney leucocytes were increased in river sticklebacks exposed to lake when compared with river conditions. Although river sticklebacks exposed to lake conditions showed elevated activation of their immune system, parasites could not be diminished as effectively as by lake sticklebacks in their native habitat. River sticklebacks seem to have reduced their immune-competence potential due to lower parasite diversity in rivers

Pollinators of an endemic and endangered species, Mammillaria gaumeri (Cactaceae), in its natural habitat (coastal dune) and in a botanical garden.

Mammillaria gaumeri (Britton & Ross) Orcutt (Cactaceae), an endemic plant of the Yucatan Peninsula, is included by the Mexican government in the list of species that require special protection. Its natural habitat is now restricted to fragmented areas and protection programs involve botanical gardens in growing individuals rescued from disturbed areas. Little information is available on the reproductive characteristics of this species and nothing is known of its pollinators. We investigated the visitors of M. gaumeri flowers, collecting and observing bee species in its natural habitat (i.e., coastal dune) and in a botanical garden, where coastal dune vegetation had been created. Observations were made on plants whose density was artificially increased by grouping flowering individuals. At each site, we: 1) collected insects visiting the flowers; 2) recorded number of visits; and 3) video-recorded bee movements on the flowers. As expected, the number of bee species and visitation frequency were higher at the botanical garden than at the coastal dune. After landing on a flower, bees either inspected the anthers or dived among them. These behaviors, carried out by all observed species, seemed related to the state of the anthers (full or empty of pollen) and stigma lobes (opened or closed). Specifically, visits lasted longer when anthers were full of pollen and stigma lobes were opened. The same bee species recorded on the dune were also recorded at the botanical garden, suggesting that the artificial dune at the botanical garden offered suitable conditions for the natural pollinators of this endangered cactus.