Persistence and transport of fauna on drifting kelp (Macrocystis pyrifera (L.) C. Agardh) rafts in the Southern California Bight

Drifting rafts of Macrocystis pyrifera may connect isolated kelp forests in the Southern California Bight. To determine which species might utilize this dispersal mechanism, faunal samples from natural drifting rafts and attached M. pyrifera plants were collected during five cruises between March 1995 and December 1997. These rafts, which can be considered as floating islands, were aged and the macroinvertebrate assemblage enumerated. There was no significant relationship between raft age and species richness, or between species richness and distance offshore, which contrasts with predictions based on island biogeography. Species richness, however, was related to raft weight. Patterns of species presence and density were investigated relative to raft age for the species most frequently associated with rafts. Only one species, the isopod Idotea resecata, was found on all sampled rafts. Some species increased in frequency with raft age and others decreased, but only one relationship, a decline in the frequency of the anemone Epiactis prolifera with raft age was significant. When species density was examined over all cruises, only I. resecata had a significant change in density (an increase) with raft age, but additional significant relationships were found when species density patterns were considered by cruise. The results of all the tests were combined to provide a measure of "raft success". Nine of the most frequent 19 species had a positive score, indicating a favorable response to rafting, seven were unaffected, and two species had negative responses to rafting. Extinction times were calculated using species abundance and raft age relationships. Two species (E. prolifera and Paracerceis cordata), were predicted to persist on rafts for only about 100 days, which is the maximum estimated raft lifetime. All other species were predicted to persist for longer periods if the rafts floated longer. Kelp fauna that begin rafting appear to be largely unaffected by rafting, and hence dispersal on kelp rafts is possible for many members of the kelp forest community.     

Rafting of benthic macrofauna: important factors determining the temporal succession of the assemblage on detached macroalgae

Rafting on biotic and abiotic substrata has been reported for many benthic marine invertebrates. Here, I describe important characteristics of common floating substrata and review published studies examining the succession of the assemblage on detached macroalgae to identify the most important factors determining this succession. Floating substrata differ in survival time (i.e. before they disintegrate) and in food value, with abiotic materials having high survival times but low food value and some biotic substrata (seagrasses, fresh wood) having high food values but short survival times. Large macroalgae with pneumatocysts may combine high survival times and high food values. Substratum survival and food value have consequences for the species composition of the rafting macrofauna. In general, suspension feeders dominate rafting assemblages on abiotic substrata, while grazing and boring species abound on macroalgae and wood. The succession of the rafting assemblage can be subdivided into three phases during which different processes predominate. During the initial phase some of the original colonizers (if present) disappear from the substratum, either due to active emigration or predation. This short, initial phase is followed immediately by the colonization phase, during which new organisms colonize the rafting substratum. Colonization may be rapid and intense in the beginning but then slows down. Towards the end of the colonization phase, some rafting organisms may reproduce and their offspring recruit within the parental raft. Results from two long-term studies confirmed that the proportion of species with direct development increased with duration of rafting. These successional changes will be most pronounced for large biotic substrata, rendering these particularly suitable for long-distance dispersal of organisms with direct development.     

Monitoring the abundance of plastic debris in the marine environment

Plastic debris has significant environmental and economic impacts in marine systems. Monitoring is crucial to assess the efficacy of measures implemented to reduce the abundance of plastic debris, but it is complicated by large spatial and temporal heterogeneity in the amounts of plastic debris and by our limited understanding of the pathways followed by plastic debris and its long-term fate. To date, most monitoring has focused on beach surveys of stranded plastics and other litter. Infrequent surveys of the standing stock of litter on beaches provide crude estimates of debris types and abundance, but are biased by differential removal of litter items by beachcombing, cleanups and beach dynamics. Monitoring the accumulation of stranded debris provides an index of debris trends in adjacent waters, but is costly to undertake. At-sea sampling requires large sample sizes for statistical power to detect changes in abundance, given the high spatial and temporal heterogeneity. Another approach is to monitor the impacts of plastics. Seabirds and other marine organisms that accumulate plastics in their stomachs offer a cost-effective way to monitor the abundance and composition of small plastic litter. Changes in entanglement rates are harder to interpret, as they are sensitive to changes in population sizes of affected species. Monitoring waste disposal on ships and plastic debris levels in rivers and storm-water runoff is useful because it identifies the main sources of plastic debris entering the sea and can direct mitigation efforts. Different monitoring approaches are required to answer different questions, but attempts should be made to standardize approaches internationally.     

Transoceanic dispersal of terrestrial species by debris rafting

Rare, long-distance dispersal events are a key process in generating and maintaining patterns in biological diversity and species distributions across space and time. The 9.0 magnitude earthquake that struck the eastern coast of Japan in 2011, and the subsequent 38 m high tsunami washed large amounts of shoreline debris into the Pacific Ocean that led to a large-scale biological rafting event carrying nearly 300 marine species to the western shores of North America. Whether oceanic, trans-Pacific dispersal via rafting generates long distance dispersal events for small, flightless, terrestrial species is unknown. By sampling beach debris associated with known hot-spots of tsunami debris along the north and east shores of Graham Island, Haida Gwaii, Canada, I document significantly dissimilar invertebrate communities associated with tide-line beach debris and the occurrence of several putative Japanese species of soil-dwelling mites (Acari: Oribatida). Previous explanations of Haida Gwaii's unique flora and fauna have been attributed to a proximity to the Beringian land bridge and the accumulated evidence of near-offshore glacial refugia during the last glacial period. However, my research also suggests that stochastic, trans-Pacific rafting events contribute to the biodiversity and biogeography of soil communities on the west coast of North America.     

Rapid, long-distance dispersal by pumice rafting

Pumice is an extremely effective rafting agent that can dramatically increase the dispersal range of a variety of marine organisms and connect isolated shallow marine and coastal ecosystems. Here we report on a significant recent pumice rafting and long-distance dispersal event that occurred across the southwest Pacific following the 2006 explosive eruption of Home Reef Volcano in Tonga. We have constrained the trajectory, and rate, biomass and biodiversity of transfer, discovering more than 80 species and a substantial biomass underwent a >5000 km journey in 7-8 months. Differing microenvironmental conditions on the pumice, caused by relative stability of clasts at the sea surface, promoted diversity in biotic recruitment. Our findings emphasise pumice rafting as an important process facilitating the distribution of marine life, which have implications for colonisation processes and success, the management of sensitive marine environments, and invasive pest species.     

Risk analysis reveals global hotspots for marine debris ingestion by sea turtles

Plastic marine debris pollution is rapidly becoming one of the critical environmental concerns facing wildlife in the 21st century. Here we present a risk analysis for plastic ingestion by sea turtles on a global scale. We combined global marine plastic distributions based on ocean drifter data with sea turtle habitat maps to predict exposure levels to plastic pollution. Empirical data from necropsies of deceased animals were then utilised to assess the consequence of exposure to plastics. We modelled the risk (probability of debris ingestion) by incorporating exposure to debris and consequence of exposure, and included life history stage, species of sea turtle and date of stranding observation as possible additional explanatory factors. Life history stage is the best predictor of debris ingestion, but the best‐fit model also incorporates encounter rates within a limited distance from stranding location, marine debris predictions specific to the date of the stranding study and turtle species. There is no difference in ingestion rates between stranded turtles vs. those caught as bycatch from fishing activity, suggesting that stranded animals are not a biased representation of debris ingestion rates in the background population. Oceanic life‐stage sea turtles are at the highest risk of debris ingestion, and olive ridley turtles are the most at‐risk species. The regions of highest risk to global sea turtle populations are off of the east coasts of the USA, Australia and South Africa; the east Indian Ocean, and Southeast Asia. Model results can be used to predict the number of sea turtles globally at risk of debris ingestion. Based on currently available data, initial calculations indicate that up to 52% of sea turtles may have ingested debris.     

Plastic Pollution in the World's Oceans: More than 5 Trillion Plastic Pieces Weighing over 250,000 Tons Afloat at Sea

Plastic pollution is ubiquitous throughout the marine environment, yet estimates of the global abundance and weight of floating plastics have lacked data, particularly from the Southern Hemisphere and remote regions. Here we report an estimate of the total number of plastic particles and their weight floating in the world''s oceans from 24 expeditions (2007–2013) across all five sub-tropical gyres, costal Australia, Bay of Bengal and the Mediterranean Sea conducting surface net tows (N = 680) and visual survey transects of large plastic debris (N = 891). Using an oceanographic model of floating debris dispersal calibrated by our data, and correcting for wind-driven vertical mixing, we estimate a minimum of 5.25 trillion particles weighing 268,940 tons. When comparing between four size classes, two microplastic <4.75 mm and meso- and macroplastic >4.75 mm, a tremendous loss of microplastics is observed from the sea surface compared to expected rates of fragmentation, suggesting there are mechanisms at play that remove <4.75 mm plastic particles from the ocean surface.     

Dispersal in Marine Organisms without a Pelagic Larval Phase

In contrast to marine organisms whose offspring go through an extended planktonic stage, the young of others develop directly into benthic juveniles or into yolky nonfeeding larvae that spend only a few hours in the plankton before settling. Yet, paradoxically, many such species have geographic distributions that are comparable to those with a pelagic dispersal stage. This article reviews some of the ways in which these organisms can expand their distributions: drifting, rafting, hitchhiking, creeping, and hopping. Drifting applies to species in which larvae may be short-lived, but adults can detach or be detached from their benthic substratum and be passively carried to new areas, floating at the water's surface or below it. Many encrusting species and mobile species can spread by rafting, settling on natural or artificial floating substrata which are propelled by wind and currents to new regions. Hitchhiking applies to those attaching to vessels or being carried in ballast water of ships to a distant region in which their offspring can survive. Other marine species extend their distributions by hopping from one island of hard substratum or favorable sedimentary microhabitat to another, while creeping species extend their distributions along shores or shelves where habitats remain similar for long distances.     

To Eat or Not to Eat? Debris Selectivity by Marine Turtles

Marine debris is a growing problem for wildlife, and has been documented to affect more than 267 species worldwide. We investigated the prevalence of marine debris ingestion in 115 sea turtles stranded in Queensland between 2006-2011, and assessed how the ingestion rates differ between species (Eretmochelys imbricata vs. Chelonia mydas) and by turtle size class (smaller oceanic feeders vs. larger benthic feeders). Concurrently, we conducted 25 beach surveys to estimate the composition of the debris present in the marine environment. Based on this proxy measurement of debris availability, we modeled turtles' debris preferences (color and type) using a resource selection function, a method traditionally used for habitat and food selection. We found no significant difference in the overall probability of ingesting debris between the two species studied, both of which have similar life histories. Curved carapace length, however, was inversely correlated with the probability of ingesting debris; 54.5% of pelagic sized turtles had ingested debris, whereas only 25% of benthic feeding turtles were found with debris in their gastrointestinal system. Benthic and pelagic sized turtles also exhibited different selectivity ratios for debris ingestion. Benthic phase turtles had a strong selectivity for soft, clear plastic, lending support to the hypothesis that sea turtles ingest debris because it resembles natural prey items such as jellyfish. Pelagic turtles were much less selective in their feeding, though they showed a trend towards selectivity for rubber items such as balloons. Most ingested items were plastic and were positively buoyant. This study highlights the need to address increasing amounts of plastic in the marine environment, and provides evidence for the disproportionate ingestion of balloons by marine turtles.     

Plastic Accumulation in the Mediterranean Sea

Concentrations of floating plastic were measured throughout the Mediterranean Sea to assess whether this basin can be regarded as a great accumulation region of plastic debris. We found that the average density of plastic (1 item per 4 m²), as well as its frequency of occurrence (100% of the sites sampled), are comparable to the accumulation zones described for the five subtropical ocean gyres. Plastic debris in the Mediterranean surface waters was dominated by millimeter-sized fragments, but showed a higher proportion of large plastic objects than that present in oceanic gyres, reflecting the closer connection with pollution sources. The accumulation of floating plastic in the Mediterranean Sea (between 1,000 and 3,000 tons) is likely related to the high human pressure together with the hydrodynamics of this semi-enclosed basin, with outflow mainly occurring through a deep water layer. Given the biological richness and concentration of economic activities in the Mediterranean Sea, the affects of plastic pollution on marine and human life are expected to be particularly frequent in this plastic accumulation region.     

Long-distance dispersal, low connectivity and molecular evidence of a new cryptic species in the obligate rafter Caprella andreae Mayer, 1890 (Crustacea: Amphipoda: Caprellidae)

The amphipod Caprella andreae Mayer, 1890 was recorded for the first time in Southern Iberian Peninsula (36°44′15″N, 3°59′38″W). This species is the only obligate rafter of the suborder Caprellidea and has been reported to attach not only to floating objects such as ropes or driftwoods but also to turtle carapaces. Mitochondrial and nuclear markers were used to examine dispersal capabilities and population genetic structure of C. andreae across seven localities in the Mediterranean and Atlantic Ocean collected from floating substrata with different dispersal patterns. The strong population differentiation with no haplotypes shared between populations suggests that C. andreae is quite faithful to the substratum on which it settles. In addition, the proportionally higher genetic diversity displayed in populations living on turtles as well as the presence of highly differentiated haplotypes in the same turtle population may be indicative that these populations survive longer, which could lead C. andreae to prefer turtles instead of floating objects to settle and disperse. Therefore, rafting on floating objects may be sporadic, and ocean currents would not be the most important factor shaping patterns of connectivity and population structure in this species. Furthermore, molecular phylogenetic analyses revealed the existence of a cryptic species whose estimates of genetic divergence are higher than those estimated between C. andreae and other congeneric species (e.g. Caprella dilatata and Caprella penantis). Discovery of cryptic species among widely distributed small marine invertebrates is quite common and, in this case, prompts for a more detailed phylogenetic analysis and taxonomic revision of genus Caprella. On the other hand, this study also means the first record of the gammarids Jassa cadetta and Elasmopus brasiliensis and the caprellid Caprella hirsuta on drifting objects.     

DECOUPLING OF SHORT‐ AND LONG‐DISTANCE DISPERSAL PATHWAYS IN THE ENDEMIC NEW ZEALAND SEAWEED CARPOPHYLLUM MASCHALOCARPUM (PHAEOPHYCEAE,FUCALES)1

The processes that produce and maintain genetic structure in organisms operate at different timescales and on different life‐history stages. In marine macroalgae, gene flow occurs through gamete/zygote dispersal and rafting by adult thalli. Population genetic patterns arise from this contemporary gene flow interacting with historical processes. We analyzed spatial patterns of mitochondrial DNA variation to investigate contemporary and historical dispersal patterns in the New Zealand endemic fucalean brown alga Carpophyllum maschalocarpum (Turner) Grev. Populations bounded by habitat discontinuities were often strongly differentiated from adjoining populations over scales of tens of kilometers and intrapopulation diversity was generally low, except for one region of northeast New Zealand (the Bay of Plenty). There was evidence of strong connectivity between the northern and eastern regions of New Zealand’s North Island and between the North and South Islands of New Zealand and the Chatham Islands (separated by 650 km of open ocean). Moderate haplotypic diversity was found in Chatham Islands populations, while other southern populations showed low diversity consistent with Last Glacial Maximum (LGM) retreat and subsequent recolonization. We suggest that ocean current patterns and prevailing westerly winds facilitate long‐distance dispersal by floating adult thalli, decoupling genetic differentiation of Chatham Island populations from dispersal potential at the gamete/zygote stage. This study highlights the importance of encompassing the entire range of a species when inferring dispersal patterns from genetic differentiation, as realized dispersal distances can be contingent on local or regional oceanographic and historical processes.     

Nests of the brown booby (Sula leucogaster) as a potential indicator of tropical ocean pollution by marine debris

Seabirds collect debris primarily nearby breeding sites, and thus they may be used to monitor these pollutants in the ocean. This study aimed to investigate the prevalence of marine debris used as nesting materials by the brown booby (Sula leucogaster) and to test the species selectivity to debris type and color in two coastal islands of Brazil. We found marine debris in 61% of the brown booby nests on both islands. Fishing gear and hard plastic were the most frequent types of debris. Higher prevalence of fishing gear was found on the island with greater fishery activity. Similarly, hard plastic was the most frequent type of debris in nests and adjacent beach environment. The frequency of debris in brown booby nests can be a potential indicator of the abundance of specific items in surrounding marine waters. Monitoring debris in brown booby nests in a long-term may provide a better understanding of the species⿿ selectivity for specific debris. Furthermore, the impacts of debris in seabird nests at population level remain an overlooked threat that may reduce the quality of nesting habitats. We showed that brown booby nests are widely impacted by marine debris and that these organisms are exposed to this form of pollution from the beginning of their life.     

Oceanic rafting by a coastal community

Oceanic rafting is thought to play a fundamental role in assembling the biological communities of isolated coastal ecosystems. Direct observations of this key ecological and evolutionary process are, however, critically lacking. The importance of macroalgal rafting as a dispersal mechanism has remained uncertain, largely owing to lack of knowledge about the capacity of fauna to survive long voyages at sea and successfully make landfall and establish. Here, we directly document the rafting of a diverse assemblage of intertidal organisms across several hundred kilometres of open ocean, from the subantarctic to mainland New Zealand. Multispecies analyses using phylogeographic and ecological data indicate that 10 epifaunal invertebrate species rafted on six large bull kelp specimens for several weeks from the subantarctic Auckland and/or Snares Islands to the Otago coast of New Zealand, a minimum distance of some 400–600 km. These genetic data are the first to demonstrate that passive rafting can enable simultaneous trans-oceanic transport and landfall of numerous coastal taxa.     

Seed rafting as a dispersal strategy for eelgrass (Zostera marina)

Using observations, experiments and model simulations we investigate the possibility and range of long-distance dispersal by seed rafting in eelgrass, Zostera marina, at the Swedish west coast. A field experiment showed that the breaking strength of post-flowering rhipidia decreased significantly over a 30-day period, indicating that ontogenetic changes of the morphology facilitate dislodgement of the rhipidia at the optimal time for dispersal. We recorded positive buoyancy of detached post-flowering rhipidia for at least 26 days. Finally, we found evidence for sexual reproduction via seeds in the study area. These results, together with field measurements of wind-driven surface-transport velocity of rhipidia and wind data from 7 years, allowed us to model a dispersal potential of up to 150 km in one season along the Swedish west coast, which corresponds well with published estimates of dispersal potential from both observational and genetic studies. The results add to evidence that rafting of seeds on floating reproductive shoots is an important dispersal strategy for eelgrass, allowing for colonisation of new areas and for recolonisation after die-back events.     

Environmental implications of plastic debris in marine settings—entanglement,ingestion, smothering,hangers-on,hitch-hiking and alien invasions

Over the past five or six decades, contamination and pollution of the world’s enclosed seas, coastal waters and the wider open oceans by plastics and other synthetic, non-biodegradable materials (generally known as ‘marine debris’) has been an ever-increasing phenomenon. The sources of these polluting materials are both land- and marine-based, their origins may be local or distant, and the environmental consequences are many and varied. The more widely recognized problems are typically associated with entanglement, ingestion, suffocation and general debilitation, and are often related to stranding events and public perception. Among the less frequently recognized and recorded problems are global hazards to shipping, fisheries and other maritime activities. Today, there are rapidly developing research interests in the biota attracted to freely floating (i.e. pelagic) marine debris, commonly known as ‘hangers-on and hitch-hikers’ as well as material sinking to the sea floor despite being buoyant. Dispersal of aggressive alien and invasive species by these mechanisms leads one to reflect on the possibilities that ensuing invasions could endanger sensitive, or at-risk coastal environments (both marine and terrestrial) far from their native habitats.     

Fine scale endemism on coral reefs: archipelagic differentiation in turbinid gastropods

The perceived wide geographic range of organisms in the sea, facilitated by ready dispersal of waterborne dispersal stages, is a challenge for hypotheses of marine speciation but a boon to efforts of marine conservation. Wide species ranges are especially striking in the reef-rich Indo-west Pacific, the largest and most diverse marine biogeographic region, extending across half the planet. The insular marine biota of the tropical Pacific is characterized by wide-ranging species and provides the most striking examples of long distance dispersal, with endemism largely confined to the most remote island groups. Here we show that the gastropod Astralium "rhodostomum" has developed endemic clades on almost every Pacific archipelago sampled, a pattern unprecedented in marine biogeography, and reminiscent of the terrestrial biota of oceanic islands. Mitochondrial DNA sequences indicate that this species-complex is comprised of at least 30 geographically isolated clades, separated by as little as 180 km. Evidence suggests that such fine scale endemism and high diversity is not exceptional, but likely characterizes a substantial fraction of the reef biota. These results imply that (1) marine speciation can regularly occur over much finer spatial scales than generally accepted, (2) the diversity of coral reefs is even higher than suggested by morphology-based estimates, and (3) conservation efforts need to focus at the archipelagic level in the sea as on land.     

Absence of post-Miocene Red Sea land bridges: biogeographic implications

In a large number of studies concerned with species movements between Africa and Eurasia, including the migrations of hominids out of Africa, a frequently‐cited dispersal route is across a hypothetical land bridge in the southern Red Sea, which is suggested to have emerged during glacial sea‐level lowstands. This paper, however, unequivocally demonstrates that palaeoceanographic and palaeoecological data are incompatible with the existence of Red Sea land bridges since the Miocene. The case is made by presenting the first quantitative history of water depth above the Red Sea sill for the last 470,000 years, a time period that includes the four most recent glacial–interglacial cycles, and by discussing the predictable consequences of any land bridge formation on the Red Sea sedimentary and microfossil records. The absence of post‐Miocene Red Sea land bridges has extensive implications for biogeographic models in the Afro‐Arabian region. Genetic, morphometric and palaeontological patterns reported in the literature cannot be related to dispersals over a land bridge, or in the case of marine organisms, separation of the Red Sea from the Indian Ocean by a land bridge. If such patterns in terrestrial species are only congruent with a southern Red Sea dispersal route, then they need to be considered in terms of sweepstake rafting, anthropogenic introduction, or in the particular case of the Out‐of‐Africa migration by modern humans, seafaring. The constraints imposed by our palaeoenvironmental record on biogeographic reconstructions within and around the Red Sea will hopefully encourage both the review of previous works and the preference for multidisciplinary approaches in future studies.     

Natural and anthropogenic dispersal mechanisms in the marine environment: a study using cheilostome Bryozoa

The global geographic ranges occupied by 197 species of cheilostomate Bryozoa found in British waters were obtained by a literature survey. Morphological grade, larval mode, environmental tolerance, species abundance and the ability to raft and to foul shipping were all investigated as traits potentially able to affect the geographic ranges of these bryozoan species. When considered independently all variables except larval mode had a significant correlation with the geographic range occupied by a species. However, when controlling for the potentially confounding effects of the other covariates, only the ability to foul or raft and species abundance had a significant effect on median geographic range and only fouling and abundance had a significant effect over global ranges. The strength of the association between fouling ability and range suggests that transport upon the hulls of ships is a very important dispersal mechanism for bryozoans, as it is thought to be also for various other marine taxa. Potential long-term (evolutionary) consequences of increased ranges brought about by anthropogenic mechanisms are discussed.     

The influence of coastal topography, circulation patterns, and rafting in structuring populations of an intertidal alga

Understanding the dispersal processes that influence genetic structure in marine species requires estimating gene flow in a dynamic, fluid environment that is often poorly characterized at scales relevant to multiple dispersive stages (e.g. spores, gametes, zygotes, larvae, adults). We examine genetic structure in the marine alga Fucus vesiculosus L., which inhabits moderately exposed shores in the northern Atlantic but releases gametes only under sunny, calm conditions. We predicted genetic structure would correlate with coastal topography because weather frequently varies across coastal promontories on the Maine shore when F. vesiculosus is reproductive, which causes one side to experience high levels of water motion (= no gamete release) while one side is calm (= gamete release). Furthermore, we expected that the effect of low dispersal capacities of gametes and zygotes would result in spatial genetic structure over short distances. Using surface drifters, we characterized near-shore circulation patterns around the study sites to investigate whether directionality of gene flow was correlated with directionality of currents. We found significant genetic differentiation among sites sampled at two different peninsulas, but patterns of differentiation were unrelated to coastal topography and there was no within-site spatial structuring. Our genetic and near-shore circulation data, combined with an examination of gamete longevity, support the dependency of gene flow on storm-detached, rafting, reproductive adults. This study highlights the significance of rafting as a mechanism for structuring established populations of macroalgae and associated biota and demonstrates the importance of coupling population genetics' research with relevant hydrodynamic studies.