The Importance of the Human Footprint in Shaping the Global Distribution of Terrestrial,Freshwater and Marine Invaders

Human activities such as transport, trade and tourism are likely to influence the spatial distribution of non-native species and yet, Species Distribution Models (SDMs) that aim to predict the future broad scale distribution of invaders often rely on environmental (e.g. climatic) information only. This study investigates if and to what extent do human activities that directly or indirectly influence nature (hereafter the human footprint) affect the global distribution of invasive species in terrestrial, freshwater and marine ecosystems. We selected 72 species including terrestrial plants, terrestrial animals, freshwater and marine invasive species of concern in a focus area located in NW Europe (encompassing Great Britain, France, The Netherlands and Belgium). Species Distribution Models were calibrated with the global occurrence of species and a set of high-resolution (9×9 km) environmental (e.g. topography, climate, geology) layers and human footprint proxies (e.g. the human influence index, population density, road proximity). Our analyses suggest that the global occurrence of a wide range of invaders is primarily limited by climate. Temperature tolerance was the most important factor and explained on average 42% of species distribution. Nevertheless, factors related to the human footprint explained a substantial amount (23% on average) of species distributions. When global models were projected into the focus area, spatial predictions integrating the human footprint featured the highest cumulative risk scores close to transport networks (proxy for invasion pathways) and in habitats with a high human influence index (proxy for propagule pressure). We conclude that human related information–currently available in the form of easily accessible maps and databases—should be routinely implemented into predictive frameworks to inform upon policies to prevent and manage invasions. Otherwise we might be seriously underestimating the species and areas under highest risk of future invasions.     

The direct physical,chemical and biotic impacts on Australian coastal waters due to recreational boating

In economically developed countries it is projected that by around 2015 over 50% of a person’s lifetime will become available for leisure. Demand for leisure needs, already strong, will continue to increase. One segment of the market, outdoor nature-based recreation (including tourism), is growing strongly worldwide. A substantial proportion of these activities are water-based. The associated demand for recreational vessels has increased rapidly in recent years and is projected to continue to trend upwards. Australian trends mirror those internationally. Using Australia as a case study, we review the direct physical, chemical and biotic impacts associated with recreational boating in coastal water environments. Major physical impacts include disturbance due to movement of craft in shallow waters (e.g., turbulence) and the effects of anchoring/drag, noise/interference/collision that impacts on wildlife. The most critical chemical impacts result from pollution due to fuels and oils, defouling treatments (even those not legislated in-country), and human waste (e.g., sewage effluent). Important biotic impacts are the potential continued introduction and secondary spread of non-native species. We conclude that while greater research effort will provide more environmentally benign products, with the increasing popularity of recreation vessels, it will be beyond the resources of Australian governments to police legislation effectively. However, based on Australian’s demonstrated engagement with government in terrestrial environmental management, with their deliberate engagement with the boating fraternity, the impacts of recreational boating would be lessened.     

Extracellular enzymes in terrestrial, freshwater, and marine environments: perspectives on system variability and common research needs

Extracellular enzymes produced by heterotrophic microbial communities are major drivers of carbon and nutrient cycling in terrestrial, freshwater, and marine environments. Although carbon and nutrient cycles are coupled on global scales, studies of extracellular enzymes associated with terrestrial, freshwater, and marine microbial communities are not often compared across ecosystems. In part, this disconnect arises because the environmental parameters that control enzyme activities in terrestrial and freshwater systems, such as temperature, pH, and moisture content, have little explanatory power for patterns of enzyme activities in marine systems. Instead, factors such as the functional diversity of microbial communities may explain varying patterns of enzyme activities observed in the ocean to date. In any case, many studies across systems focus on similar issues that highlight the commonalities of microbial community organization. Examples include the effective lifetime of enzymes released into the environment; the extent to which microbial communities coordinate enzyme expression to decompose complex organic substrates; and the influence of microbial community composition on enzyme activities and kinetics. Here we review the often-disparate research foci in terrestrial, freshwater, and marine environments. We consider the extent to which environmental factors may regulate extracellular enzyme activities within each ecosystem, and highlight commonalities and current methodological challenges to identify research questions that may aid in integrating cross-system perspectives in the future.     

Fish species introductions provide novel insights into the patterns and drivers of phylogenetic structure in freshwaters

Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. We quantify phylogenetic clustering and overdispersion of native and non-native fishes of a large river basin in the American Southwest to test for the mechanisms (environmental filtering versus competitive exclusion) and spatial scales influencing community structure. Contrary to expectations, non-native species were phylogenetically clustered and related to natural environmental conditions, whereas native species were not phylogenetically structured, likely reflecting human-related changes to the basin. The species that are most invasive (in terms of ecological impacts) tended to be the most phylogenetically divergent from natives across watersheds, but not within watersheds, supporting the hypothesis that Darwin''s naturalization conundrum is driven by the spatial scale. Phylogenetic distinctiveness may facilitate non-native establishment at regional scales, but environmental filtering restricts local membership to closely related species with physiological tolerances for current environments. By contrast, native species may have been phylogenetically clustered in historical times, but species loss from contemporary populations by anthropogenic activities has likely shaped the phylogenetic signal. Our study implies that fundamental mechanisms of community assembly have changed, with fundamental consequences for the biogeography of both native and non-native species.     

Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems

The cycles of the key nutrient elements nitrogen (N) and phosphorus (P) have been massively altered by anthropogenic activities. Thus, it is essential to understand how photosynthetic production across diverse ecosystems is, or is not, limited by N and P. Via a large-scale meta-analysis of experimental enrichments, we show that P limitation is equally strong across these major habitats and that N and P limitation are equivalent within both terrestrial and freshwater systems. Furthermore, simultaneous N and P enrichment produces strongly positive synergistic responses in all three environments. Thus, contrary to some prevailing paradigms, freshwater, marine and terrestrial ecosystems are surprisingly similar in terms of N and P limitation.     

Non-native fishes and native species diversity in freshwater fish assemblages across the United States

The proliferation of non-native species in North American freshwater ecosystems is considered a primary threat to the integrity of native community structure. However, a general understanding of consistent and predictable impacts of non-native species on native freshwater diversity is limited, in part, because of a lack of broad-scale studies including data from numerous localities across multiple drainages. This study uses data from 751 localities collected during the United States Geological Survey (USGS) National Water Quality Assessment (NAWQA) program to examine the influence of non-native fish species on native freshwater fish assemblages across the United States. In general, no significant differences in native fish richness and diversity measures were detected between sites with only native species and sites containing non-native species. However, at sites with non-native species, the number of non-native species present was negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Non-native piscivores were negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Native piscivores were positively correlated with native richness and diversity and negatively correlated with native evenness at sites with only native species. Our results suggest that from a superficial perspective, native species richness and diversity are not different among sites with and without non-native species. However, when patterns of native species richness and diversity are examined at sites containing non-native species, correlations between non-native and native species richness and diversity imply the expected negative effect of invasive taxa. Additionally, non-native piscivores appear to have a significant negative effect on native taxa and possibly represent a novel selective force on naive native prey.     

Latitudinal gradients in biotic niche breadth vary across ecosystem types

Several properties of food webs—the networks of feeding links between species—are known to vary systematically with the species richness of the underlying community. Under the ‘latitude–niche breadth hypothesis’, which predicts that species in the tropics will tend to evolve narrower niches, one might expect that these scaling relationships could also be affected by latitude. To test this hypothesis, we analysed the scaling relationships between species richness and average generality, vulnerability and links per species across a set of 196 empirical food webs. In estuarine, marine and terrestrial food webs there was no effect of latitude on any scaling relationship, suggesting constant niche breadth in these habitats. In freshwater communities, on the other hand, there were strong effects of latitude on scaling relationships, supporting the latitude–niche breadth hypothesis. These contrasting findings indicate that it may be more important to account for habitat than latitude when exploring gradients in food-web structure.     

The First Freshwater Mosasauroid (Upper Cretaceous,Hungary) and a New Clade of Basal Mosasauroids

Mosasauroids are conventionally conceived of as gigantic, obligatorily aquatic marine lizards (1000s of specimens from marine deposited rocks) with a cosmopolitan distribution in the Late Cretaceous (90–65 million years ago [mya]) oceans and seas of the world. Here we report on the fossilized remains of numerous individuals (small juveniles to large adults) of a new taxon, Pannoniasaurus inexpectatus gen. et sp. nov. from the Csehbánya Formation, Hungary (Santonian, Upper Cretaceous, 85.3–83.5 mya) that represent the first known mosasauroid that lived in freshwater environments. Previous to this find, only one specimen of a marine mosasauroid, cf. Plioplatecarpus sp., is known from non-marine rocks in Western Canada. Pannoniasaurus inexpectatus gen. et sp. nov. uniquely possesses a plesiomorphic pelvic anatomy, a non-mosasauroid but pontosaur-like tail osteology, possibly limbs like a terrestrial lizard, and a flattened, crocodile-like skull. Cladistic analysis reconstructs P. inexpectatus in a new clade of mosasauroids: (Pannoniasaurus (Tethysaurus (Yaguarasaurus, Russellosaurus))). P. inexpectatus is part of a mixed terrestrial and freshwater faunal assemblage that includes fishes, amphibians turtles, terrestrial lizards, crocodiles, pterosaurs, dinosaurs and birds.     

Molecular and developmental contributions to divergent pigment patterns in marine and freshwater sticklebacks

Pigment pattern variation across species or populations offers a tractable framework in which to investigate the evolution of development. Juvenile threespine sticklebacks (Gasterosteus aculeatus) from marine and freshwater environments exhibit divergent pigment patterns that are associated with ecological differences. Juvenile marine sticklebacks have a silvery appearance, whereas sticklebacks from freshwater environments exhibit a pattern of vertical bars. We investigated both the developmental and molecular basis of this population‐level variation in pigment pattern. Time course imaging during the transition from larval to juvenile stages revealed differences between marine and freshwater fish in spatial patterns of chromatophore differentiation as well as in pigment amount and dispersal. In freshwater fish, melanophores appear primarily within dark bars whereas iridophores appear within light bars. By contrast, in marine fish, these chromatophores are interspersed across the flank. In addition to spatially segregated chromatophore differentiation, pigment amount and dispersal within melanophores varies spatially across the flank of freshwater, but not marine fish. To gain insight into the molecular pathways that underlie the differences in pigment pattern development, we evaluated differential gene expression in the flanks of developing fish using high‐throughput cDNA sequencing (RNA‐seq) and quantitative PCR. We identified several genes that were differentially expressed across dark and light bars of freshwater fish, and between freshwater and marine fish. Together, these experiments begin to shed light on the process of pigment pattern evolution in sticklebacks.     

Prudent sessile feeding by the corallivore snail,Coralliophila violacea on coral energy sinks

Convergence of form and function has accompanied the evolution of modular growth in terrestrial plants and colonial marine invertebrates. Part of this convergence is related to the optimal exploitation of resources (space and light) and the ability to translocate energy products from sources to sink sites. Feeding on the energy pathways and energy sinks of terrestrial plants is a well-known phenomenon. Hermatypic corals, the major organisms constructing tropical reef environments, contain photosynthetic algae (zooxanthellae), energetic products of which are translocated towards sink sites located at the corals'' growing tips and regenerating areas. Despite the plant–coral convergence in energy pathways and sinks, there has been no evidence to date that coral energy sinks are exploited by coral predators. Gastropods of the genus Coralliophila are found feeding on coral margins, causing small and localized tissue damage. However, the ability of these snails to continue to feed without moving over a long period remains puzzling. Using a ¹⁴C labelling technique, we found that colony margins of the stony coral Porites function as major energy sinks. When snails inhabited these sites they incorporated significant amounts of ¹⁴C, indicating that they had fed on photosynthetic products translocated from the interior of the colony. Furthermore, when snails aggregate in the interior of the colony, thereby causing large surface injuries, they induce the development of significant new sink sites. This mode of prudent sessile feeding maximizes the efficiency of energy exploitation by the predatory snail, while minimizing tissue damage to the coral. The fact that energy sink sites occur in many coral species suggests that the strategy of sink exploitation for nutrition could also occur in many other marine host–symbiont relationships.     

Changes in pathways and vectors of biological invasions in Northwest Europe

We assessed how establishment patterns of non-native freshwater, marine and terrestrial species into Northwest Europe (using Great Britain, France, Belgium and the Netherlands as the study countries) have changed over time, and identified the prevalent pathways and vectors of recent arrivals. Data were extracted from 33 sources on (a) presence/absence and (b) first year of observation in the wild in each country, and (c) continent(s) of origin, (d) invasion pathway(s), (e) invasion vector(s) and (f) environment(s) for 359 species, comprising all non-native Mollusca, Osteichthyes (bony fish), Anseriformes (wildfowl) and Mammalia, and non-native invasive Angiospermae present in the area. Molluscs, fish and wildfowl, particularly those originating from South America, arrived more recently into Northwest Europe than other groups, particularly mammals, invasive plants and species originating from North America. Non-deliberate introductions, those of aquatic species and those from elsewhere in Europe and/or Asia increased strongly in importance after the year 2000 and were responsible for 69, 83 and 89 % of new introductions between 2001 and 2015, respectively. Non-deliberate introductions and those from Asia and North America contributed significantly more to introductions of invasive species in comparison to other non-native species. From the 1960s, ornamental trade has increased in importance relative to other vectors and was responsible for all deliberate introductions of study groups since 2001. Non-deliberate introductions of freshwater and marine species originating from Southeast Europe and Asia represent an increasingly important ecological and economic threat to Northwest Europe. Invertebrates such as molluscs may be particularly dangerous due to their small size and difficulties in detection. Prevention of future invasions in this respect will require intensive screening of stowaways on boats and raising of public awareness.     

Biosecurity and Vector Behaviour: Evaluating the Potential Threat Posed by Anglers and Canoeists as Pathways for the Spread of Invasive Non-Native Species and Pathogens

Invasive non-native species (INNS) endanger native biodiversity and are a major economic problem. The management of pathways to prevent their introduction and establishment is a key target in the Convention on Biological Diversity''s Aichi biodiversity targets for 2020. Freshwater environments are particularly susceptible to invasions as they are exposed to multiple introduction pathways, including non-native fish stocking and the release of boat ballast water. Since many freshwater INNS and aquatic pathogens can survive for several days in damp environments, there is potential for transport between water catchments on the equipment used by recreational anglers and canoeists. To quantify this biosecurity risk, we conducted an online questionnaire with 960 anglers and 599 canoeists to investigate their locations of activity, equipment used, and how frequently equipment was cleaned and/or dried after use. Anglers were also asked about their use and disposal of live bait. Our results indicate that 64% of anglers and 78.5% of canoeists use their equipment/boat in more than one catchment within a fortnight, the survival time of many of the INNS and pathogens considered in this study and that 12% of anglers and 50% of canoeists do so without either cleaning or drying their kit between uses. Furthermore, 8% of anglers and 28% of canoeists had used their equipment overseas without cleaning or drying it after each use which could facilitate both the introduction and secondary spread of INNS in the UK. Our results provide a baseline against which to evaluate the effectiveness of future biosecurity awareness campaigns, and identify groups to target with biosecurity awareness information. Our results also indicate that the biosecurity practices of these groups must improve to reduce the likelihood of inadvertently spreading INNS and pathogens through these activities.     

Genetic conflict,kin and the origins of novel genetic systems

Genetic conflict may have played an important role in the evolution of novel genetic systems. The ancestral system of eumendelian genetics is highly symmetrical. Those derived from it (e.g. thelytokous parthenogenesis, haplodiploidy and parent-specific allele expression) are more asymmetrical in the genetic role played by maternal versus paternal alleles. These asymmetries may have arisen from maternal–paternal genetic conflict, or cytonuclear conflict, or from an interaction between them. Asymmetric genetic systems are much more common in terrestrial and freshwater taxa than in marine taxa. We suggest three reasons for this, based on the relative inhospitability of terrestrial environments to three types of organism: (i) pathogens—departure from the marine realm meant escape from many pathogens and parasites, reducing the need for sexual reproduction; (ii) symbionts—symbionts are no more important in the terrestrial realm than the marine realm but are more likely to be obligately intracellular and vertically transmitted, making them more likely to disrupt their host''s genetic systems; (iii) Gametes and embryos—because neither gametes nor embryos can be shed into air as easily as into seawater, the mother''s body is a more important environment for both types of organisms in the terrestrial realm than in the marine realm. This environment of asymmetric kinship (with neighbours more closely related by maternal alleles than by paternal alleles) may have helped to drive asymmetries in expression and transmission.     

Antarctica: The final frontier for marine biological invasions

Antarctica is experiencing significant ecological and environmental change, which may facilitate the establishment of non‐native marine species. Non‐native marine species will interact with other anthropogenic stressors affecting Antarctic ecosystems, such as climate change (warming, ocean acidification) and pollution, with irreversible ramifications for biodiversity and ecosystem services. We review current knowledge of non‐native marine species in the Antarctic region, the physical and physiological factors that resist establishment of non‐native marine species, changes to resistance under climate change, the role of legislation in limiting marine introductions, and the effect of increasing human activity on vectors and pathways of introduction. Evidence of non‐native marine species is limited: just four marine non‐native and one cryptogenic species that were likely introduced anthropogenically have been reported freely living in Antarctic or sub‐Antarctic waters, but no established populations have been reported; an additional six species have been observed in pathways to Antarctica that are potentially at risk of becoming invasive. We present estimates of the intensity of ship activity across fishing, tourism and research sectors: there may be approximately 180 vessels and 500+ voyages in Antarctic waters annually. However, these estimates are necessarily speculative because relevant data are scarce. To facilitate well‐informed policy and management, we make recommendations for future research into the likelihood of marine biological invasions in the Antarctic region.     

Ecological selectivity and the evolution of mammalian substrate preference across the K–Pg boundary

The Cretaceous–Paleogene (K–Pg) mass extinction 66 million years ago was characterized by a worldwide ecological catastrophe and rapid species turnover. Large‐scale devastation of forested environments resulting from the Chicxulub asteroid impact likely influenced the evolutionary trajectories of multiple clades in terrestrial environments, and it has been hypothesized to have biased survivorship in favour of nonarboreal lineages across the K–Pg boundary. Here, we evaluate patterns of substrate preferences across the K–Pg boundary among crown group mammals, a group that underwent rapid diversification following the mass extinction. Using Bayesian, likelihood, and parsimony reconstructions, we identify patterns of mammalian ecological selectivity that are broadly similar to those previously hypothesized for birds. Models based on extant taxa indicate predominant K–Pg survivorship among semi‐ or nonarboreal taxa, followed by numerous independent transitions to arboreality in the early Cenozoic. However, contrary to the predominant signal, some or all members of total‐clade Euarchonta (Primates + Dermoptera + Scandentia) appear to have maintained arboreal habits across the K–Pg boundary, suggesting ecological flexibility during an interval of global habitat instability. We further observe a pronounced shift in character state transitions away from plesiomorphic arboreality associated with the K–Pg transition. Our findings are consistent with the hypothesis that predominantly nonarboreal taxa preferentially survived the end‐Cretaceous mass extinction, and emphasize the pivotal influence of the K‐Pg transition in shaping the early evolutionary trajectories of extant terrestrial vertebrates.     

Propagule pressure and the invasion risks of non-native freshwater fishes: a case study in England

European countries in general, and England in particular, have a long history of introducing non-native fish species, but there exist no detailed studies of the introduction pathways and propagules pressure for any European country. Using the nine regions of England as a preliminary case study, the potential relationship between the occurrence in the wild of non-native freshwater fishes (from a recent audit of non-native species) and the intensity ( i.e. propagule pressure) and diversity of fish imports was investigated. The main pathways of introduction were via imports of fishes for ornamental use ( e.g. aquaria and garden ponds) and sport fishing, with no reported or suspected cases of ballast water or hull fouling introductions. The recorded occurrence of non-native fishes in the wild was found to be related to the time (number of years) since the decade of introduction. A shift in the establishment rate, however, was observed in the 1970s after which the ratio of established-to-introduced species declined. The number of established non-native fish species observed in the wild was found to increase significantly ( P < 0·05) with increasing import intensity (log₁₀ x + 1 of the numbers of fish imported for the years 2000–2004) and with increasing consignment diversity (log₁₀ x + 1 of the numbers of consignment types imported for the years 2000–2004). The implications for policy and management are discussed.     

Characterizing mauka-to-makai connections for aquatic ecosystem conservation on Maui,Hawaiʻi

Mauka-to-makai (mountain to sea in the Hawaiian language) hydrologic connectivity – commonly referred to as ridge-to-reef – directly affects biogeochemical processes and socioecological functions across terrestrial, freshwater, and marine systems. The supply of freshwater to estuarine and nearshore environments in a ridge-to-reef system supports the food, water, and habitats utilized by marine fauna. In addition, the ecosystem services derived from this land-to-sea connectivity support social and cultural practices (hereafter referred to as socio-cultural) including fishing, aquaculture, wetland agriculture, religious ceremonies, and recreational activities. To effectively guide island resource management, a better understanding of the linkages from ridge-to-reef across natural and social usages is critical, particularly in the context of climate change, with anticipated increasing temperature and shifting precipitation patterns. The objective of this study was to identify spatial linkages that promote multiple and diverse uses, following the ridge-to-reef concept, at an island-wide scale to identify regions of high conservation importance for aquatic resources. We selected the Island of Maui as a study representative of many Pacific islands. Diverse datasets, including agricultural lands within watersheds, wetland locations, presence of stream species, indicators of freshwater input from streams, coral cover, nearshore fish biomass, socio-cultural data such as fishpond locations, wetland taro cultivation, beach recreation use, and lastly the dynamically downscaled Coupled Model Intercomparison Project Phase (CMIP5) future climate projections scenarios (Representative Concentration Pathway (RCP) 4.5 & 8.5) were used to examine the spatial linkages through hydrological connectivity from land to the sea. Zonation spatial planning software was used to prioritize areas of high management and conservation value and to help inform aquatic resources management. The resulting prioritized areas included many minimally disturbed watersheds in east Maui and western nearshore and coastal zones that are adjacent to diverse coral reefs. These results are driven by the importance of fish biomass and coral reef distribution as well as traditional wetland taro cultivation and coastal access points for recreation. These results underline the importance of examining ridge-to-reef systems for aquatic resource management and including important social and cultural values in resource management upon planning adaptation strategies for climate change. Improving our understanding of diverse natural and socio-cultural influences on habitat conditions and their values in these areas provides an opportunity to strategically plan future management and conservation actions.     

Quantification of climate tourism potential of Croatia based on measured data and regional modeling

Tourism is one of the most important economic sectors in Croatia. The Adriatic coast is a popular travel destination for tourists, especially during the summer months. During their activities, tourists are affected by atmospheric conditions and therefore by weather and climate. Therefore, it is important to have reliable information about thermal conditions as well as their impacts on human beings. Here, the climate tourism potential of Croatia is presented and quantified on the basis of three selected stations in different climatic regions. The physiologically equivalent temperature is used for analysis as well as other climatic parameters relevant for tourism and recreation. The results already point to hot conditions for outdoor activities in summer during afternoons, especially along the coast but also for continental regions, resulting in a reduction of the climate tourism potential. In the future, this trend looks set to increase, possibly leading to a changing tourism sector in Croatia requiring adaptation and new strategies.     

Lost in space? Searching for directions in the spatial modelling of individuals,populations and species ranges

The workshop ‘Spatial models in animal ecology, management and conservation’ held at Silwood Park (UK), 9–11 March 2010, aimed to synthesize recent progress in modelling the spatial dynamics of individuals, populations and species ranges and to provide directions for research. It brought together marine and terrestrial researchers working on spatial models at different levels of organization, using empirical as well as theory-driven approaches. Different approaches, temporal and spatial scales, and practical constraints predominate at different levels of organization and in different environments. However, there are theoretical concepts and specific methods that can fruitfully be transferred across levels and systems, including: habitat suitability characterization, movement rules, and ways of estimating uncertainty.