Oceanic Island Systems Research Articles

Phthalates and fatty acid markers in free-ranging cetaceans from an insular oceanic region: Ecological niches as drivers of contamination

Plastic additives, such as phthalates, are ubiquitous contaminants that can have detrimental impacts on marine organisms and overall ecosystems' health. Valuable information about the status and resilience of marine ecosystems can be obtained through the monitoring of key indicator species, such as cetaceans. In this study, fatty acid profiles and phthalates were examined in blubber biopsies of free-ranging individuals from two delphinid species (short-finned pilot whale – Globicephala macrorhynchus, n = 45; common bottlenose dolphin – Tursiops truncatus, n = 39) off Madeira Island (NE Atlantic). This investigation aimed to explore the relations between trophic niches (epipelagic vs. mesopelagic), contamination levels, and the health status of individuals within different ecological and biological groups (defined by species, residency patterns and sex). Multivariate analysis of selected dietary fatty acids revealed a clear niche segregation between the two species. Di-n-butylphthalate (DBP), diethyl phthalate (DEP), and bis(2-ethylhexyl) phthalate (DEHP) were the most prevalent among the seven studied phthalates, with the highest concentration reached by DEHP in a bottlenose dolphin (4697.34 ± 113.45 ng/g). Phthalates esters (PAEs) concentration were higher in bottlenose dolphins (Mean ∑ PAEs: 947.56 ± 1558.34 ng/g) compared to pilot whales (Mean ∑ PAEs: 229.98 ± 158.86 ng/g). In bottlenose dolphins, DEHP was the predominant phthalate, whereas in pilot whales, DEP and DBP were more prevalent. Health markers suggested pilot whales might suffer from poorer physiological conditions than bottlenose dolphins, although high metabolic differences were seen between the two species. Phthalate levels showed no differences by ecological or biological groups, seasons, or years. This study is the first to assess the extent of plastic additive contamination in free-ranging cetaceans from a remote oceanic island system, underscoring the intricate relationship between ecological niches and contaminant exposure. Monitoring these chemicals and their potential impacts is vital to assess wild population health, inform conservation strategies, and protect critical species and habitats.

Diel vertical migration and seamount stepping stones promote species connectivity from coastal to offshore insular systems in the Tropical Southwestern Atlantic

AbstractThe recruitment of marine species in isolated oceanic island systems can be challenged by prevailing currents, as exemplified by the Tropical Southwestern Atlantic. In this region, the Fernando de Noronha ridge hosts several seamounts, the Rocas Atoll and the Fernando de Noronha Archipelago, which are home to great marine biodiversity. However, along the ridge, the central branch of the South Equatorial Current (cSEC), flowing westward, poses a challenge to the recruitment of organisms toward Fernando de Noronha. To unveil critical insights into the intricate processes shaping biodiversity in these insular ecosystems, we use a dispersal Lagrangian tool to explore the role of diel vertical migration (DVM) to depth strata influenced by the South Equatorial Undercurrent (SEUC), which flows eastward bellow the cSEC, in shaping species dispersal and metacommunity dynamics. Our results show that while not a direct journey, the DVM into SEUC‐influenced strata increases the possibility that the seamounts and the Rocas Atoll act as stepping stones between the continental shelf and Fernando de Noronha. Propagules of organisms originating primarily from the continental shelf are transported to the western seamounts of the ridge. Upon reaching the western seamounts, organisms can find suitable habitats to recruit. The progeny of these communities that migrate to SEUC‐influenced strata have the opportunity to reach suitable habitats at the Rocas Atoll and the Eastern seamounts, ultimately connecting to the Fernando de Noronha archipelago. These results provide scientific fundaments for the development of a functional network of marine protected areas in the Tropical Southwestern Atlantic.

Waves of Colonization and Gene Flow in a Great Speciator.

Secondary contact between previously allopatric lineages offers a test of reproductive isolating mechanisms that may have accrued in isolation. Such instances of contact can produce stable hybrid zones-where reproductive isolation can further develop via reinforcement or phenotypic displacement-or result in the lineages merging. Ongoing secondary contact is most visible in continental systems, where steady input from parental taxa can occur readily. In oceanic island systems, however, secondary contact between closely related species of birds is relatively rare. When observed on sufficiently small islands, relative to population size, secondary contact likely represents a recent phenomenon. Here, we examine the dynamics of a group of birds whose apparent widespread hybridization influenced Ernst Mayr's foundational work on allopatric speciation: the whistlers of Fiji (Aves: Pachycephala). We demonstrate two clear instances of secondary contact within the Fijian archipelago, one resulting in a hybrid zone on a larger island, and the other resulting in a wholly admixed population on a smaller, adjacent island. We leveraged low genome-wide divergence in the hybrid zone to pinpoint a single genomic region associated with observed phenotypic differences. We use genomic data to present a new hypothesis that emphasizes rapid plumage evolution and post-divergence gene flow.

Cassiopea andromeda (Cnidaria, Scyphozoa) in the subtropical eastern Atlantic

Abstract This study provides the first records of the upside-down jellyfish Cassiopea andromeda (Forskål, 1775) in the eastern Atlantic supported by molecular analysis. Specimens were observed, recorded, and sampled in an inland aquaculture facility in September 2023 in Tenerife Island (Canary Islands). This new record officially demonstrates the geographical expansion of C. andromeda, and the introduction of a new potential invasive species in the Macaronesia oceanic island system.

Seasonal variation in microplastics and zooplankton abundances and characteristics: The ecological vulnerability of an oceanic island system

The ingestion of microplastics (MPs - plastic particles <5 mm) by planktivorous organisms represents a significant threat to marine food webs. To investigate how seasonality might affect plastic intake in oceanic islands' ecosystems, relative abundances and composition of MPs and mesozooplankton samples collected off Madeira Island (NE Atlantic) between February 2019 and January 2020 were analysed. MPs were found in all samples, with fibres accounting for 89 % of the particles. MPs and zooplankton mean abundance was 0.262 items/m3 and 18.137 individuals/m3, respectively. Their monthly variations follow the seasonal fluctuation of environmental parameters, such as currents, chlorophyll-a concentration, sea surface temperature and precipitation intensity. A higher MPs/zooplankton ratio was recorded in the warm season (May-Oct), reaching 0.068 items/individual when considering large-sized particles (1000–5000 μm). This is the first study to assess the seasonal variability of MPs in an oceanic island system providing essential information respecting its ecological impact in pelagic environments.

Seasonal Variation in Microplastics and Zooplankton Traits: The Ecological Vulnerability of an Oceanic Island System

Seasonal Variation in Microplastics and Zooplankton Traits: The Ecological Vulnerability of an Oceanic Island System

Similar trait structure and vulnerability in pelagic fish faunas on two remote island systems

The link between biodiversity and ecosystem functioning has been the topic of considerable research, but it remains unclear how biodiversity decline is compromising ecosystem functionality, particularly in the pelagic realm. Here, we explore how pelagic fish species diversity relates to functional diversity by sampling two locations, which, on the basis of biogeography, environmental conditions and human pressures, were expected to host pronounced differences in species composition and abundances and therefore functionality. Strings of five drifting mid-water Baited Remote Underwater Video Systems were used to survey pelagic vertebrate diversity and abundance in two isolated oceanic island systems, the Malpelo Fauna and Flora Sanctuary—a large, 25-year-old marine protected area—and an unprotected area in Cape Verde. Functional diversity, which offers insight into a community’s resilience against disturbance, was analysed using six key functional traits of marine fishes. Abundance was recorded as MaxN, the maximum number of individuals of a given species in a single frame during the 2-h deployment time. Cape Verde showed high overall abundance (Total MaxN 873) and low biomass (3559 kg), with a predominance of smaller fishes. Malpelo showed high biomass (7839 kg) but lower abundance (Total MaxN 465), with a predominance of large species. Species and functional diversity were marginally different between locations. Multivariate analysis of species relative abundances showed significant divergence between locations, although community functional traits overlapped strongly, suggesting that both communities share a similar structure and vulnerability. The existence of a common functional ‘backbone’ in diverging species communities across the oceans, under different productivity regimes, and under different protection levels, suggests that although pelagic communities may differ considerably in terms of species composition, this does not translate into a differing functional structure and resilience potential. Whether this vulnerability is a common feature of pelagic communities and how this contrasts with benthic systems warrants further research.

Facies architecture, modal analysis and geochemistry of the Early Ediacaran turbiditic series in the Skoura inlier (Central High Atlas, Morocco): Implications for provenance and geotectonic setting.

A multidisciplinary approach based on lithostratigraphy, petrography, modal composition and geochemistry was performed for the first time, to study the Early Ediacaran Skoura Turbidites (EEST) of the Skoura inlier, Central High Atlas. It represents widely distributed turbidite successions up to 8 km in thickness. A total of twelve lithofacies types have been distinguished, spread in four profiles, and grouped in three major depositional environments: shallow delta complex (FA1), submarine channel (FA2), and deep-sea fan (FA3). The paleogeographic distribution of the EEST may indicate basin depth changes related to the onset of convergence.Petrographic and modal analyses of EEST reveal greywacke and lith-arenite lithologies displaying similar clast composition, mainly comprising quartz and lithic fragments. Volcanic, plutonic, and metamorphic rocks have been exposed in the source areas feeding the Skoura basin. On Q-F-L diagram, EEST have characteristics of a recycled orogen, suggesting their derivation from an active arc setting. Geochemical data of EEST samples display similar SiO2 and Al2O3 abundances, with relatively intermediate compositions. Most of the studied samples demonstrate their derivation from a composite of oceanic island and continental arc systems providing felsic and mafic to intermediate detritus to the Skoura basin. Rare-earth element patterns show features similar to those of sandstones in modern active margin zones. The EEST rocks are similar with other Early Ediacaran sediments of the Saghro Group in the Anti-Atlas belt. This architecture suggests deposition of the EEST in an orogenic basin developed in a back-arc setting during the Early Ediacaran time.

Tourism and urban development as drivers for invertebrate diversity loss on tropical islands.

Oceanic islands harbour a disproportionately high number of endemic and threatened species. Rapidly growing human populations and tourism are posing an increasing threat to island biota, yet the ecological consequences of these human land uses on small oceanic island systems have not been quantified. Here, we investigated and compared the impact of tourism and urban island development on ground-associated invertebrate biodiversity and habitat composition on oceanic islands. To disentangle tourism and urban land uses, we investigated Indo-Pacific atoll islands, which either exhibit only tourism or urban development, or remain uninhabited. Within the investigated system, we show that species richness, abundance and Shannon diversity of the investigated invertebrate community are significantly decreased under tourism and urban land use, relative to uninhabited islands. Remote-sensing-based spatial data suggest that habitat fragmentation and a reduction in vegetation density are having significant effects on biodiversity on urban islands, whereas land use/cover changes could not be linked to the documented biodiversity loss on tourist islands. This offers the first direct evidence for a major terrestrial invertebrate loss on remote oceanic atoll islands due to different human land uses with yet unforeseeable long-term consequences for the stability and resilience of oceanic island ecosystems.

Assessing gaps in reporting non-target mortality in island rodent eradication operations

Eradicating invasive species is a key part of island restoration, and can reverse the devastating impacts on native biota. Rodents are one of the most widespread invasive species, found on 80% of oceanic island systems, but have been removed from hundreds of islands through the application of anticoagulant-treated cereal bait. While such eradication operations are often net positive events for island ecosystems over the long-term, some native biota are also susceptible, resulting in short-term non-target mortality. One of the most widely distributed groups of birds, rails and allies (Rallidae) are highly adaptable, often endemic, and are known often to suffer mortality during rodent eradication operations, to varying degrees. Our goal was determine if the year of eradication or the size of the island predicted whether non-target mortalities were reported, including those that were true absences of mortality. We examined 122 eradication operations on 81 islands with rails present from 1983 to 2015, and found 78% with no reported information on non-target mortality using our search criteria. We found non-target mortality reporting has decreased over time, and there was no relationship with island size. Post-operational monitoring of eradication operations should thoroughly record non-target mortality to improve our understanding of factors affecting non-target mortality, and the efficacy of mitigation measures.

Evolution in Extremis: The When, How, and Why of Hawaiian Carabid Beetles

The Hawaiian Archipelago is the most isolated oceanic island system in the World, separated from the nearest source areas by more than 4000 km. Five independent colonization events have resulted in diversification of a native carabid beetle fauna in excess of 400 known species. This diverse assemblage is disharmonic, with the major radiations restricted to the platynine genus Blackburnia Sharp (139 species), the subgenus Nesocidium Sharp of Bembidion Latreille (21 species), and the moriomorphine genus Mecyclothorax Sharp (239 species). Biogeographical, ecological, and evolutionary attributes of these three radiations are compared in order to determine factors crucial to carabid beetle diversification in this most-isolated situation. Biogeographical attributes include the age of origin of the constituent radiation, the island likely colonized by its common ancestor, and the biological characteristics, where known, of the colonizing ancestors for each independent radiation. Ecological attributes include the amount of habitat specialization undergone during each radiation, taking into account the primordial habitat colonized and the subsequent pattern of occupation of different habitat types during diversification. Evolutionary attributes include brachyptery, body-size evolution, sexual selection, and the evolution of specialized body conformations. It is shown that ecological specialization—i.e., occupation of a diverse array of ecological zones and microhabitats—in concert with reduced dispersal ability brought on by evolution of brachyptery are positively associated with enhanced levels of diversification. Comparing sympatric island faunas, it is shown that the latter operates synergistically with body size, as the smaller-bodied Mecyclothorax beetles speciate much more rapidly than the larger-bodied Blackburnia on Maui and Hawai῾i Island. Nonetheless, small body size does not gaurantee high diversity, as Bembidion beetles attain body sizes similar to Mecyclothorax beetles. Age of origin of a radiation is a subsidiary criterion for diversification given that the Mecyclothorax radiation commenced only 1.2 Ma, whereas it is hypothesized that Blackburnia have been resident in the Hawaiian archipelago for upwards of 28 Ma. Thus especially for Blackburnia we are constrained in our ability to know all of the evolutionary products of the radiation due to extinction of presumably all or nearly all species that occupied the now-sunken islands northwest of the oldest high island of Kauai. We are fortunate to know of several extinct Blackburnia species discovered in lowland subfossil deposits in Kauai, and these species provide crucial information now regarding future patterns of extinction. Sexual selection can be demonstrated for the Bembidion subgenus Nesocidium, and is a likely explanation for genitalic evolution over parts of the Mecyclothorax radiation, but it is not a phenomenon pervasively associated with increased levels of speciation.

Changing Ecological Opportunities Facilitated the Explosive Diversification of New Caledonian Oxera (Lamiaceae).

Phylogenies recurrently demonstrate that oceanic island systems have been home to rapid clade diversification and adaptive radiations. The existence of adaptive radiations posits a central role of natural selection causing ecological divergence and speciation, and some plant radiations have been highlighted as paradigmatic examples of such radiations. However, neutral processes may also drive speciation during clade radiations, with ecological divergence occurring following speciation. Here, we document an exceptionally rapid and unique radiation of Lamiaceae within the New Caledonian biodiversity hotspot. Specifically, we investigated various biological, ecological, and geographical drivers of species diversification within the genus Oxera. We found that Oxera underwent an initial process of rapid cladogenesis likely triggered by a dramatic period of aridity during the early Pliocene. This early diversification of Oxera was associated with an important phase of ecological diversification triggered by significant shifts of pollination syndromes, dispersal modes, and life forms. Finally, recent diversification of Oxera appears to have been further driven by the interplay of allopatry and habitat shifts likely related to climatic oscillations. This suggests that Oxera could be regarded as an adaptive radiation at an early evolutionary stage that has been obscured by more recent joint habitat diversification and neutral geographical processes. Diversification within Oxera has perhaps been triggered by varied ecological and biological drivers acting in a leapfrog pattern, but geographic processes may have been an equally important driver. We suspect that strictly adaptive radiations may be rare in plants and that most events of rapid clade diversification may have involved a mixture of geographical and ecological divergence.

Molecular data exclude current hybridization between iguanas Conolophus marthae and C. subcristatus on Wolf Volcano (Galápagos Islands)

Natural hybridization may influence population fitness and responsiveness to natural selection, in particular in oceanic island systems. In previous studies, interspecific hybridization was detected between the Galapagos iguana species Amblyrhynchus cristatus and Conolophus subcristatus. Further, possible hybridization was also suggested to occur between C. subcristatus and C. marthae at Wolf Volcano on Isabela Island. In this work, we investigated the level of hybridization between C. subcristatus and C. marthae using a large set of microsatellite markers. Results indicated strong differentiation between species and, while we cannot rule out hybridization in the past, there is no evidence of ongoing hybridization between C. marthae and C. subcristatus. These findings have great importance for the design of management actions and conservation plans, in particular for the purposes of a head start program. However, because potential for hybridization may change under different environmental and demographic conditions, genetic characterization of newly marked individuals of C. marthae and C. subcristatus in Wolf Volcano should not be interrupted.

Development of 26 microsatellite markers in Bupleurum latissimum (Apiaceae), an endangered plant endemic to Ulleung Island, Korea.

Premise of the StudyTo enhance our understanding of evolutionary consequences and to establish a suitable conservation strategy, we isolated microsatellite markers for the endangered Bupleurum latissimum (Apiaceae), which is endemic to the oceanic Ulleung Island. We also attempted cross‐amplification in B. euphorbioides and B. longeradiatum to investigate its continental progenitors.Methods and ResultsUsing high‐throughput sequencing data, we developed 26 polymorphic microsatellite loci in three multiplexes, of which 13 loci were polymorphic in the two related species. For B. latissimum, alleles numbered two to four and the observed and expected heterozygosity ranged from 0.000 to 0.500 and 0.061 to 0.529, respectively.ConclusionsThese developed markers will be useful for understanding evolutionary patterns of B. latissimum in an oceanic island system and for establishing suitable conservation strategies at the genetic level.

Diversity and tectonics: predictions from neutral theory

AbstractNumerical simulations of neutral metacommunities are used here to predict the effects of growth and shrinkage of metacommunities, as well as their separation and merging caused by continental collision and rifting and their secondary eustatic effects. Although growth and shrinkage of metacommunities predictably change diversity, separating and merging metacommunities have counterintuitive effects. Separating and merging metacommunities change diversity within the individual areas, especially so for smaller areas, but they cause no change in total diversity of the system, contrary to previous predictions. The response times of metacommunities are likely to be geologically undetectable except for enormously large systems. These models can be used to predict the plate-tectonic effects on the diversity of terrestrial, coastal-marine, deep-marine, and oceanic-island systems. Of these, global and regional coastal-marine systems are the most acutely sensitive to the changes in area and fragmentation caused by plate tectonics. Oceanic-island systems also experience global and regional changes in diversity during supercontinent breakup and assembly, with the global effects driven by the changing length of volcanic arcs, and the regional effects also driven by secondary eustatic changes in shallow-marine area. Although individual terrestrial provinces or continents may experience substantial changes in diversity from rifting and collision, global terrestrial diversity should be unchanged except for the relatively modest contributions caused by the secondary eustatic effects on land area. These changes in diversity may be reinforced or counteracted by the changing latitudinal position of metacommunities.

Oceanic island biogeography through the lens of the general dynamic model: assessment and prospect.

The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non-equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDM's predictions and in developing and enhancing ecological-evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: (i) macroecological tests using a space-for-time rationale; (ii) extensions of theory to islands following different patterns of ontogeny; (iii) the implications of GDM dynamics for lineage diversification and trait evolution; and (iv) the potential for downscaling GDM dynamics to local-scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non-native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research.

Opportunities and strategies to incorporate ecosystem services knowledge and decision support tools into planning and decision making in Hawai'i.

Incorporating ecosystem services into management decisions is a promising means to link conservation and human well-being. Nonetheless, planning and management in Hawai'i, a state with highly valued natural capital, has yet to broadly utilize an ecosystem service approach. We conducted a stakeholder assessment, based on semi-structured interviews, with terrestrial (n = 26) and marine (n = 27) natural resource managers across the State of Hawai'i to understand the current use of ecosystem services (ES) knowledge and decision support tools and whether, how, and under what contexts, further development would potentially be useful. We found that ES knowledge and tools customized to Hawai'i could be useful for communication and outreach, justifying management decisions, and spatial planning. Greater incorporation of this approach is clearly desired and has a strong potential to contribute to more sustainable decision making and planning in Hawai'i and other oceanic island systems. However, the unique biophysical, socio-economic, and cultural context of Hawai'i, and other island systems, will require substantial adaptation of existing ES tools. Based on our findings, we identified four key opportunities for the use of ES knowledge and tools in Hawai'i: (1) linking native forest protection to watershed health; (2) supporting sustainable agriculture; (3) facilitating ridge-to-reef management; and (4) supporting statewide terrestrial and marine spatial planning. Given the interest expressed by natural resource managers, we envision broad adoption of ES knowledge and decision support tools if knowledge and tools are tailored to the Hawaiian context and coupled with adequate outreach and training.

Introduction to Hydrology of Volcanic Islands

Among the volcanic regions in Spain, the most important are the Canary Islands. A remarkable amount of geological, ecological, botanical and hydrological singularities are concentrated in the archipelago. From the point of view of water resources, the Islands are a worldwide example of their use and management. This knowledge, developed trough years of research and experience in the Canary Islands, can be transferred to other oceanic island systems. In this paper, we start introducing the formation processes and geological oddities that condition water uses in the volcanic islands, with a special spot for the underground resources. Afterwards, waterworks more characteristic of islands and the effect of “fog rainfall” are described. Finally, we aim to the transfer of technology and success cases on water resources between global volcanic islands is proposed.

Enemy release and plant invasion: patterns of defensive traits and leaf damage in Hawaii

Invasive species may be released from consumption by their native herbivores in novel habitats and thereby experience higher fitness relative to native species. However, few studies have examined release from herbivory as a mechanism of invasion in oceanic island systems, which have experienced particularly high loss of native species due to the invasion of non-native animal and plant species. We surveyed putative defensive traits and leaf damage rates in 19 pairs of taxonomically related invasive and native species in Hawaii, representing a broad taxonomic diversity. Leaf damage by insects and pathogens was monitored in both wet and dry seasons. We found that native species had higher leaf damage rates than invasive species, but only during the dry season. However, damage rates across native and invasive species averaged only 2% of leaf area. Native species generally displayed high levels of structural defense (leaf toughness and leaf thickness, but not leaf trichome density) while native and invasive species displayed similar levels of chemical defenses (total phenolics). A defense index, which integrated all putative defense traits, was significantly higher for native species, suggesting that native species may allocate fewer resources to growth and reproduction than do invasive species. Thus, our data support the idea that invasive species allocate fewer resources to defense traits, allowing them to outperform native species through increased growth and reproduction. While strong impacts of herbivores on invasion are not supported by the low damage rates we observed on mature plants, population-level studies that monitor how herbivores influence recruitment, mortality, and competitive outcomes are needed to accurately address how herbivores influence invasion in Hawaii.

Intraguild predation and mesopredator release effect on long-lived prey

Complete extirpation of a species can generate cascading effects throughout an ecosystem, yet are precisely the goal of island eradications of pest species. “Mesopredator release effect”, an asymmetrical special case of intraguild predation, has been hypothesised as a possible indirect effect from eradications, where superpredator removal can generate a mesopredator increase which may increase the impact on their shared prey. Theoretically this suggests that for intraguild predators, the superpredator may protect the shared prey from mesopredation, and removal of superpredators alone is not recommended. We create a model of long-lived age-structured shared prey and explore the non-equilibrium dynamics of this system. The superpredator can impact all prey life-stages (adult survival and reproductive success) whereas the smaller mesopredator can only impact early life-stages (reproductive success). This model is independently tested with data from a closed oceanic island system where eradication of introduced intraguild predators is possible for conservation of threatened birds. Mesopredator release only occurs in strongly top-down moderated (resource-abundant) systems. Even when mesopredator release can occur, the negative impact of more mesopredators is outweighed by the benefit of superpredator removal, allowing recovery of the prey population. Results are robust to 10% variation in model parameters. The consideration of age-structured prey contradicts previous theoretical results for mesopredator release effect and intraguild predation. Superpredator eradication is vital for population recovery of long-lived insular species. Nonetheless island conservation must retain a whole-ecosystem perspective given the complex trophic relationships among multiple species on islands.