Effects of invasive rats and burrowing seabirds on seeds and seedlings on New Zealand islands

Rats (Rattus rattus, Rattus norvegicus, Rattus exulans) are important invaders on islands. They alter vegetation indirectly by preying on burrowing seabirds. These seabirds affect vegetation through nutrient inputs from sea to land and physical disturbance through trampling and burrowing. Rats also directly affect vegetation though consumption of seeds and seedlings. Seedling communities on northern New Zealand islands differ in composition and densities among islands which have never been invaded by rats, are currently invaded by rats, or from which rats have been eradicated. We conducted experimental investigations to determine the mechanisms driving these patterns. When the physical disturbance of seabirds was removed, in soils collected from islands and inside exclosures, seedling densities increased with seabird burrow density. For example, seedling densities inside exclosures were 10 times greater than those outside. Thus the negative effects of seabirds on seedlings, by trampling and uprooting, overwhelm the potentially beneficial effects of high levels of seed germination, seedling emergence, and possibly seed production, which result from seed burial and nutrient additions. Potential seedling density was reduced on an island where rats were present, germination of seeds from soils of this island was approximately half that found on other islands, but on this island seedling density inside exclosures was 7 times the density outside. Although the total negative effects of seabirds and rats on seedling densities are similar (reduced seedling density), the differences in mechanisms and life stages affected result in very different filters on the plant community.     

Invasive rats and seabirds after 2,000 years of an unwanted coexistence on Mediterranean islands

In the Mediterranean, the survival of endemic long-lived seabirds despite the long-standing introduction of one of the most damaging alien predator, the ship rat (Rattus rattus), on most islands constitutes an amazing conservation paradox. A database gathering information on approximately 300 Western Mediterranean islands was analyzed through generalized linear models to identify the factors likely to influence ship rat presence and to account for how ship rat presence and island characteristics may have driven the presence and abundance of seabirds. Our review showed that few Mediterranean islands remain rat-free. At the regional scale, rat presence was only a limiting factor in the abundance of the smallest seabird, the storm petrel (Hydrobates pelagicus), while the distribution and abundance of the three shearwaters were more influenced by island characteristics. We hypothesized that the long-term persistence of these seabirds may have been facilitated by the various biogeographical contexts of Mediterranean islands, likely to provide intra-island refuges.     

Seabird modulations of isotopic nitrogen on islands

The transport of nutrients by migratory animals across ecosystem boundaries can significantly enrich recipient food webs, thereby shaping the ecosystems' structure and function. To illustrate the potential role of islands in enabling the transfer of matter across ecosystem boundaries to be gauged, we investigated the influence of seabirds on nitrogen input on islands. Basing our study on four widely differing islands in terms of their biogeography and ecological characteristics, sampled at different spatial and temporal intervals, we analyzed the nitrogen isotopic values of the main terrestrial ecosystem compartments (vascular plants, arthropods, lizards and rodents) and their relationship to seabird values. For each island, the isotopic values of the ecosystem were driven by those of seabirds, which ultimately corresponded to changes in their marine prey. First, terrestrial compartments sampled within seabird colonies were the most enriched in δ(15)N compared with those collected at various distances outside colonies. Second, isotopic values of the whole terrestrial ecosystems changed over time, reflecting the values of seabirds and their prey, showing a fast turnover throughout the ecosystems. Our results demonstrate that seabird-derived nutrients not only spread across the terrestrial ecosystems and trophic webs, but also modulate their isotopic values locally and temporally on these islands. The wealth of experimental possibilities in insular ecosystems justifies greater use of these model systems to further our understanding of the modalities of trans-boundary nutrient transfers.     

Direct and indirect effects of rats: does rat eradication restore ecosystem functioning of New Zealand seabird islands?

Introduced rats (Rattus spp.) can affect island vegetation structure and ecosystem functioning, both directly and indirectly (through the reduction of seabird populations). The extent to which structure and function of islands where rats have been eradicated will converge on uninvaded islands remains unclear. We compared three groups of islands in New Zealand: islands never invaded by rats, islands with rats, and islands on which rats have been controlled. Differences between island groups in soil and leaf chemistry and leaf production were largely explained by burrow densities. Community structure of woody seedlings differed by rat history and burrow density. Plots on islands with high seabird densities had the most non-native plant species. Since most impacts of rats were mediated through seabird density, the removal of rats without seabird recolonization is unlikely to result in a reversal of these processes. Even if seabirds return, a novel plant community may emerge.     

Predation of seabirds by invasive rats: multiple indirect consequences for invertebrate communities

Invasive species are a global problem but most studies have focused on their direct rather than indirect ecological effects. We studied litter and soil‐inhabiting invertebrate communities on 18 islands off northern New Zealand, to better understand the indirect ecological consequences of rat (Rattus) invasion. Nine islands host high densities of burrowing procellariid seabirds that transport large amounts of nutrients from the ocean to the land. The other nine have been invaded over the past 50–150 years by rat species that have severely reduced the density of seabirds by preying on eggs and chicks. Invaded islands had lower densities of seabird burrows but deeper forest litter than did the uninvaded islands, indicative of rats reducing disturbance effects of seabirds. However, despite deeper litter on the invaded islands, eight of the 19 orders of invertebrates that we measured were significantly less abundant on invaded islands. Furthermore, three soil‐inhabiting micro‐invertebrate groups that we measured were significantly less abundant on invaded islands. These differences probably result from rats thwarting transfer of resources by seabirds from the ocean to the land. We also investigated community‐level properties of each of three test groups of invertebrates (minute land snails, spiders and soil nematodes) to illustrate this process. Spiders were equally abundant on both groups of islands, but showed lower species richness on the invaded islands. The other two groups showed no difference in species richness with island invasion status, but were more abundant on uninvaded islands. Reduced abundance of soil nematodes on invaded islands provides strong evidence of indirect consequences of seabird reduction by rats, because nematodes are unavailable to rats as prey. We predict that if rats are eradicated from islands, components of below‐ground invertebrate dependent on seabird‐mediated soil conditions may take considerable time to recover because they require subsequent seabird recolonisation.     

A GIS‐based decision‐making approach for prioritizing seabird management following predator eradication

Given that 29% of seabird species are threatened with extinction, protecting seabird colonies on offshore islands is a global conservation priority. Seabirds are vulnerable to non‐native predator invasions, which reduce or eliminate colonies. Accordingly, conservation efforts have focused on predator eradication. However, affected populations are often left to passively recover following eradications. Although seabirds are highly mobile, their life history traits such as philopatry can limit passive recolonization of newly predator‐free habitat. In such cases, seabird colonies can potentially be re‐instated with active restoration via chick translocations or social attraction methods, which can be risky and expensive. We used biogeographic and species‐specific behavioral data in the Hauraki Gulf, New Zealand, a global hotspot of seabird diversity and predator eradications, to illustrate the use of geographic information systems multi‐criteria decision analysis to prioritize islands for active seabird restoration. We identified nine islands with low observed passive recovery of seabirds posteradication over a 50‐year timeframe, and classified these as sites where active seabird management could be prioritized. Such spatially explicit tools are flexible, allowing for managers to choose case‐specific criteria such as time, funding, and goals constrained for their conservation needs. Furthermore, this flexibility can also be applied to threatened species management by customizing the decision criteria for individual species' capacity to passively recolonize islands. On islands with complex restoration challenges, decision tools that help island restoration practitioners decide whether active seabird management should be paired with eradication can optimize restoration outcomes and ecosystem recovery.     

Integrating seabird restoration and mammal eradication programs on islands to maximize conservation gains

Colonial nesting seabirds frequently drive island ecosystem biodiversity by maintaining ecosystem functioning and community dynamics. Invasive mammal introductions to most of the world’s islands have ravaged insular seabird populations and had associated devastating ecosystem-wide effects. Eradication programs remove invasive mammals from islands, with the goal of conserving and restoring island species and systems. However, most eradication programs rely almost exclusively on passive seabird recovery to achieve these goals. Unfortunately, the life histories of most seabird species are not conducive to passive recovery within a contemporary timeframe. Seabird restoration techniques can effectively overcome life history related issues and significantly reduce recovery times for insular seabird populations, thereby reducing associated ecosystem-wide recovery times. By integrating seabird restoration and eradication programs, practitioners can maximize conservation gains, expand funding opportunities, and restore island ecosystems and the biodiversity they support.     

Measuring nocturnal seabird activity and status using acoustic recording devices: applications for island restoration

ABSTRACT Nocturnal burrow‐nesting seabirds breeding on isolated oceanic islands pose challenges to conventional monitoring techniques, resulting in their frequent exclusion from population studies. These seabirds have been devastated by nonnative predator introductions on islands worldwide. After predators are eradicated, recovery has been poorly quantified, but evidence suggests some nocturnal seabird populations have been slow to return. We evaluated the use of automated acoustic recorders and call‐recognition software to investigate nocturnal seabird recovery after removal of introduced Arctic foxes (Alopex lagopus) in the Aleutian Archipelago, Alaska. We compared relative seabird abundance among islands by examining levels of vocal activity. We deployed acoustic recorders on Nizki‐Alaid, Amatignak, and Little Sitkin islands that had foxes removed in 1975, 1991, and 2000, respectively, and on Buldir, a predator‐free seabird colony. Despite frequent gales, only 2.9% of 2230 recording hours from May to August of 2008 and 2009 were unusable due to wind noise. Recording quality and call recognition model success were highest when recording devices were placed at sites offering some wind shelter. We detected greater vocal activity of Fork‐tailed (Oceanodroma furcata) and Leach's (O. leucorhoa) storm‐petrels and Ancient Murrelets (Synthliboramphus antiquus) on islands with longer time periods since fox eradication. Also, by detecting chick calls in the automated recordings, we confirmed breeding by Ancient Murrelets on an island thought to be abandoned due to fox predation. Acoustic monitoring allowed us to examine the relative abundance of seabirds at remote sites. If a link between vocalizations and population dynamics can be made, acoustic monitoring could be a powerful census method.     

Seabirds as adhesive seed dispersers of alien and native plants in the oceanic Ogasawara Islands, Japan

Previous studies have shown that the dispersal of plant seeds to oceanic islands is largely attributable to birds. However, few studies have assessed the role of adhesive dispersal by birds even though this mechanism has long been recognized as a major vector of seed transport. Some data point to the possibility that adhesive transport by seabirds transfers alien plant seeds in island ecosystems. In the present study, we examined the seed-dispersing ability of seabirds among islands in the oceanic Ogasawara Islands, Japan. We used capture surveys to examine the frequency of seeds adhering to seabirds and tested the salt tolerances of the seeds. The distributions of the plant species were examined and the relationships between plant and seabird distributions were analyzed using generalized linear models. Seeds of nine plant species, including aliens, were detected on 16–32?% of captured seabirds. Seeds included those generally considered to be dispersed by wind or internally transported by birds in their guts. Seeds exposed to NaCl solution isotonic with seawater for up to 8?h suffered little or no loss of viability. Analyses of plant distributions demonstrated positive relationships between the distributions of some plants and seabirds. These results show that seabirds effectively disperse seeds of both native and introduced plant species. This is the first study to comprehensively assess adhesive seed dispersal by seabirds; it provides essential information on long-distance dispersal.     

Long-term effects of feral goats (<Emphasis Type="Italic">Capra hircus</Emphasis>) on Mediterranean island communities: results from whole island manipulations

Islands exhibit disproportionally high biodiversity, however high levels of endemism and simplified food webs make their communities susceptible to invasive species. Introduced goats (Capra hircus), a generalist herbivore, are among the most harmful invasive species on islands. Concern about goat impacts on island communities have resulted in eradication programs, which have been generally implemented without comprehensive evaluation and monitoring. Unintended consequences may follow eradication, as grazing can have complex effects on island food webs. Using whole island manipulations, we evaluate the long-term, community-wide effects of goat herbivory, as well as their subsequent removal, in a system of 16 islands in the Aegean Sea (Greece) located within the Mediterranean biodiversity hotspot. Goat grazing on these islands is a major conservation concern, as these support endemic plant communities that have evolved in low herbivory conditions and lack appropriate defenses. We show that goat introductions lead to significant decreases in vegetation height, percent cover, and biomass but not to immediate plant species loss, as native island endemics are replaced with widespread generalist taxa carried in by the livestock. Additionally, goats contribute to the desertification of islands by initiating a long-term soil loss cycle that continues even after goats are removed; however, remaining soil structure and chemistry are not affected. Island arthropod populations do not appear to be significantly impacted by goat introduction or removal, except for a distinct increase in the order Diptera with goat presence. This study also reaffirms the role of seabirds in providing important marine subsidies, rich in nitrogen and phosphorus, to island food webs. Plant species diversity declines following goat removal, and vegetation cover returns only partially, as further recovery is being prevented by the long-term loss of soil. This suggests that following goat removal, island communities may require additional restoration efforts, including seabird reestablishment and reintroduction of extirpated plant populations, to promote island recovery.     

Climate oscillation and the invasion of alien species influence the oceanic distribution of seabirds

Spatial and temporal distribution of seabird transiting and foraging at sea is an important consideration for marine conservation planning. Using at‐sea observations of seabirds (n = 317), collected during the breeding season from 2012 to 2016, we built boosted regression tree (BRT) models to identify relationships between numerically dominant seabird species (red‐footed booby, brown noddy, white tern, and wedge‐tailed shearwater), geomorphology, oceanographic variability, and climate oscillation in the Chagos Archipelago. We documented positive relationships between red‐footed booby and wedge‐tailed shearwater abundance with the strength in the Indian Ocean Dipole, as represented by the Dipole Mode Index (6.7% and 23.7% contribution, respectively). The abundance of red‐footed boobies, brown noddies, and white terns declined abruptly with greater distance to island (17.6%, 34.1%, and 41.1% contribution, respectively). We further quantified the effects of proximity to rat‐free and rat‐invaded islands on seabird distribution at sea and identified breaking point distribution thresholds. We detected areas of increased abundance at sea and habitat use‐age under a scenario where rats are eradicated from invaded nearby islands and recolonized by seabirds. Following rat eradication, abundance at sea of red‐footed booby, brown noddy, and white terns increased by 14%, 17%, and 3%, respectively, with no important increase detected for shearwaters. Our results have implication for seabird conservation and island restoration. Climate oscillations may cause shifts in seabird distribution, possibly through changes in regional productivity and prey distribution. Invasive species eradications and subsequent island recolonization can lead to greater access for seabirds to areas at sea, due to increased foraging or transiting through, potentially leading to distribution gains and increased competition. Our approach predicting distribution after successful eradications enables anticipatory threat mitigation in these areas, minimizing competition between colonies and thereby maximizing the risk of success and the conservation impact of eradication programs.     

Island shape,size, and isolation affect nest‐site selection by Little Terns

ABSTRACT Populations of many seabirds and other species that nest along coasts are declining due to habitat degradation and loss. An improved understanding of the species‐specific factors that determine nest density across a landscape is therefore critical for conservation efforts. We examined factors that affected the density (number per hectare) and abundance (number at a sampling site) of nests of Little Terns (Sternula albifrons) on the Sinai Peninsula, Egypt. Terns preferred to nest on islands rather than the mainland, with islands constituting 64% of the area surveyed, but containing 99% of the 439 tern nests we found. Nest densities were highest on islands that were small, located at moderate distances from the mainland, and irregularly shaped or elongated. Most nests (69%) were on islands with areas < 3 ha, although these islands represented < 5% of total island area, and islands with the highest nest densities were 80–300 m from the mainland. Terrestrial predators were more likely to occur on larger islands, visiting three of the largest four islands. Most tern nests were within 1 m of shorelines, causing island perimeter to be a strong influence on nest density. Island shape was the only factor that significantly affected nest abundance, with more nests on islands with relatively long perimeters for their size. Our results suggest that protection or creation of relatively small, slender islands at moderate distances from shore may be an effective means of increasing the number of breeding sites for Little Terns. Although not generally considered a potential determinant of nest site preferences for seabirds, island shape is likely to be important for species that prefer sites adjacent to water, including species that nest on beaches and seaside cliffs.     

Invasive rats alter woody seedling composition on seabird-dominated islands in New Zealand

Invasive rats (Rattus rattus, R. norvegicus, R. exulans) have large impacts on island habitats through both direct and indirect effects on plants. Rats affect vegetation by extirpating burrowing seabirds through consumption of eggs, chicks, and adults. These seabirds serve as ecosystem engineers, affecting plant communities by burying and trampling seeds and seedlings, and by altering microclimate. Rats also directly affect plant communities by consuming seeds and seedlings. We studied the direct and indirect impacts of rats on the seedlings of woody plants on 21 islands in northern New Zealand. We compared seedling densities and richness on islands which differed in status with respect to rats: nine islands where rats never invaded, seven islands where rats were present at the time of our study, and five islands where rats were either eradicated or where populations were likely to be small as a result of repeated eradications and re-invasions. In addition, we compared plots from a subset of the 21 islands with different burrow densities to examine the effects of burrowing seabirds on plants while controlling for other factors that differ between islands. We categorized plant communities by species composition and seedling density in a cluster analysis. We found that burrow densities explained more variation in seedling communities than rat status. In areas with high seabird burrow density seedling densities were low, especially for the smallest seedlings. Species richness and diversity of seedlings, but not seedling density, were most influenced by changes in microclimate induced by seabirds. Islands where rats had been eradicated or that had low rat populations had the lowest diversity and richness of seedlings (and adults), but the highest seedling density. Seedling communities on these islands were dominated by Pseudopanax lessonii and Coprosma macrocarpa. This indicates lasting effects of rats that may prevent islands from returning to pre-invasion states.     

Seabird predation by great skuas Stercorarius skua– intra‐specific competition for food?

Competition for food is widely cited as an important cost of coloniality among birds and much of the evidence in support of this hypothesis comes from studies of colonial piscivorous seabirds. However, for generalist seabirds able to switch between different prey types, the role of food availability in relation to colony size is unclear. Here we investigate patterns of the consumption of seabird prey in relation to colony size in a generalist seabird, the great skua Stercorarius skua, in Shetland, UK. At the population level skuas feed mainly on sandeels Ammodytes marinus and fishery discards, but respond to declines in fish availability to facultatively prey on other seabirds. By comparing the consumption of seabirds among seven different sized colonies, including one colony with artificially reduced numbers of skuas (Fair Isle), we investigate whether consumption of seabird prey is influenced by skua population size, while simultaneously measuring seabird prey availability. Data from five years also enables us to investigate the influence of annual variation in environmental conditions on seabird consumption. Using measures of body condition and reproductive performance we investigate the consequences of living in different sized colonies, which may provide insight into ultimate costs of nesting at high population density. Skua diets varied among colonies and the proportion of seabird prey in the diet was inversely related to skua colony size, despite similar per capita numbers of seabirds across colonies. At the colony where their numbers were artificially suppressed, skuas consumed a greater proportion of seabirds per capita. Highly significant year effects in seabird predation were observed but the pattern among colonies remained consistent over time. Two measures of adult body condition (pectoral muscle index and mean corpuscular volume) revealed that adult great skuas were in poorer condition at the largest colony (Foula), but reproductive performance did not alter significantly among colonies. This study provides evidence that intra‐specific competition among skuas may limit opportunities for obtaining seabird prey, which may be particularly important during periods of poor availability of sandeels and fishery discards, and has implications for assessing the impact of skuas on seabird populations.     

Seabird nutrient subsidies alter patterns of algal abundance and fish biomass on coral reefs following a bleaching event

Cross‐ecosystem nutrient subsidies play a key role in the structure and dynamics of recipient communities, but human activities are disrupting these links. Because nutrient subsidies may also enhance community stability, the effects of losing these inputs may be exacerbated in the face of increasing climate‐related disturbances. Nutrients from seabirds nesting on oceanic islands enhance the productivity and functioning of adjacent coral reefs, but it is unknown whether these subsidies affect the response of coral reefs to mass bleaching events or whether the benefits of these nutrients persist following bleaching. To answer these questions, we surveyed benthic organisms and fishes around islands with seabirds and nearby islands without seabirds due to the presence of invasive rats. Surveys were conducted in the Chagos Archipelago, Indian Ocean, immediately before the 2015–2016 mass bleaching event and, in 2018, two years following the bleaching event. Regardless of the presence of seabirds, relative coral cover declined by 32%. However, there was a post‐bleaching shift in benthic community structure around islands with seabirds, which did not occur around islands with invasive rats, characterized by increases in two types of calcareous algae (crustose coralline algae [CCA] and Halimeda spp.). All feeding groups of fishes were positively affected by seabirds, but only herbivores and piscivores were unaffected by the bleaching event and sustained the greatest difference in biomass between islands with seabirds versus those with invasive rats. By contrast, corallivores and planktivores, both of which are coral‐dependent, experienced the greatest losses following bleaching. Even though seabird nutrients did not enhance community‐wide resistance to bleaching, they may still promote recovery of these reefs through their positive influence on CCA and herbivorous fishes. More broadly, the maintenance of nutrient subsidies, via strategies including eradication of invasive predators, may be important in shaping the response of ecological communities to global climate change.     

Marine resources subsidize insular rodent populations in the Gulf of California,Mexico

Inputs of energy and nutrients from one ecosystem may subsidize consumers in adjacent ones, with significant consequences for local communities and food webs. We used stable isotope and faecal pellet analysis to quantify use of ocean-derived resources by small mammals on islands in the Gulf of California, Mexico. Rodents were live-trapped on grids originating near shore and extending 125-200 m inland to evaluate the extent to which rodents transport marine nutrients inland, and to determine whether marine foods subsidize island populations, permitting higher densities than would be possible based on terrestrial resources alone. Both faeces and stable carbon and nitrogen isotopes revealed that omnivorous mice (Peromyscus maniculatus) consume ocean-derived prey, including littoral and supralittoral invertebrates, and that their diets differed markedly from those of granivorous rodents (Chaetodipus rudinoris). On a small, seabird roosting island, marine prey were important in the diet of mice regardless of their proximity to shore, underscoring the pervasive influence of the ocean on small islands with relatively large coastline area. On a large island, however, consumption of marine foods declined sharply > or =50 m from shore, which suggests that mice are poor conduits of inland movement of energy and nutrients from the sea. Marine resources seemed to act as subsidies for omnivorous rodents: more P. maniculatus were captured near shore than farther inland and there was an inverse relationship between island area and rodent abundance, suggesting that small islands with large amounts of marine inputs support the highest population densities. Patterns of local and island-wide abundance of P. maniculatus are likely the result of several interacting factors, including frustrated dispersal, competition with C. rudinoris, and the absence of predators. We speculate, however, that the availability of marine resources allows P. maniculatus to reach high densities and to persist on small islands in the Gulf despite low and unpredictable terrestrial productivity. Spatial trophic subsidies thus provide a possible mechanistic explanation for the widely reported inverse relationship between population density and island or habitat area.     

Isotopic analysis of island House Martins Delichon urbica indicates marine provenance of nutrients

The presence of one of the largest colonies of House Martins in Europe on the small island of Stora Karlsö, Sweden, led us to investigate the source of their food by analysis of stable isotopes of carbon and nitrogen. Carbon isotopic values of House Martin nestlings were the same as those of Common Guillemot Uria aalge nestlings fed on marine fish, but differed from local Collared Flycatcher Ficedula albicollis nestlings fed on woodland insects. We infer that these House Martins fed their chicks almost exclusively on insects that had used nutrients derived from seabirds, indicating a dependence on the presence of a large seabird colony. We suggest by extension that some populations of island passerines of high conservation importance may also be dependent on nutrient subsidies from seabird colonies.     

Promotion of ecosystem carbon sequestration by invasive predators

Despite recent interest in understanding the effects of human-induced global change on carbon (C) storage in terrestrial ecosystems, most studies have overlooked the influence of a major element of global change, namely biological invasions. We quantified ecosystem C storage, both above- and below-ground, on each of 18 islands off the coast of New Zealand. Some islands support high densities of nesting seabirds, while others have been invaded by predatory rats and host few seabirds. Our results show that, by preying upon seabirds, rats have indirectly enhanced C sequestration in live plant biomass by 104%, reduced C sequestration in non-living pools by 26% and increased total ecosystem C storage by 37%. Given the current worldwide distribution of rats and other invasive predatory mammals, and the consequent disappearance of seabird colonies, these predators may be important determinants of ecosystem C sequestration.     

Small island biogeography in the Gulf of California: lizards, the subsidized island biogeography hypothesis, and the small island effect

Aim We used insular lizard communities to test the predictions of two hypotheses that attempt to explain patterns of species richness on small islands. We first address the subsidized island biogeography (SIB) hypothesis, which predicts that spatial subsidies may cause insular species richness to deviate from species–area predictions, especially on small islands. Next, we examine the small island effect (SIE), which suggests small islands may not fit the traditional log‐linear species–area curve. Location Islands with arthropodivorous lizard communities throughout the Gulf of California. Methods To evaluate the SIB hypothesis, we first identified subsidized and unsubsidized islands based on surrogate measures of allochthonous productivity (i.e. island size and bird presence). Subsequently, we created species–area curves from previously published lizard species richness and island area data. We used the residuals and slopes from these analyses to compare species richness on subsidized and unsubsidized islands. To test for an SIE, we used breakpoint regression to model the relationship between lizard species richness and island area. We compared results from this model to results from the log‐linear regression model. Results Subsidized islands had a lower slope than unsubsidized islands, and the difference between these groups was significant when small islands were defined as < 1 km². In addition to comparing slopes, we tested for differences in the magnitude of the residuals (from the species–area regression of all islands) for subsidized vs. unsubsidized islands. We found no significant patterns in the residual values for small vs. large islands, or between islands with and without seabirds. The SIE was found to be a slightly better predictor of lizard species richness than the traditional log‐linear model. Main conclusions Predictions of the SIB hypothesis were partially supported by the data. The absence of a significant SIE may be a result of spatial subsidies as explained by the SIB hypothesis and data presented here. We conclude by suggesting potential scenarios to test for interactions between these two small island hypotheses. Future studies considering factors affecting species richness should examine the possible role of spatial subsidies, an SIE, or a synergistic effect of the two in data sets with small islands.