Determining variation in the success of New Zealand land birds

The New Zealand avifauna has changed considerably since the arrival of humans approximately 1000 years ago. The extinction of native species followed by their replacement with introduced exotic species is recognized as having greatly reduced the distinctiveness of the avifauna. Using phylogenetically independent methods, I tested whether life history and ecological correlates of extinction, introduction and persistence were significant across species and within taxa. Extinct land bird species were, on average, and independent of phylogenetic relatedness, larger-bodied than extant species. Introduction success was found to be significantly associated with increasing species body size, generation time and indices of human effort. In contrast, the geographical distribution of extant land birds was correlated with traits associated with high population growth rates (i.e. small, rapidly developing species, with high fecundity). My results suggest that selection pressures have changed significantly since the arrival of human colonists in New Zealand and that these changes have favoured different types of land birds, to the general detriment of endemic species. Comparative methods that explicitly examine historical changes are necessary to elucidate the changing roles of biological and extrinsic traits in the differential success of species in any assemblage.     

Is phylogenetic relatedness to native species important for the establishment of reptiles introduced to California and Florida?

Aim Charles Darwin posited that introduced species with close relatives were less likely to succeed because of fiercer competition resulting from their similarity to residents. There is much debate about the generality of this rule, and recent studies on plant and fish introductions have been inconclusive. Information on phylogenetic relatedness is potentially valuable for explaining invasion outcomes and could form part of screening protocols for minimizing future invasions. We provide the first test of this hypothesis for terrestrial vertebrates using two new molecular phylogenies for native and introduced reptiles for two regions with the best data on introduction histories. Location California and Florida, USA. Methods We performed an ordination of ecological traits to confirm that ecologically similar species are indeed closely related phylogenetically. We then inferred molecular phylogenies for introduced and native reptiles using sequence data for two nuclear and three mitochondrial genes. Using these phylogenies, we computed two distance metrics: the mean phylogenetic distance (MPD) between each introduced species and all native species in each region (which indicates the potential interactions between introduced species and all native species in the community) and the distance of each introduced species to its nearest native relative – NN (indicating the degree of similarity and associated likelihood of competition between each introduced species and its closest evolutionary analogue). These metrics were compared for introduced species that established and those that failed. Results We demonstrate that phylogenetically related species do share similar ecological functions. Furthermore, successfully introduced species are more distantly related to natives (for NN and MPD) than failed species, although variation is high. Main conclusions The evolutionary history of a region has value for explaining and predicting the outcome of human‐driven introductions of reptiles. Phylogenetic metrics are thus useful inputs to multi‐factor risk assessments, which are increasingly required for screening introduced species.     

The role of competition and introduction effort in the success of passeriform birds introduced to New Zealand

Abstract The finding that passeriform birds introduced to the islands of Hawaii and Saint Helena were more likely to successfully invade when fewer other introduced species were present has been interpreted as strong support for the hypothesis that interspecific competition influences invasion success. I tested whether invasions were more likely to succeed when fewer species were present using the records of passeriform birds introduced to four acclimatization districts in New Zealand. I also tested whether introduction effort, measured as the number of introductions and the total number of birds released, could predict invasion outcomes, a result previously established for all birds introduced to New Zealand. I found patterns consistent with both competition and introduction effort as explanations for invasion success. However, data supporting the two explanations were confounded such that the greater success of invaders arriving when fewer other species were present could have been due to a causal relationship between invasion success and introduction effort. Hence, without data on introduction effort, previous studies may have overestimated the degree to which the number of potential competitors could independently explain invasion outcomes and may therefore have overstated the importance of competition in structuring introduced avian assemblages. Furthermore, I suggest that a second pattern in avian invasion success previously attributed to competition, the morphological overdispersion of successful invaders, could also arise as an artifact of variation in introduction effort.     

Overestimation of establishment success of non-native birds in Hawaii and Britain

The tens rules states that 10 % of all introduced species establish and about 10 % of those species become invasive. Several studies have failed to support the tens rule. However, these studies are beset by a general weakness: many unsuccessful invasions are never reported, and without these data tests of the tens rules are inadequate. Here, using data on the establishment success of non-native birds in Hawaii and Britain and comparing these data with those from a previous study, we show that lack of information about failed species introductions, and the tendency to report species that have become invasive more than those that have not, result in an overestimate of the establishment success and invasion rates of non-native species.     

Propagule size and the relative success of exotic ungulate and bird introductions to New Zealand

We investigated factors affecting the success of 14 species of ungulates introduced to New Zealand around 1851-1926. The 11 successful species had a shorter maximum life span and were introduced in greater numbers than the three unsuccessful species. Because introduction effort was confounded with other life-history traits, we examined whether independent introductions of the same species were more likely to succeed when a greater number of individuals were introduced. For the six species with introductions that both succeeded and failed, successful introductions always involved an equal or greater number of individuals than unsuccessful introductions of the same species. For all independent introductions, there was a highly significant relationship between the number of individuals introduced and introduction success. When data for ungulate and bird introductions to New Zealand were combined, a variable categorizing species as ungulate or bird was a highly significant predictor of introduction success, after variation in introduction effort was controlled. For a given number of individuals introduced, ungulates were much more likely to succeed than birds.     

Single-queen introductions characterize regional and local invasions by the facultatively clonal little fire ant Wasmannia auropunctata

Clonal reproduction may facilitate the spread of invasive species by reducing the minimum population size necessary for successful establishment. We used microsatellite markers to reconstruct the composition of founding populations in two regional (Central Africa and Hawaii) and 23 local (near a Gabonese oilfield) invasions of the facultatively parthenogenetic little fire ant. Central Africa had a single dominant queen clone, which appears to have initiated the regional infestation, and then produced numerous other clones by rare sexual reproduction. This interpretation of the data was also supported by the genotype of a worker from the first collection in Africa (Gabon 1913). We found only a single queen clone in Hawaii, likewise indicating a single-clone introduction, most likely from an earlier infestation in Florida. Single-clone introductions also gave rise to the vast majority (92%) of local infestations at our oilfield study site. These results suggest the unusual, largely clonal, reproductive strategy of the little fire ant may enhance its success as an invasive species. However, the occasional sexual production of novel genotypes after the initial introduction may provide genetic flexibility that overcomes shortcomings of pure clonality.     

Searching for phylogenetic pattern in biological invasions

It has been suggested that alien species with close indigenous relatives in the introduced range may have reduced chances of successful establishment and invasion (Darwin's naturalization hypothesis). Studies trying to test this have in fact been addressing four different hypotheses, and the same data can support some while rejecting others. In this paper, we argue that the phylogenetic pattern will change depending on the spatial and phylogenetic scales considered. Expectations and observations from invasion biology and the study of natural communities are that at the spatial scale relevant to competitive interactions, closely related species will be spatially separated, whereas at the regional scale, species in the same genera or families will tend to co-occur more often than by chance. We also argue that patterns in the relatedness of indigenous and naturalized plants are dependent on the continental/island setting, spatial occupancy levels, and on the group of organisms under scrutiny. Understanding how these factors create a phylogenetic pattern in invasions will help us predict which groups are more likely to invade where, and should contribute to general ecological theory.     

Life history and ecology influences establishment success of introduced land birds

Despite the many studies that have investigated successful establishment of introduced bird species, very little is known about the patterns of success worldwide and the influence of life history and ecological traits. This study describes the analysis of non-native land bird introductions to test existing hypotheses of establishment success using a modern comparative approach to control for phylogenetic relatedness among taxa. I used randomization tests, permutational phylogenetic regressions, and across-taxa and sister-taxa comparisons to examine predicted correlates of introduction success. My analyses confirmed that the variability in establishment success among introduced land bird families is distributed in a manner significantly different from a random process, and that life history and ecological attributes are an important influence of introduction success. I found strong evidence, through a generalized linear model, that increased habitat generalism, lack of migratory tendency, and sexual monochromatism together explain significant variation in the successful establishment of introduced land bird species. This has resulted in a predictive equation for the novel introduction of land bird species.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 465–480.     

Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest

Negative density dependence (NDD) and environmental filtering (EF) shape community assembly, but their relative importance is poorly understood. Recent studies have shown that seedling's mortality risk is positively related to the phylogenetic relatedness of neighbours. However, natural enemies, whose depredations often cause NDD, respond to functional traits of hosts rather than phylogenetic relatedness per se. To understand the roles of NDD and EF in community assembly, we assessed the effects on seedling mortality of functional similarity, phylogenetic relatedness and stem density of neighbouring seedlings and adults in a species-rich tropical forest. Mortality risks increased for common species when their functional traits departed substantially from the neighbourhood mean, and for all species when surrounded by close relatives. This indicates that NDD affects community assembly more broadly than does EF, and leads to the tentative conclusion that natural enemies respond to phylogenetically correlated traits. Our results affirm the prominence of NDD in structuring species-rich communities.     

It takes one to know one: Similarity to resident alien species increases establishment success of new invaders

Aim

Darwin's naturalization hypothesis states that dissimilarity to native species may benefit alien species establishment due to empty niches and reduced competition. We here add a new dimension to large‐scale tests of community invasibility, investigating the role that previously established alien species play in facilitating or hindering new invasions in plant communities.

Location

Permanent grasslands across France (including mainland and Corsica), as a receding ecosystem of great conservation importance.

Methods

Focusing on 121 alien plant species occurring in 7,215 vegetation plots, we quantified biotic similarity between new invaders and resident alien species (i.e., alien species with longer residence times) based on phylogenetic and trait distances. Additionally, we calculated distances to native species for each alien species and plot. Using multispecies distribution models, we analysed the influence of these biotic similarity measures and additional covariates on establishment success (presence/absence) of new invaders.

Results

We found that biotic similarity to resident alien species consistently increased establishment success of more recently introduced species. Phylogenetic relatedness to previous invaders had an equally strong positive effect as relatedness to native species. Conversely, trait similarity to natives hindered alien establishment as predicted by Darwin's naturalization hypothesis. These results highlight that various mechanisms may act simultaneously to determine alien establishment success.

Main conclusions

Our results suggest that, with greater similarity among alien species, invasion success increases. Such a pattern may arise either due to actual facilitation among invaders or as a result of weaker competitive interactions among invaders than between native and alien species, leading to an indirect facilitative effect. Alternatively, recent environmental changes (e.g., eutrophication, climate change) may have added new environmental filters. Determining how initial invasions might pave the road for subsequent invasions is crucial for effective multispecies management decisions and contributes a new aspect to our understanding of community assembly.

     

The role of life history traits in mammalian invasion success

Why some organisms become invasive when introduced into novel regions while others fail to even establish is a fundamental question in ecology. Barriers to success are expected to filter species at each stage along the invasion pathway. No study to date, however, has investigated how species traits associate with success from introduction to spread at a large spatial scale in any group. Using the largest data set of mammalian introductions at the global scale and recently developed phylogenetic comparative methods, we show that human‐mediated introductions considerably bias which species have the opportunity to become invasive, as highly productive mammals with longer reproductive lifespans are far more likely to be introduced. Subsequently, greater reproductive output and higher introduction effort are associated with success at both the establishment and spread stages. High productivity thus supports population growth and invasion success, with barriers at each invasion stage filtering species with progressively greater fecundity.     

Breeding system and pollination ecology of introduced plants compared to their native relatives

Identifying how plant-enemy interactions contribute to the success of introduced species has been a subject of much research, while the role of plant-pollinator interactions has received less attention. The ability to reproduce in new environments is essential for the successful establishment and spread of introduced species. Introduced plant species that are not capable of autonomous self-fertilization and are unable to attract resident pollinators may suffer from pollen limitation. Our study quantifies the degree of autogamy and pollination ecology of 10 closely related pairs of native and introduced plant species at a single site near St. Louis, Missouri, USA. Most of these species pairs had similar capacities for autogamy; however, of those that differed, the introduced species were more autogamous than their native congeners. Most introduced plants have pollinator visitation rates similar to those of their native congeners. Of the 20 species studied, only three had significant pollen limitation. We suggest that the success of most introduced plant species is because they are highly autogamous or because their pollinator visitation rates are similar to those of their native relatives. Understanding and identifying traits related to pollination success that are key in successful introductions may allow better understanding and prediction of biological invasions.     

A refined model of body mass and population density in flightless birds reconciles extreme bimodal population estimates for extinct moa

Flightless birds were once the largest and heaviest terrestrial fauna on many archipelagos around the world. Robust approaches for estimating their population parameters are essential for understanding prehistoric insular ecosystems and extinction processes. Body mass and population density are negatively related for extant flightless bird species, providing a method for quantifying densities and population sizes of extinct flightless species. Here we assemble an updated global data set of body mass and population densities for extant flightless birds and estimate the relationship between these variables. We use generalised least squares models that account for phylogenetic relatedness and incorporate the effects of limiting factors (e.g. habitat suitability) on population density. We demonstrate the applicability of this allometric relationship to extinct species by estimating densities for each of the nine species of moa (Dinornithiformes) and generating a combined spatially explicit map of total moa density across New Zealand. To compare our density estimates with those previously published, we summed individual species' abundances to generate a mean national density of 2.02–9.66 birds km⁻² for low- and high-density scenarios, respectively. Our results reconcile the extreme bimodality of previous estimates (< 2 birds km⁻² and > 10 birds km⁻²) and are comparable to contemporary densities of large herbivorous wild mammals introduced into New Zealand about 150 yr ago. The revised moa density has little effect on the harvest rates required to bring about extinction within 150–200 yr, indicating that rapid extinction was an inevitable response to human hunting, irrespective of the initial population of moa.     

Historical factors mediate the effects of interspecific competition

Biotic interactions, such as interspecific competition, are potentially important in determining whether introduced species succeed or fail to establish wild populations. Such effects may be difficult to detect, however, because the outcome of interspecific competition may depend on historical and largely unpredictable circumstances such as the timing of introductions and the number of individuals of each species introduced. I used a stochastic birth-death model to explore the effects of interspecific competition, the timing of introductions and the numbers of individuals of each species introduced, on invasion success in a two-species competitive system. I then compared the model predictions with actual data on establishment outcomes for passerine birds introduced to New Zealand, for which we have data on the timing of introductions, the size of release populations, and a measure of the strength of per capita competition (the degree of morphological similarity among species). The model and data agree well, suggesting that interspecific competition was an important determinant of invasion success in this assemblage, but that the outcome of competition depended critically on circumstances such as the timing of introductions and number of individuals released. Hence, while there is a deterministic component to invasion success in this assemblage (morphologically similar species are less likely to establish), historical circumstances played a critical role in mediating the outcomes.     

Competition and introduction regime shape exotic bird communities in Hawaii

Complex combinations of historical and local-regional processes determine the assembly of ecological communities. We investigated such processes in the Hawaiian introduced avifauna, comprising 140 years of historical records of invasions and extinctions of birds. Here the particular introduction regime (i.e., colonization attempts and number of introduced species) and priority effects constitute the historical (and regional) component, and competition is the local component. These processes are theoretically supported by means of a Lotka–Volterra model of species competition, finding that changes in the specific introduction regime might result in different extinction dynamics. Both field data and model outcomes support the biotic resistance hypothesis, so that the invasibility of new incomers decrease with species richness. Finally, we found that the resistance to new invaders depends on the particular introduction regime. Thus, community assembly models built to predict the success of exotic species should consider more scenarios than random introduction regimes.     

Analysis of importations for biological control of insect pests and weeds in New Zealand

Importations of biological control agents for insect pests and weeds in New Zealand are summarized and factors contributing to the relative success of the programmes are examined. The establishment rate of 30.9% is similar to that achieved worldwide, but is significantly lower than the rate achieved in the island habitat of Hawaii. The pioneering role of New Zealand in biological control is shown by the high proportion of programmes first attempted in this country. Although this novelty has not reduced the establishment rate, introductions against endemic species have not succeeded. Size of release was not a dominant feature in the establishment of agents. Complete or substantial success is recorded for 17 of the 70 target pests, with a relatively high success rate in forestry programmes. Examples of the influence of climate matching and competitive exclusion are also discussed. Changing practices and attitudes to the introduction of biological control agents are documented to show the increasing emphasis on specialists. No adverse effects of introductions are reported. The challenge to practitioners and regulators is to develop systems to evaluate conflicts of interest and develop workable mechanisms to determine which biological control agents are suitable for release.     

Phylogenetic patterns of colonization and extinction in experimentally assembled plant communities

Background

Evolutionary history has provided insights into the assembly and functioning of plant communities, yet patterns of phylogenetic community structure have largely been based on non-dynamic observations of natural communities. We examined phylogenetic patterns of natural colonization, extinction and biomass production in experimentally assembled communities.

Methodology/Principal Findings

We used plant community phylogenetic patterns two years after experimental diversity treatments (1, 2, 4, 8 or 32 species) were discontinued. We constructed a 5-gene molecular phylogeny and statistically compared relatedness of species that colonized or went extinct to remaining community members and patterns of aboveground productivity. Phylogenetic relatedness converged as species-poor plots were colonized and speciose plots experienced extinctions, but plots maintained more differences in composition than in phylogenetic diversity. Successful colonists tended to either be closely or distantly related to community residents. Extinctions did not exhibit any strong relatedness patterns. Finally, plots that increased in phylogenetic diversity also increased in community productivity, though this effect was inseparable from legume colonization, since these colonists tended to be phylogenetically distantly related.

Conclusions

We found that successful non-legume colonists were typically found where close relatives already existed in the sown community; in contrast, successful legume colonists (on their own long branch in the phylogeny) resulted in plots that were colonized by distant relatives. While extinctions exhibited no pattern with respect to relatedness to sown plotmates, extinction plus colonization resulted in communities that converged to similar phylogenetic diversity values, while maintaining differences in species composition.     

Nest predators and the evolution of avian reproductive strategies: a comparison of Australian and New Zealand birds

Summary Nest predation has been considered an important factor in the evolution of avian life histories: smaller clutches and shorter incubation and nestling periods are expected where nest predation has significant effects on reproductive success. Unlike the Australian avifauna, terrestrial New Zealand birds have evolved in the absence of reptilian and mammalian predators. Here we compare the reproductive strategies of terrestrial native New Zealand birds with those of their Australian sister taxa. In 11 of 14 comparisons, New Zealand birds were larger than their Australian relatives, but we did not find any significant differences in reproductive tactics between the two regions, a result inconsistent with the nest predation hypothesis. We discuss several reasons why this may be so. One possibility is that selection imposed on avian life history tactics by mammalian predators following the arrival of humans in New Zealand has led to strategies similar to those adopted in Australia.     

Vicars,tramps and assembly of the New Zealand avifauna: a review of molecular phylogenetic evidence

The avifauna of New Zealand is taxonomically and ecologically distinctive, as is typical of island biotas. However, the potential for an old geological age of New Zealand has encouraged a popular notion of a ‘Moa’s ark’ based on the idea that much of the fauna was isolated when Zealandia broke from Gondwana c. 83 million years ago. Molecular phylogenetics has proved useful for exploring the relative importance of different biogeographical processes, revealing for example that ‘tramp’ species (widely dispersing taxa) have arrived in New Zealand even in the last few hundred years, and that some avian taxa have close phylogenetic relatives overseas (predominantly Australian), indicating their recent ancestors were tramps, too. Distinctive taxa with deep phylogenetic ancestry might be ‘vicars’ that owe their presence to vicariance, but lack of close morphological, taxonomic and phylogenetic affinity provides only tenuous evidence for this. Disproving the alternative possibility that apparent vicars are descended from tramps that dispersed in earlier times remains challenging, but molecular analyses have yielded startling insights. Among New Zealand’s iconic taxa, the world’s largest eagle shared a Pleistocene ancestor with a small Australian eagle, and giant, flightless moa are phylogenetic sisters of the much smaller, flying tinamous of South America. The New Zealand avifauna is neither isolated nor stable, but demonstrative of prolonged and ongoing colonization, speciation and extinction.     

Predicting establishment success for alien reptiles and amphibians: a role for climate matching

We examined data comprising 1,028 successful and 967 failed introduction records for 596 species of alien reptiles and amphibians around the world to test for factors influencing establishment success. We found significant variations between families and between genera. The number of jurisdictions where a species was introduced was a significant predictor of the probability the species had established in at least one jurisdiction. All species that had been introduced to more than 10 jurisdictions (34 species) had established at least one alien population. We also conducted more detailed quantitative comparisons for successful (69 species) and failed (116 species) introductions to three jurisdictions (Great Britain, California and Florida) to test for associations with climate match, geographic range size, and history of establishment success elsewhere. Relative to failed species, successful species had better climate matches between the jurisdiction where they were introduced and their geographic range elsewhere in the world. Successful species were also more likely to have high establishment success rates elsewhere in the world. Cross-validations indicated our full model correctly categorized establishment success with 78–80% accuracy. Our findings may guide risk assessments for the import of live alien reptiles and amphibians to reduce the rate new species establish in the wild.