Is propagule size the critical factor in predicting introduction outcomes in passeriform birds?

Influential analyses of the propagule pressure hypothesis have been based on multiple bird species introduced to one region (e.g. New Zealand). These analyses implicitly assume that species-level and site-level characteristics are less important than the number of individuals released. In this study we compared records of passerine introductions with propagule size information across multiple regions (New Zealand, Australia, and North America). We excluded species introduced to just one of the three regions or with significant uncertainty in the historical record, as well as species that succeeded or failed in all regions. Because it is often impossible to attribute success to any single event or combination of events, our analysis compared randomly selected propagule sizes of unsuccessful introductions with those of successful introductions. Using Monte Carlo repeated sampling we found no statistical support for the propagule pressure hypothesis, even when using assumptions biased toward showing an effect.     

A reassessment of the role of propagule pressure in influencing fates of passerine introductions to New Zealand

Several studies have argued that principal factor in determining the fate of bird introductions is introduction effort. In large part, these studies have emerged from analyses of historical records from a single place—New Zealand. Here we raise two concerns about these conclusions. First, we argue that although many bird species were introduced repeatedly to New Zealand, in many cases the introductions apparently occurred only after the species were already successfully naturalized. The inclusion of such seemingly superfluous introductions may exaggerate the importance of propagule pressure. And second, we question the reliability of the records themselves. In many cases these records are equivocal, as inconsistencies appear in separate studies of the same records. Our analysis indicates that species were successful not because they were introduced frequently and in high numbers, but rather it is likely that they were introduced frequently and in high numbers because the initial releases were successful.     

A reassessment of historical records of avian introductions to Australia: no case for propagule pressure

Introduced species are widely believed to represent a significant threat to conservation of biological diversity. A better understanding of the ecological factors associated with successful species establishment should lead to improved management and mitigation of these introductions. The “propagule pressure hypothesis”, implying a greater chance of successful introduction with greater numbers introduced, has been widely accepted as a principal ecological factor in explaining establishment of exotic species. The historical record of bird introductions in a few locations, including the state of Victoria in Australia, has been advanced as the principal quantitative support for the hypothesis. We compiled lists of bird species introductions into Australia from several sources, and discovered inconsistencies in the records of introductions. In a series of comparisons, we found that the historical record of passerine introductions to Australia does not support the propagule pressure hypothesis unless superfluous introductions of already successful species are included. An additional problem with previous analyses is the inclusion of unsuccessful haphazard cage escapes.     

Establishment success in introduced passeriforms of New Zealand: evidence for a Franklin Delano Roosevelt effect

Blackburn et al. (Biol Invasions doi:10.1007/s10530-013-0451-x, 2013) have proposed a series of tests that are purported to evaluate the motivations and decisions of early acclimatization societies that introduced passeriform birds to New Zealand. We show that the data are not consistent with their predictions and, moreover, their predictions are consistent with our alternative hypothesis. Thus, species they identify as being apparently successful species were released in significantly higher numbers than those they claim as unequivocal failures; the apparently successful species were released significantly fewer times and in significantly lower numbers than those designated as unequivocally successful species; persistence time of unsuccessful species was significantly and positively correlated with number of releases; and among species released just a single time there was no difference in propagule sizes of unsuccessful versus successful species. Thus, despite their counter claims, the results of Blackburn et al. (Biol Invasions doi:10.1007/s10530-013-0451-x, 2013) clearly show that patterns in establishment are consistent with the historical record and what they have termed a “Franklin Delano Roosevelt Effect”.     

Propagule pressure as a driver of establishment success in deliberately introduced exotic species: fact or artefact?

A central paradigm in invasion biology is that more releases of higher numbers of individuals increase the likelihood that an exotic population successfully establishes and persists. Recently, however, it has been suggested that, in cases where the data are sourced from historical records of purposefully released species, the direction of causality is reversed, and that initial success leads to higher numbers being released. Here, we explore the implications of this alternative hypothesis, and derive six a priori predictions from it. We test these predictions using data on Acclimatization Society introductions of passerine bird species to New Zealand, which have previously been used to support both hypotheses for the direction of causality. All our predictions are falsified. This study reaffirms that the conventional paradigm in invasion biology is indeed the correct one for New Zealand passerine bird introductions, for which numbers released determine establishment success. Our predictions are not restricted to this fauna, however, and we keenly anticipate their application to other suitable datasets.     

The earliest House Sparrow introductions to North America

Several authors have argued that three separate introductions of roughly 100 individuals were required initially to establish the House Sparrow (Passer domesticus) in the Brooklyn, New York area. We argue that these claims are in error and that the actual record suggests that it is likely the initial introduction of just 16 birds in 1851 was all that was required to establish the species in New York. We further suggest that a similar level of scrutiny of historical records will reveal more examples of misinterpretations and errors, casting doubt on the validity of studies that claim propagule pressure has played an important role in determining the fate of bird introductions.     

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.     

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.     

A population model for predicting the successful establishment of introduced bird species

One of the strongest generalities in invasion biology is the positive relationship between probability of establishment and the numbers of individuals introduced. Nevertheless, a number of significant questions remain regarding: (1) the relative importance of different processes during introduction (e.g., demographic, environmental, and genetic stochasticity, and Allee effects); (2) the relative effects of propagule pressure (e.g., number of introductions, size of introductions, and lag between introductions); and (3) different life history characteristics of the species themselves. Here, we adopt an individual-based simulation modeling approach to explore a range of such details in the relationship between establishment success and numbers of individuals introduced. Our models are developed for typical exotic bird introductions, for which the relationship between probability of establishment and the numbers of individuals introduced has been particularly well documented. For both short-lived and long-lived species, probability of establishment decreased across multiple introductions (compared with a single introduction of the same total size), and this decrease was greater when inbreeding depression was included. Sensitivity analyses revealed four predictors that together accounted for >95 % of model performance. Of these, R ₀ (the average number of daughters produced per female over her lifetime) and propagule pressure were of primary importance, while random environmental effects and inbreeding depression exerted lesser influence. Initial founder size is undoubtedly going to be important for ensuring the persistence of introduced populations. However, we found the demographic traits, which influence how introduced individuals behave, to have the greatest effect on establishment success.     

Patterns of success in game bird introductions in the United States

Better predictions of the success of species’ introductions require careful evaluation of the relative importance of at least three kinds of factors: species characteristics, characteristics of the site of introduction, and event-level factors such as the numbers of individuals released. (Henceforth, we call this propagule pressure.) The 1644 introductions of 17 Phasianid species released in various US states during the Foreign Game Investigation Program provides a particularly rich source of data to test these ideas. An examination of these records indicates that 13 of these 17 species always failed, despite generally numerous individual releases and large numbers of individuals in each release. Moreover, ten of these species have been successfully introduced elsewhere. Only four of the 17 species were successful in at least one state. Some 20 sets of releases of three of these four species always failed in some states, again given generally numerous individual releases of large numbers of individuals in each release. Simply, the combination of site and species factors explain the lack of successes. This leaves a combination of 18 states where one or more of the four species succeeded. For these, there are significant differences in the numbers of birds introduced from state to state. But only for two species Alectoris chukar and; Tetraogallus himalayensis are there significant differences that show a greater chance of success when more individuals are introduced. These results support the conclusion that the number of individuals released, meaning propagule pressure, is not as important as characteristics of the species and the location to where its introduction occurred.     

Conclusions from a dynamic population model for tsetse: response to comments

The critique by Hargrove et al. (Popul Ecol, 2011) of our recently published paper on a tsetse population model (Barclay and Vreysen in Popul Ecol 53:89–110, 2011) has made some good points but has also misinterpreted the intent of some of our results as we presented them. Hargrove et al. rightly say that there is a mismatch between the size of the unit cells in the model (1 ha) and the iteration rate of the model (every 5 days), yielding too low a dispersal rate to simulate reality. However, they have misconstrued several of our results that we presented as examples to imply that those results were a necessary condition for control of tsetse, especially using traps and targets.     

Strength in size not numbers: propagule size more important than number in sexually reproducing populations

Propagule pressure has consistently been identified as a primary factor in invader success, and reducing it can be one of the most effective methods for preventing the establishment of non-native species. However, when policy is implemented to reduce propagule pressure it almost exclusively focuses on the size of individual introduction events (‘propagule size’), with little confirmation that controlling this single aspect of propagule pressure is the most effective strategy. The number of introduction events (‘propagule number’) can play as much, or more, of a role in invader success, yet only a small portion of propagule pressure research has studied the relative importance of size and number. We investigated the relative roles of propagule size and number in the establishment of a sexually reproducing species using a field mesocosm experiment that introduced Hemimysis anomala (a non-native mysid) across a range of propagule sizes and numbers. We found that single, large introductions had higher abundances and probabilities of survival than smaller, more frequent additions. This experiment illustrated that, for sexual reproducers, focusing on lowering propagule size can be the most effective method for reducing non-native establishment.     

Herbivory and plant competition reduce mountain beech seedling growth and establishment in New Zealand

The effect of herbivory and resource availability on the competitive ability of different plant species has been an area of intense debate amongst plant ecologists for at least two decades, but the interactive effects of herbivory and plant competition between woody and herbaceous plants are rarely studied and theory is poorly developed. This study used experimental manipulations on transplanted and naturally occurring mountain beech (Nothofagus solandri var. cliffortioides) seedlings to show the effects of deer browsing and competition from deer-induced, herbaceous turf communities on mountain beech regeneration in New Zealand. Differences in the species composition of these turfs had little effect on mountain beech seedling establishment, but turf removal increased seedling growth and survivorship, showing that competition with other plants had direct effects on mountain beech regeneration. Deer browsing reduced the establishment and growth of seedlings, but the size of this effect did not vary with light and nutrient availability. There was no immediate compositional response of turf communities following the removal of deer browsing. The addition of nutrients appeared to reduce the intensity of belowground competition (stem growth increased relative to root growth) and increase seedling mortality, but there was no effect of changing levels of light. These results showed simple and direct negative effects of deer browsing on mountain beech regeneration. Indirect negative effects on regeneration were caused by deer-induced turf communities. We found little evidence for interactive effects between herbivory, plant competition and the availability of light or nutrients on seedling regeneration, which suggests that these factors acted independently. Nomenclature: Beever et al. (1992); Parsons et al. (1995); Edgar and Connor (2000); and Brownsey and Smith-Dodsworth (2000). Raukaua simplex is described by Mitchell et al. (1997). Coprosma “taylorae” is referred to by Eagle (1986) and Halocarpus biformis, Phyllocladus alpinus, Podocarpus hallii and Podocarpus nivalis by Wilson and Galloway (1993).     

Crossing the Tasman Sea: Inferring the introduction history of Litoria aurea and Litoria raniformis (Anura: Hylidae) from Australia into New Zealand

Abstract The amphibian fauna of New Zealand consists of three native species (Leiopelma spp.), and three Litoria species introduced from Australia in the last 140 years. We conducted a molecular phylogeographical study that aimed to identify the Australian origins of two species, Litoria aurea and Litoria raniformis. We used partial sequences of the mitochondrial cytochrome oxidase I (cox1) gene from 59 specimens sampled from across the range of both species to identify the probable source populations for the New Zealand introductions, and to describe the current genetic diversity among New Zealand Litoria populations. Our genetic data suggest that L. aurea was introduced into the North Island of New Zealand from two regions in Australia, once from the northern part of coastal New South Wales and once from the southern part of coastal New South Wales. Our data indicate that L. raniformis introductions originated from the Melbourne region of southern Victoria and once established in the South Island of New Zealand, the species subsequently spread throughout both islands. In addition, we found a distinct haplotype in L. raniformis from Tasmania that strongly suggests, contrary to earlier reports, that this species was not introduced into New Zealand from Tasmania. Finally, we identified two very distinctive mitochondrial lineages of L. raniformis within the mainland Australia distribution, which may be previously unrecognized species.     

The synaptonemal complex and meiotic recombination in humans: new approaches to old questions

Meiotic prophase serves as an arena for the interplay of two important cellular activities, meiotic recombination and synapsis of homologous chromosomes. Synapsis is mediated by the synaptonemal complex (SC), originally characterized as a structure linked to pairing of meiotic chromosomes (Moses (1958) J Biophys Biochem Cytol 4:633–638). In 1975, the first electron micrographs of human pachytene stage SCs were presented (Moses et al. (1975) Science 187:363–365) and over the next 15 years the importance of the SC to normal meiotic progression in human males and females was established (Jhanwar and Chaganti (1980) Hum Genet 54:405–408; Pathak and Elder (1980) Hum Genet 54:171–175; Solari (1980) Chromosoma 81:315–337; Speed (1984) Hum Genet 66:176–180; Wallace and Hulten (1985) Ann Hum Genet 49(Pt 3):215–226). Further, these studies made it clear that abnormalities in the assembly or maintenance of the SC were an important contributor to human infertility (Chaganti et al. (1980) Am J Hum Genet 32:833–848; Vidal et al. (1982) Hum Genet 60:301–304; Bojko (1983) Carlsberg Res Commun 48:285–305; Bojko (1985) Carlsberg Res Commun 50:43–72; Templado et al. (1984) Hum Genet 67:162–165; Navarro et al. (1986) Hum Reprod 1:523–527; Garcia et al. (1989) Hum Genet 2:147–53). However, the utility of these early studies was limited by lack of information on the structural composition of the SC and the identity of other SC-associated proteins. Fortunately, studies of the past 15 years have gone a long way toward remedying this problem. In this minireview, we highlight the most important of these advances as they pertain to human meiosis, focusing on temporal aspects of SC assembly, the relationship between the SC and meiotic recombination, and the contribution of SC abnormalities to human infertility.The synaptonemal complex–50 years     

Assessing loss of coral cover on the Great Barrier Reef: A response to Hughes et al. (2011)

Hughes et al. (Coral Reefs, 2011, in press) challenge our interpretations of the changes in coral cover observed on the Great Barrier Reef (GBR) between 1986 and 2004 (Sweatman et al. in Coral Reefs 30:521–531, 2011). They question whether we can accurately assign all causes of coral loss; we contend that this makes no difference to the observed changes. They defend the validity of historical data on coral cover from before the start of systematic large-scale monitoring and conclude that coral cover has been declining since at least 1960, but we find no trend in the early data. We remain convinced that combining data collected at different spatial scales (quadrats and transects in the past mixed with more recent whole-reef averages from manta tows) are likely to overestimate decline, because whole-reef averages will very rarely reach the high cover values that can occur at the quadrat scale. Hughes et al. (Coral Reefs, 2011, in press) state that we dismiss runoff as a cause of ecosystem degradation; we defend our interpretations and dispute some of their examples. In summary, we stand by our conclusion that coral cover on the GBR declined in the period 1986–2004 but through localised and unsynchronised changes that included recovery.     

Popularity and Propagule Pressure: Determinants of Introduction and Establishment of Aquarium Fish

Propagule pressure is frequently cited as an important determinant of invasion success for terrestrial taxa, but its importance for aquatic species is unclear. Using data on aquarium fishes in stores and historical records of fish introduced and established in Canadian and United States waters, we show clear relationships exist between frequency of occurrence in shops and likelihood of introduction and of establishment. Introduced and established taxa are also typically larger than those available from stores, consistent with the propagule pressure hypothesis in that larger fish may be released more frequently due to outgrowing their aquaria. Attempts to reduce the numbers of introductions may be the most practical mechanism to reduce the number of new successful invasions.     

Temperature controls a latitudinal gradient in the proportion of watershed nitrogen exported to coastal ecosystems

Increased export of biologically available nitrogen (N) to the coastal zone is strongly linked to eutrophication, which is a major problem in coastal marine ecosystems (NRC (2000) Clean Coastal Waters: Understanding and Reducing the Effects of Nutrient Pollution. National Academy Press, Washington, DC; Bricker et al. (1999) National Estuarine Eutrophication Assessment. Effects of nutrient enrichment in the nation’s estuaries. NOAA-NOS Special Projects Office, Silver Spring, MD). However, not all of the nitrogen input to a watershed is exported to the coast (Howarth et al. (1996) Biogeochemistry 35:75–139; Jordan and Weller (1996) Bioscience 46:655–664). Global estimates of nitrogen export to coasts have been taken to be 25% of watershed input, based largely on northeastern U.S. observations (Galloway et al. (2004) Biogeochemistry 70:153–226; Boyer et al. (2006) Global Biogeochem Cycle 20:Art. No. GB1S91). We applied the N budgeting methodology developed for the International SCOPE Nitrogen project (Howarth et al. (1996) Biogeochemistry 35:75–139; Boyer et al. (2002) Biogeochemistry 57:137–169) to 12 watersheds in the southeastern U.S., and compared them with estimates of N export for 16 watersheds in the northeastern U.S. (Boyer et al. (2002) Biogeochemistry 57:137–169). In southeastern watersheds, average N export was only 9% of input, suggesting the need for downward revision of global estimates. The difference between northern and southern watersheds is not a function of the absolute value of N inputs, which spanned a comparable range and were positively related to export in both cases. Rather, the proportion of N exported was significantly related to average watershed temperature (% N export = 58.41 e^{−0.11 * temperature}; R ² = 0.76), with lower proportionate nitrogen export in warmer watersheds. In addition, we identified a threshold in proportionate N export at 38°N latitude that corresponds to a reported breakpoint in the rate of denitrification at 10–12°C. We hypothesize that temperature, by regulating denitrification, results in increased proportionate N export at higher latitudes. Regardless of the mechanism, these observations suggest that temperature increases associated with future climate change may well reduce the amount of nitrogen that reaches estuaries, which will have implications for coastal eutrophication.     

Divergent introduction histories among invasive populations of the delicate skink ( Lampropholis delicata): has the importance of genetic admixture in the success of biological invasions been overemphasized?

Aim

Invasive species are predicted to experience a reduction in genetic diversity during the introduction process because of founder effects, yet they are able to successfully establish in new regions and outcompete the native biota. Admixture has been proposed as a potential solution to this genetic paradox. We adopted a phylogeographic approach to investigate the invasion history of the delicate skink ( Lampropholis delicata) in the Pacific region and test the hypothesis that admixture is important for the success of biological invasions.

Location

Eastern Australia and the Pacific region (Lord Howe Island, New Zealand, Hawaii).

Methods

We obtained mitochondrial DNA sequence data ( ND2, ND4) from across the native Australian range (238 samples, 120 populations) and 371 samples from the introduced range of L. delicata. Genetic distances and Analysis of molecular variance (AMOVA) were used to examine the level of genetic variation across the native and introduced ranges.

Results

Fourteen haplotypes were evident in the introduced range (1 in Hawaii, 7 in New Zealand, 7 in Lord Howe Island), with a shared haplotype present in both New Zealand and Lord Howe Island. Five source regions were identified (Brisbane, Tenterfield, Border Ranges, Yamba‐Coffs Harbour, Sydney) from across four distinct native‐range genetic lineages. The Hawaiian population stems from a single introduction from Brisbane, whereas one or more introductions from the Tenterfield region led to the New Zealand populations. Multiple introductions from across all five source regions have resulted in extreme admixture (up to 8.3% sequence divergence) within Lord Howe Island.

Main Conclusions

L. delicata introductions are capable of being successful both in the presence and absence of admixture. Contrary to the predictions of the sequential two‐step model, the presence of admixture was not related to the time since initial introduction. We suggest that the importance of admixture in determining the success of biological invasions has been overemphasized.

     

Factors determining the number of land bird species on Islands around South-Western Australia

Summary New evidence from the passerine faunas of islands off Southwestern Australia agrees with the hypothesis that the passerine faunas of Australian and New Zealand islands are impoverished because most passerine species are poor colonizers. Dispersal of landbirds onto Carnac Island near Perth was infrequent, and many of those species that arrived were represented by single birds. Comparison of similarly structured island and mainland habitats showed that island habitats still have fewer passerine bird species than mainland habitats. Island bird faunas are more stable over short periods of time than over long periods; this is contrary to island avifaunas in the Northern Hemisphere.The following features typify the avifaunas of Australian islands: immigration of species of land birds occurs infrequently; (natural) extinction is rare; and the degree of saturation of the avifaunas is low. Without more direct evidence, competitive interactions should not be invoked to account for the species poverty of these insular avifaunas.