Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system

Introduced species are an increasing agent of global change. Biogeographic comparisons of introduced biotas at regional and global scales can clarify trends in source regions, invasion pathways, sink regions, and survey effort. We identify the Northeast Pacific Ocean (NEP; northern California to British Columbia) as a model system for analyzing patterns of marine invasion success in cool temperate waters. We review literature and field surveys, documenting 123 introduced invertebrate, algal, fish, and vascular plant species in the NEP. Major invasion pathways were shipping (hull fouling, solid and water ballast; 1500s-present) and shellfish (particularly oysters) and finfish imports (commonest from the 1870s to mid-1900s). The cumulative number of successful invasions over time increased at linear, quadratic, and exponential rates for different taxa, pathways, and regions within the NEP. Regional analysis of four major NEP estuaries showed that Puget Sound and the contiguous Straits had the most introduced species, followed by Humboldt Bay, Coos Bay and Willapa Bay. Data on cumulative shipping volumes predicted smaller-scale, but not larger-scale spatial patterns in the number of shipping-mediated invasions. We identify the major challenges in scaling up from regional to global invasion analysis in cool temperate regions. Retrospective analyses for distinct biogeographic regions such as the NEP provide insight into vector dynamics and regional invasibility, and are a necessary foundation for monitoring and managing global change caused by biotic invasions.     

Predicting the invasion success of Mediterranean alien plants from their introduction characteristics

We describe a model to predict the invasion success of alien plants on Mediterranean islands, according to their mode, frequency and time since introduction and simple species traits. The model was developed using canonical discriminant analysis to identify categories of low- and high-risk species. Whilst classifying ca 80% of the data set correctly, the type II error rate remained moderately high, which limits its value in practical terms. This level of accuracy is typical of similar models, and critical analysis of the weaknesses is necessary to develop long-term improvements in methodology. Sensitivity analysis indicates that, overall, mode of introduction had an important influence on invasiveness, with species introduced for public amenity being more likely to attain high-risk status than species introduced privately to gardens. Frequency and time since introduction had a relatively minor influence on the model outcome, but remained highly confounded with other variables. Although invasion success, as measured by the number of islands colonized, is expected to increase over time, it was not possible to estimate the rates of expansion from historical trends, which are complicated by changes in import fashions over the centuries. Many older introductions were crops, whereas there has been a recent tendency towards exotic ornamentals and weedy species. Despite a fundamental influence on the results, similar historical influences are seldom taken into account adequately during trait analyses. Unravelling such inter-correlations between predictors is therefore an important challenge for the future of screening protocols.     

Was Lates Late? A Null Model for the Nile Perch Boom in Lake Victoria

Nile perch (Lates niloticus) suddenly invaded Lake Victoria between 1979 and 1987, 25 years after its introduction in the Ugandan side of the lake. Nile perch then replaced the native fish diversity and irreversibly altered the ecosystem and its role to lakeshore societies: it is now a prised export product that supports millions of livelihoods. The delay in the Nile perch boom led to a hunt for triggers of the sudden boom and generated several hypotheses regarding its growth at low abundances – all hypotheses having important implications for the management of Nile perch stocks. We use logistic growth as a parsimonious null model to predict when the Nile perch invasion should have been expected, given its growth rate, initial stock size and introduction year. We find the first exponential growth phase can explain the timing of the perch boom at the scale of Lake Victoria, suggesting that complex mechanisms are not necessary to explain the Nile perch invasion or its timing. However, the boom started in Kenya before Uganda, indicating perhaps that Allee effects act at smaller scales than that of the whole Lake. The Nile perch invasion of other lakes indicates that habitat differences may also have an effect on invasion success. Our results suggest there is probably no single management strategy applicable to the whole lake that would lead to both efficient and sustainable exploitation of its resources.     

Bounded random walks as a null model for evaluating population trends

Management of wildlife and protection of endangered species depend on determination of population trends. Because population changes are stochastic and autoregressive, there is reason to believe that population trends might not be properly determined by simple regression over short time periods. A bounded random walk (BRW) model is introduced as a null model for evaluating population trends. The BRW model shows long-term stability but rising and falling sequences of up to many decades. For a given variability and survey length, there will be an expected probability of finding a greater than X% slope simply by chance. This false positive probability needs to be considered when evaluating trends. Breeding Bird Survey data for 128 species over 46 years for two states were analyzed for trends for different series lengths. Trends estimated from short series were likely to not agree with the 46-year trends. Very short series (e.g., 5 years) tended to indicate no trend due to loss of statistical power. A 101-year series for sandwich term (Sterna sandvicensis) revealed that even for 40 year-long series, 33% of subset series had a negative trend compared to the strong 101 year full series positive trend. The BRW model simulations and both data sets pointed to 20 years as a minimum time period for estimating trends reliably, though this can be longer for species that tend to cycle. Proper inference should thus consider the implications of inherent time series variability.     

The myth of plant species saturation

Plant species assemblages, communities or regional floras might be termed 'saturated' when additional immigrant species are unsuccessful at establishing due to competitive exclusion or other inter-specific interactions, or when the immigration of species is off-set by extirpation of species. This is clearly not the case for state, regional or national floras in the USA where colonization (i.e. invasion by exotic species) exceeds extirpation by roughly a 24 to 1 margin. We report an alarming temporal trend in plant invasions in the Pacific Northwest over the past 100 years whereby counties highest in native species richness appear increasingly invaded over time. Despite the possibility of some increased awareness and reporting of native and exotic plant species in recent decades, historical records show a significant, consistent long-term increase in exotic species (number and frequency) at county, state and regional scales in the Pacific Northwest. Here, as in other regions of the country, colonization rates by exotic species are high and extirpation rates are negligible. The rates of species accumulation in space in multi-scale vegetation plots may provide some clues to the mechanisms of the invasion process from local to national scales.     

Succession of Exotic and Native Species Assemblages within Restored Floodplain Forests: A Test of the Parallel Dynamics Hypothesis

Exotic plants pose a threat to restoration success in post‐agricultural bottomlands, but little information exists on their dynamics during succession of actively restored sites. We hypothesized that exotic assemblages would establish during succession and that their compositional trends during succession time would mirror those published for native species in other systems, with an early peak in herbaceous richness followed by a decline as woody species establish. In the summer of 2008, we sampled 16 sites across an 18‐year chronosequence of restored forests, with an additional four mature forest stands for comparison, within the Cypress Creek NWR, Illinois, U.S.A. We identified all vascular plant species and quantified canopy openness at three canopy strata, and soil texture and chemistry. Trends in exotic assemblages were significantly correlated with canopy openness at all strata. Richness of exotic and native herbaceous species was related to stand age and consistent with a Weibull regression model. Native and exotic herbaceous cover followed an exponential decay model. Woody native richness over time conformed to a logistic model; woody exotics exhibited no relationship with stand age and were present in sites of all ages. Our results indicate that although their rates of decline differ, herbaceous exotics and natives exhibit similar successional dynamics; therefore, herbaceous exotics may not pose a lasting threat to restoration success in reforested floodplains. Woody exotics can establish across a range of successional stages and persist under closed canopy conditions. Bottomland restorations are vulnerable to the invasion and expansion of exotic plant species even after canopy closure.     

Modelling of malaria risk,rates, and trends: A spatiotemporal approach for identifying and targeting sub-national areas of high and low burden

While mortality from malaria continues to decline globally, incidence rates in many countries are rising. Within countries, spatial and temporal patterns of malaria vary across communities due to many different physical and social environmental factors. To identify those areas most suitable for malaria elimination or targeted control interventions, we used Bayesian models to estimate the spatiotemporal variation of malaria risk, rates, and trends to determine areas of high or low malaria burden compared to their geographical neighbours. We present a methodology using Bayesian hierarchical models with a Markov Chain Monte Carlo (MCMC) based inference to fit a generalised linear mixed model with a conditional autoregressive structure. We modelled clusters of similar spatiotemporal trends in malaria risk, using trend functions with constrained shapes and visualised high and low burden districts using a multi-criterion index derived by combining spatiotemporal risk, rates and trends of districts in Zambia. Our results indicate that over 3 million people in Zambia live in high-burden districts with either high mortality burden or high incidence burden coupled with an increasing trend over 16 years (2000 to 2015) for all age, under-five and over-five cohorts. Approximately 1.6 million people live in high-incidence burden areas alone. Using our method, we have developed a platform that can enable malaria programs in countries like Zambia to target those high-burden areas with intensive control measures while at the same time pursue malaria elimination efforts in all other areas. Our method enhances conventional approaches and measures to identify those districts which had higher rates and increasing trends and risk. This study provides a method and a means that can help policy makers evaluate intervention impact over time and adopt appropriate geographically targeted strategies that address the issues of both high-burden areas, through intensive control approaches, and low-burden areas, via specific elimination programs.     

Can We Infer Island Introduction and Naturalization Rates from Inventory Data? Evidence from Introduced Plants in Galapagos

Studies of human-mediated rates of introduction of organisms to islands are few, results fall into two models (linear and exponential), and relationships with human population and activities are obscure. Owing to their late settlement and good scientific record, data from Galapagos may be exceptionally informative. The rate of introduction of vascular plant species to Galapagos has been suggested to be exponential, paralleling the rise in human population. However, detailed reconstruction of growth in numbers of introduced plant species, based on historical and recent records, reveals changes in study criteria over the last two centuries, which obscure the true introduction rate. At first, cultivated species were deliberately excluded from most studies. From the 1960s, naturalized cultivated species were included more consistently, but non-naturalized species were still excluded. From the mid-1980s, the latter were deliberately included. Accidental introductions increased linearly from 1807 (the first records) to the present. Escapes from cultivation show increases in rate around 1906 and in the period 1970–1990, the latter coinciding with the first studies directed at areas affected by human activities. Non-naturalized cultivated species rose abruptly from the late 1980s, as they became deliberately studied. There seems to be no direct link with human population size. Data represent rate of discovery rather than true introduction rate, and the changing overall rate reflects changing botanical interests and research effort. Data from other islands also suggest that linear increases in naturalized plants are the norm. Galapagos data do not permit confident statements about the introduction rate of cultivated species, but suggest that this may depend more on human activities than human population size.     

Culturally-transmitted feeding behaviour in primates: Evidence for accelerating learning rates

Cultural transmission implies the rapid spread of behavioural innovations when initially naïve individuals copy more informed ones. Mathematical models of transmission feature accelerating (and in most cases, logistic) rates of learning as animals that acquire an innovation provide ever increasing numbers of informers for potential learners. Conversely, non-accelerating rates have been proposed as a null hypothesis for apparent cases of cultural transmission that can best be explained by simpler mechanisms such as trial-and-error learning. Using the AIC technique for comparing models with different numbers of parameters, this paper examines the 21 cases in the primate literature where quantifiable data are available on learning rates for presumed culturally-transmitted feeding innovations. In each case, cumulative distributions over time of the frequency or proportion of individuals that acquire an innovation are compared with three accelerating functions (logistic, positive exponential, and hyperbolic sine) and two non-accelerating ones (linear and logarithmic). In 16 cases, the best fit is given by an accelerating function: nine of these support the logistic, four support the positive exponential and three, the reverse S-shaped hyperbolic sine. Individual cases often show small differences between alternative functions, but overall trends support the cultural assumption of accelerating learning rates.     

Clustered or scattered? The impact of habitat quality clustering on establishment and early spread

The match between the environmental conditions of an introduction area and the preferences of an introduced species is the first prerequisite for establishment. Yet, introduction areas are usually landscapes, i.e. heterogeneous sets of habitats that are more or less favourable to the introduced species. Because individuals are able to disperse after their introduction, the quality of the habitat surrounding the introduction site is as critical to the persistence of introduced populations as the quality of the introduction site itself. Moreover, demographic mechanisms such as Allee effects or dispersal mortality can hamper dispersal and affect spread across the landscape, in interaction with the spatial distribution of favourable habitat patches. In this study, we investigate the impact of the spatial distribution of heterogeneous quality habitats on establishment and early spread. First, we simulated introductions in one‐dimensional landscapes for different dispersal rates and either dispersal mortality or Allee effects. The landscapes differed by the distribution of favourable and less favourable habitats, which were either clustered into few large aggregates of the same quality or scattered into multiple smaller ones. Second, we tested the predictions of simulations by performing experimental introductions of hymenopteran parasitoids (Trichogramma chilonis) in ‘clustered’ and ‘scattered’ microcosm landscapes. Results highlighted two impacts of the clustering of favourable habitat: by decreasing the risks of dispersal from the introduction site to unfavourable habitat early during the invasion, it increased establishment success. However, by increasing the distance between favourable habitat patches, it also hindered the subsequent spread of introduced species over larger areas.     

Intrinsic determinants of a population trend in timing of breeding in the wandering albatross

Numerous studies of wild animal species have documented that population level responses to environmental change are underpinned by individual level phenotypic plasticity. However, where the relationship between an individual trait and a climate variable occurs when both show a trend over time, phenotypic plasticity may be confounded by ageing. We investigated between and within individual change in laying date in the wandering albatross Diomedea exulans, a long‐lived species experiencing a dramatic decline in population size. Laying date has advanced over the last three decades. A mean‐centering analysis demonstrated that this pattern was driven by within‐individual changes as opposed to appearance or disappearance of phenotypes. Furthermore, a lack of between individual effect suggested the change resulted from ageing as opposed to phenotypic plasticity. Females varied significantly in rate of advance, such that those with low past reproductive rates exhibited a negative temporal trend in laying date, whereas birds with moderate to high past reproductive performance showed little change. The population trend was therefore driven by a subset with low past breeding success. An analysis of effects of timing of breeding on breeding success revealed stabilizing selection for relative laying date. Furthermore, current breeding success was positively related to past success rate, which suggests that there may be indirect selection against plasticity in this population. Our results show that population trends can arise from individual level change unrelated to prevailing environmental conditions, thus demonstrating the importance of longitudinal analyses in the interpretation of climate change effects.     

Survival of Svalbard pink-footed geese Anser brachyrhynchus in relation to winter climate, density and land-use

1. Global change may strongly affect population dynamics, but mechanisms remain elusive. Several Arctic goose species have increased considerably during the last decades. Climate, and land-use changes outside the breeding area have been invoked as causes but have not been tested. We analysed the relationships between conditions on wintering and migration staging areas, and survival in Svalbard pink-footed geese Anser brachyrhynchus. Using mark-recapture data from 14 winters (1989-2002) we estimated survival rates and tested for time trends, and effects of climate, goose density and land-use. 2. Resighting rates differed for males and females, were higher for birds recorded during the previous winter and changed smoothly over time. Survival rates did not differ between sexes, varied over time with a nonsignificant negative trend, and were higher for the first interval after marking (0.88-0.97) than afterwards (0.74-0.93). Average survival estimates were 0.967 (SE 0.026) for the first and 0.861 (SE 0.023) for all later survival intervals. 3. We combined 16 winter and spring climate covariates into two principal components axes. F1 was related to warm/wet winters and an early spring on the Norwegian staging areas and F2 to dry/cold winters. We expected that F1 would be positively related to survival and F2 negatively. F1 explained 23% of survival variation (F1,10=3.24; one-sided P=0.051) when alone in a model and 28% (F1,9=4.50; one-sided P=0.031) in a model that assumed a trend for survival. In contrast, neither F2 nor density, land-use, or scaring practices on important Norwegian spring staging areas had discernible effects on survival. 4. Climate change may thus affect goose population dynamics, with warmer winters and earlier springs enhancing survival and fecundity. A possible mechanism is increased food availability on Danish wintering and Norwegian staging areas. As geese are among the main herbivores in Arctic ecosystems, climate change, by increasing goose populations, may have important indirect effects on Arctic vegetation. Our study also highlights the importance of events outside the breeding area for the population dynamics of migrant species.     

Years of Potential Life Lost Due to Occupational Fatal Injury in the United States

Fatal injury surveillance data coupled with life expectancy data may be used to assess the impact of occupational fatal injuries on years of potential life lost (YPLL). We compare three definitions of YPLL and trends over time in YPLL. Two definitions determine YPLL as expected life lost to fixed life expectancies of 65 or 85 years. The third definition uses actuarial adjustments of life expectancy given survival to a given age stratified by gender and race. Fatalities from the National Traumatic Occupational Fatality (NTOF) database are used to illustrate the three definitions of YPLL. The three YPLL measures were similar in magnitude and direction of the trend in YPLL over 1980-1992. Proper interpretation of these trends can only be made in conjunction with other measures (e.g., rates). Almost all YPLL trends are declining, implying that over time fatal injuries are shifting to older workers. The exception is the increasing trend in YPLL for the retail trade industry, injury rates have also been increasing over time for this industry. Mining and construction have the highest YPLL among all industries. This analysis suggests efforts to prevent the occupational fatalities of younger workers should focus on the retail trade, mining, and construction industries.     

Changes in temperature sensitivity of spring phenology with recent climate warming in Switzerland are related to shifts of the preseason

The spring phenology of plants in temperate regions strongly responds to spring temperatures. Climate warming has caused substantial phenological advances in the past, but trends to be expected in the future are uncertain. A simple indicator is temperature sensitivity, the phenological advance statistically associated with a 1°C warmer mean temperature during the “preseason”, defined as the most temperature‐sensitive period preceding the phenological event. Recent analyses of phenological records have shown a decline in temperature sensitivity of leaf unfolding, but underlying mechanisms were not clear. Here, we propose that climate warming can reduce temperature sensitivity simply by reducing the length of the preseason due to faster bud development during this time period, unless the entire preseason shifts forward so that its temperature does not change. We derive these predictions theoretically from the widely used “thermal time model” for bud development and test them using data for 19 phenological events recorded in 1970–2012 at 108 stations spanning a 1600 m altitudinal range in Switzerland. We consider how temperature sensitivity, preseason start, preseason length and preseason temperature change (i) with altitude, (ii) between the periods 1970–1987 and 1995–2012, which differed mainly in spring temperatures, and (iii) between two non‐consecutive sets of 18 years that differed mainly in winter temperatures. On average, temperature sensitivity increased with altitude (colder climate) and was reduced in years with warmer springs, but not in years with warmer winters. These trends also varied among species. Decreasing temperature sensitivity in warmer springs was associated with a limited forward shift of preseason start, higher temperatures during the preseason and reduced preseason length, but not with reduced winter chilling. Our results imply that declining temperature sensitivity can result directly from spring warming and does not necessarily indicate altered physiological responses or stronger constraints such as reduced winter chilling.     

Spatial dynamics of invasion: the geometry of introduced species

Many exotic species combine low probability of establishment at each introduction with rapid population growth once introduction does succeed. To analyse this phenomenon, we note that invaders often cluster spatially when rare, and consequently an introduced exotic's population dynamics should depend on locally structured interactions. Ecological theory for spatially structured invasion relies on deterministic approximations, and determinism does not address the observed uncertainty of the exotic-introduction process. We take a new approach to the population dynamics of invasion and, by extension, to the general question of invasibility in any spatial ecology. We apply the physical theory for nucleation of spatial systems to a lattice-based model of competition between plant species, a resident and an invader, and the analysis reaches conclusions that differ qualitatively from the standard ecological theories. Nucleation theory distinguishes between dynamics of single- and multi-cluster invasion. Low introduction rates and small system size produce single-cluster dynamics, where success or failure of introduction is inherently stochastic. Single-cluster invasion occurs only if the cluster reaches a critical size, typically preceded by a number of failed attempts. For this case, we identify the functional form of the probability distribution of time elapsing until invasion succeeds. Although multi-cluster invasion for sufficiently large systems exhibits spatial averaging and almost-deterministic dynamics of the global densities, an analytical approximation from nucleation theory, known as Avrami's law, describes our simulation results far better than standard ecological approximations.     

Parametric and semiparametric approaches to testing for seasonal trend in serial count data

We present two tests for seasonal trend in monthly incidence data. The first approach uses a penalized likelihood to choose the number of harmonic terms to include in a parametric harmonic model (which includes time trends and autogression as well as seasonal harmonic terms) and then tests for seasonality using a parametric bootstrap test. The second approach uses a semiparametric regression model to test for seasonal trend. In the semiparametric model, the seasonal pattern is modeled nonparametrically, parametric terms are included for autoregressive effects and a linear time trend, and a parametric bootstrap test is used to test for seasonality. For both procedures, a null distribution is generated under a null Poisson model with time trends and autoregression parameters.We apply the methods to skin melanoma incidence rates collected by the surveillance, epidemiology, and end results (SEER) program of the National Cancer Institute, and perform simulation studies to evaluate the type I error rate and power for the two procedures. These simulations suggest that both procedures are alpha-level procedures. In addition, the harmonic model/bootstrap test had similar or larger power than the semiparametric model/bootstrap test for a wide range of alternatives, and the harmonic model/bootstrap test is much easier to implement. Thus, we recommend the harmonic model/bootstrap test for the analysis of seasonal incidence data.     

Geographic signatures in species turnover: decoupling colonization and extinction across a latitudinal gradient

High latitude communities have low species richness and are rapidly warming with climate change. Thus, temporal changes in community composition are expected to be greatest at high latitudes. However, at the same time traits such as body size can also change with latitude, potentially offsetting or increasing changes to community composition over time. We tested how zooplankton communities (copepods and cladocerans) have changed over a 25–75 year time span by assessing colonization and extinction rates from lakes across an 1800 km latitudinal gradient, and further tested whether species traits predict rates of community change over time. Lake‐level dissimilarity, measured with Sorenson distance, decreased at higher latitudes. This decrease was due to higher colonization rates of cladocerans in lower latitude lakes and consistent extinction rates across the latitudinal gradient. At the species level, colonization increased with regional occupancy, and tended to be higher for smaller bodied, locally abundant, species. Local extinction rates were negatively correlated with local abundance and regional occupancy, but were not influenced by body size. None of these species‐specific characteristics changed predictably with latitude. Contrary to our expectations, low‐latitude zooplankton communities changed more rapidly than high‐latitude communities by becoming more species rich, not by losing species that were historically present. Moreover, colonization and extinction trends suggest that lakes have become increasingly dominated by species with smaller body sizes and that are already common locally and regionally. Together, these findings indicate that rates of species turnover in freshwater lakes across a latitudinal gradient are not predicted by rates of temperature change, but that turnover is nonetheless resulting in trait‐shifts that favour small, generalist species.     

Young adult cancer incidence trends in Taiwan and the U.S. from 2002 to 2016

BackgroundPrevious studies have not examined young adult cancer incidence trends in Taiwan, or comprehensively compared these trends at two nations with different population genetics, environmental exposures, and health care. Therefore, we compared the incidence rates and trends of the most common young adult cancers diagnosed at 20–39 years of age in Taiwan and the U.S.MethodsIncidence rates from 2002 to 2016 were calculated from the Taiwan National Health Insurance Research Datasets and the U.S. Surveillance, Epidemiology, and End Results Program. For trend assessment, average annual percent change (AAPC) values were calculated from 15 years of data using Joinpoint Regression Program. We also obtained sex or age of diagnosis stratified estimates.ResultsThe age-standardized overall young adult cancer incidence rate significantly increased from 2002 to 2016 in both Taiwan (AAPC=1.1%, 95% CI: 0.8–1.5%) and the U.S. (AAPC=1.8%, 95% CI: 1.1–2.4%). Cancers with significantly decreasing trends in Taiwan included cancers of the nasopharynx, liver, and tongue, which were not among the most common young adult cancers in the U.S. Cancers with significantly increasing trends in both Taiwan and the U.S. included colorectal, thyroid, and female breast cancers. Lymphoma, ovarian cancer, and lung and bronchus cancer had significantly increasing trends in Taiwan but not in the U.S. Although cervical cancer had significantly decreasing trends in both nations among those 30–39 years of age, its trend was significantly increasing in Taiwan but decreasing in the U.S. among those 20–29 years of age.ConclusionThe types of common young adult cancers as well as their incidence rates and trends differed in Taiwan and the U.S. Future studies should further understand the etiological factors driving these trends.     

The vulnerability of habitats to plant invasion: disentangling the roles of propagule pressure,time and sampling effort

Aim To quantify the vulnerability of habitats to invasion by alien plants having accounted for the effects of propagule pressure, time and sampling effort. Location New Zealand. Methods We used spatial, temporal and habitat information taken from 9297 herbarium records of 301 alien plant species to examine the vulnerability of 11 terrestrial habitats to plant invasions. A null model that randomized species records across habitats was used to account for variation in sampling effort and to derive a relative measure of invasion based either on all records for a species or only its first record. The relative level of invasion was related to the average distance of each habitat from the nearest conurbation, which was used as a proxy for propagule pressure. The habitat in which a species was first recorded was compared to the habitats encountered for all records of that species to determine whether the initial habitat could predict subsequent habitat occupancy. Results Variation in sampling effort in space and time significantly masked the underlying vulnerability of habitats to plant invasions. Distance from the nearest conurbation had little effect on the relative level of invasion in each habitat, but the number of first records of each species significantly declined with increasing distance. While Urban, Streamside and Coastal habitats were over‐represented as sites of initial invasion, there was no evidence of major invasion hotspots from which alien plants might subsequently spread. Rather, the data suggest that certain habitats (especially Roadsides) readily accumulate alien plants from other habitats. Main conclusions Herbarium records combined with a suitable null model provide a powerful tool for assessing the relative vulnerability of habitats to plant invasion. The first records of alien plants tend to be found near conurbations, but this pattern disappears with subsequent spread. Regardless of the habitat where a species was first recorded, ultimately most alien plants spread to Roadside and Sparse habitats. This information suggests that such habitats may be useful targets for weed surveillance and monitoring.