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Integrating local pastoral knowledge, participatory mapping, and species distribution modeling for risk assessment of invasive rubber vine ( Cryptostegia grandiflora ) in Ethiopia’s Afar region

The threats posed by invasive plants span ecosystems and economies worldwide. Local knowledge of biological invasions has proven beneficial for invasive species research, but to date no work has integrated this knowledge with species distribution modeling for invasion risk assessments. In this study, we integrated pastoral knowledge with Maxent modeling to assess the suitable habitat and potential impacts of invasive Cryptostegia grandiflora Robx. Ex R.Br. (rubber vine) in Ethiopia’s Afar region. We conducted focus groups with seven villages across the Amibara and Awash-Fentale districts. Pastoral knowledge revealed the growing threat of rubber vine, which to date has received limited attention in Ethiopia, and whose presence in Afar was previously unknown to our team. Rubber vine occurrence points were collected in the field with pastoralists and processed in Maxent with MODIS-derived vegetation indices, topographic data, and anthropogenic variables. We tested model fit using a jackknife procedure and validated the final model with an independent occurrence data set collected through participatory mapping activities with pastoralists. A Multivariate Environmental Similarity Surface analysis revealed areas with novel environmental conditions for future targeted surveys. Model performance was evaluated using area under the receiver-operating characteristic curve (AUC) and showed good fit across the jackknife models (average AUC = 0.80) and the final model (test AUC = 0.96). Our results reveal the growing threat rubber vine poses to Afar, with suitable habitat extending downstream of its current known location in the middle Awash River basin. Local pastoral knowledge provided important context for its rapid expansion due to acute changes in seasonality and habitat alteration, in addition to threats posed to numerous endemic tree species that provide critical provisioning ecosystem services. This work demonstrates the utility of integrating local ecological knowledge with species distribution modeling for early detection and targeted surveying of recently established invasive species.

Ecology and Society

Toward a comprehensive information system to assist invasive species management in Hawaii and Pacific Islands

The need for coordinated regional and global electronic databases to assist prevention, early detection, rapid response, and control of biological invasions is well accepted. The Pacific Basin Information Node (PBIN), a node of the National Biological Information Infrastructure, has been increasingly engaged in the invasive species enterprise since its establishment in 2001. Since this time, PBIN has sought to support frontline efforts at combating invasions, through working with stakeholders in conservation, agriculture, forestry, health, and commerce to support joint information needs. Although initial emphasis has been on Hawaii, cooperative work with other Pacific islands and countries of the Pacific Rim is already underway and planned.

Weed Science

Annual grass invasion is transforming the sagebrush biome’s songbird communities

Novel stressors like climate change and biological invasions alter ecological communities, resulting in changes to ecosystem services and biodiversity (that is, ecological transformation). Most ecological transformation research focuses on plants, but animals are likely affected by and plausibly mediate the extent, impact, and pace of transformations. In western North America, where invasive annual grasses are transforming sagebrush-steppe ecosystems, we quantified how transformation in vegetation drives songbird community change. We hypothesized that transformation of vegetation communities via invasion alters wildlife communities by favoring generalists over specialists, albeit with extinction debts that may temporarily obscure transformations’ consequences. Although local-scale songbird diversity increased with grass invasion, we found that this shift was accompanied by ongoing reorganization of songbird communities at larger scales (across the sagebrush biome), driven by heterogeneous impacts of invasion among species and guilds. Through our biome-wide analysis, we were also able to identify high-priority regions for conservation of sensitive songbird species. Our research provides evidence that wildlife communities are transforming alongside vegetation communities and offers insight into the approaches required to quantify nascent community turnover.

western United States

Understanding gaps in early detection of and rapid response to invasive species in the United States: A literature review and bibliometric analysis

While concepts regarding invasive species establishment patterns and eradication possibilities have long been a topic of invasion biology, the specific terminology referring to early detection of and rapid response to (EDRR) invasive species emerged in scientific literature during the early 2000s. Since then, the EDRR approach has expanded to include a suite of detection, planning, and management tools. By conducting a systematic literature review, we attempt to characterize the field of EDRR in the United States and its territories as reflected by publication records. Specifically, we assessed publication data such as the number of publications per year, the most common journals where papers were published, and the relationship between author’s keywords for studies focusing on aquatic and terrestrial habitats. For publications that used invasive species occurrence or abundance data (whether collected for the purposes of the respective publication or acquired from another data source), we manually vetted additional information such as focal taxa, data collection years and locations, sources of other data used, and whether data or code were deposited in open access formats. We also conducted network analyses for the author institutions that coauthored papers together most frequently and for the references most cited by EDRR publications. Overall, we found that silos existed in terms of which author institutions worked together, which existing literature was cited, and which topics were frequently explored. We also found evidence of substantial gaps in data access and use. For example, although a wide variety of data sources for invasive species occurrences are available, these sources were seldom cited by published literature, and newly collected data was not often deposited into invasive species databases or other open-source data repositories. Considering the continued advocation for a centralized national EDRR information system, our study suggests that facilitating access to data, decision support tools, and other informational resources represents a key opportunity for improving EDRR capabilities.

Ecological Informatics

Could climate change benefit invasive snakes? Modelling the potential distribution of the California Kingsnake in the Canary Islands

The interaction between climate change and biological invasions is a global conservation challenge with major consequences for invasive species management. However, our understanding of this interaction has substantial knowledge gaps; this is particularly relevant for invasive snakes on islands because they can be a serious threat to island ecosystems. Here we evaluated the potential influence of climate change on the distribution of invasive snakes on islands, using the invasion of the California kingsnake ( Lampropeltis californiae ) in Gran Canaria. We analysed the potential distribution of L. californiae under current and future climatic conditions in the Canary Islands, with the underlying hypothesis that the archipelago might be suitable for the species under these climate scenarios. Our results indicate that the Canary Islands are currently highly suitable for the invasive snake, with increased suitability under the climate change scenarios tested here. This study supports the idea that invasive reptiles represent a substantial threat to near-tropical regions, and builds on previous studies suggesting that the menace of invasive reptiles may persist or even be exacerbated by climate change. We suggest future research should continue to fill the knowledge gap regarding invasive reptiles, in particular snakes, to clarify their potential future impacts on global biodiversity.

Journal of Environmental Management

Essential elements of online information networks on invasive alien species

In order to be effective, information must be placed in the proper context and organized in a manner that is logical and (preferably) standardized. Recently, invasive alien species (IAS) scientists have begun to create online networks to share their information concerning IAS prevention and control. At a special networking session at the Beijing International Symposium on Biological Invasions, an online Eastern Asia-North American IAS Information Network (EA-NA Network) was proposed. To prepare for the development of this network, and to provide models for other regional collaborations, we compare four examples of global, regional, and national online IAS information networks: the Global Invasive Species Information Network, the Invasives Information Network of the Inter-American Biodiversity Information Network, the Chinese Species Information System, and the Invasive Species Information Node of the US National Biological Information Infrastructure. We conclude that IAS networks require a common goal, dedicated leaders, effective communication, and broad endorsement, in order to obtain sustainable, long-term funding and long-term stability. They need to start small, use the experience of other networks, partner with others, and showcase benefits. Global integration and synergy among invasive species networks will succeed with contributions from both the top-down and the bottom-up. ?? 2006 Springer.

Conference Paper

Invasive rat control is an efficient, yet insufficient, method for recovery of the critically endangered Hawaiian plant hau kuahiwi (Hibiscadelphus giffardianus)

Biological invasions of rodents and other species have been especially problematic on tropical islands. Invasive Rattus rattus consumption of Hibiscadelphus giffardianus (Malvaceae; common Hawaiian name hau kuahiwi) fruit and seeds has been hypothesized to be the most-limiting factor inhibiting the critically endangered tree, but this has not been experimentally tested, and little is known about other factors affecting seed dispersal, germination, and seedling establishment. Thus, we do not know if rat removal is sufficient to increase hau kuahiwi recruitment. This study aims to evaluate the effect of rat population control on the ability of hau kuahiwi to retain fruit and establish seedlings. We compared hau kuahiwi fruiting and seedling recruitment in a stand treated to reduce rat abundance and a neighbouring control stand. Fruit retention increased following treatment but seedling establishment did not. Although rat control improves the ability of hau kuahiwi to retain fruit, other, presently unknown inhibitors to seed dispersal, germination, and/or seedling development remain. Seed and seedling predation by other species, competition from numerous invasive plant species, unsuitable climate, and/or other factors may be primary inhibitors in the absence of rats, but we emphasize that progressive isolation of these factors at individual hau kuahiwi life stages may be necessary to identify the remaining threats to the conservation of this critically endangered plant.

PLoS ONE

Turbidity alters pre-mating social interactions between native and invasive stream fishes

Environmental degradation can result in the loss of aquatic biodiversity if impairment promotes hybridisation between non-native and native species. Although aquatic biological invasions involving hybridisation have been attributed to elevated water turbidity, the extent to which impaired clarity influences reproductive isolation among non-native and native species is poorly understood. We examined whether turbidity influences intraspecific and interspecific pre-mating social interactions between invasive red shiner ( Cyprinella lutrensis ) and native blacktail shiner ( Cyprinella venusta ) from the Upper Coosa River Basin (U.S.A.). We found that the number or duration of conspecific and heterospecific interactions increased under turbid conditions. Additionally, we found evidence indicating that native blacktail shiner females are especially likely to interact with invasive red shiner males due to species- and sex-specific responses to turbid conditions. These findings suggest that elevated turbidity can increase pre-mating social interactions between native and invasive species, which could result in greater hybridisation and promote the genetic assimilation of native species following species introductions. Thus, integrating knowledge of species behaviour into conservation and management planning can help deter the establishment and spread of invasive species.

Freshwater Biology

Linking the concept of scale to studies of biological diversity: evolving approaches and tools.

Although the concepts of scale and biological diversity independently have received rapidly increasing attention in the scientific literature since the 1980s, the rate at which the two concepts have been investigated jointly has grown much more slowly. We find that scale considerations have been incorporated explicitly into six broad areas of investigation related to biological diversity: (1) heterogeneity within and among ecosystems, (2) disturbance ecology, (3) conservation and restoration, (4) invasion biology, (5) importance of temporal scale for understanding processes, and (6) species responses to environmental heterogeneity. In addition to placing the papers of this Special Feature within the context of brief summaries of the expanding literature on these six topics, we provide an overview of tools useful for integrating scale considerations into studies of biological diversity. Such tools include hierarchical and structural-equation modelling, kriging, variable-width buffers, k -fold cross-validation, and cascading graph diagrams, among others. Finally, we address some of the major challenges and research frontiers that remain, and conclude with a look to the future.

Diversity and Distributions

Predicting foundation bunchgrass species abundances: Model-assisted decision-making in protected-area sagebrush steppe

Foundation species are structurally dominant members of ecological communities that can stabilize ecological processes and influence resilience to disturbance and resistance to invasion. Being common, they are often overlooked for conservation but are increasingly threatened from land use change, biological invasions, and over-exploitation. The pattern of foundation species abundances over space and time may be used to guide decision-making, particularly in protected areas for which they are iconic. We used ordinal logistic regression to identify the important environmental influences on the abundance patterns of bluebunch wheatgrass ( Pseudoroegneria spicata ), Thurber's needlegrass ( Achnatherum thurberianum ), and Sandberg bluegrass ( Poa secunda ) in protected-area sagebrush steppe. We then predicted bunchgrass abundances along gradients of topography, disturbance, and invasive annual grass abundance. We used model predictions to prioritize the landscape for implementation of a management and restoration decision-support tool. Models were fit to categorical estimates of grass cover obtained from an extensive ground-based monitoring dataset. We found that remnant stands of abundant wheatgrass and bluegrass were associated with steep north-facing slopes in higher and more remote portions of the landscape outside of recently burned areas where invasive annual grasses were less abundant. These areas represented only 25% of the landscape and were prioritized for protection efforts. Needlegrass was associated with south-facing slopes, but in low abundance and in association with invasive cheatgrass ( Bromus tectorum ). Abundances of all three species were strongly negatively correlated with occurrence of another invasive annual grass, medusahead ( Taeniatherum caput-medusae ). The rarity of priority bunchgrass stands underscored the extent of degradation and the need for prioritization. We found no evidence that insularity reduced invasibility; annual grass invasion represents a serious threat to protected-area bunchgrass communities. Our study area was entirely within the Wyoming big sagebrush ecological zone, understood to have inherently low resilience to disturbance and resistance to weed invasion. However, our study revealed important variation in abundance of the foundation species associated with resilience and resistance along the topographic-soil moisture gradient within this zone, providing an important foothold for conservation decision-making in these steppe ecosystems. We found the foundation species focus a parsimonious strategy linking monitoring to decision-making via biogeographic modeling.

Ecosphere

Annual grass invasion in sagebrush-steppe: The relative importance of climate, soil properties and biotic interactions

The invasion by winter-annual grasses (AGs) such as Bromus tectorum into sagebrush steppe throughout the western USA is a classic example of a biological invasion with multiple, interacting climate, soil and biotic factors driving the invasion, although few studies have examined all components together. Across a 6000-km 2 area of the northern Great Basin, we conducted a field assessment of 100 climate, soil, and biotic (functional group abundances, diversity) factors at each of 90 sites that spanned an invasion gradient ranging from 0 to 100 % AG cover. We first determined which biotic and abiotic factors had the strongest correlative relationships with AGs and each resident functional group. We then used regression and structural equation modeling to explore how multiple ecological factors interact to influence AG abundance. Among biotic interactions, we observed negative relationships between AGs and biodiversity, perennial grass cover, resident species richness, biological soil crust cover and shrub density, whereas perennial and annual forb cover, tree cover and soil microbial biomass had no direct linkage to AG. Among abiotic factors, AG cover was strongly related to climate (increasing cover with increasing temperature and aridity), but had weak relationships with soil factors. Our structural equation model showed negative effects of perennial grasses and biodiversity on AG cover while integrating the negative effects of warmer climate and positive influence of belowground processes on resident functional groups. Our findings illustrate the relative importance of biotic interactions and climate on invasive abundance, while soil properties appear to have stronger relationships with resident biota than with invasives.

Oregon

Using plant invasions to compare occurrence- and abundance-based calculations of biotic homogenisation: Are results complementary or contradictory?

Aim Beta diversity quantifies the similarity of ecological assemblages. Its increase, known as biotic homogenisation, can be a consequence of biological invasions. However, species occurrence (presence/absence) and abundance-based analyses can produce contradictory assessments of the magnitude and direction of changes in beta diversity. Previous work indicates these contradictions should be less frequent in nature than in theory, but a growing number of empirical studies report discrepancies between occurrence- and abundance-based approaches. Understanding if these discrepancies represent a few isolated cases or are systematic across a diversity of ecosystems would allow us to better understand the general patterns, mechanisms and impacts of biotic homogenisation. Location United States. Time Period 1963–2020. Major Taxa Studied Vascular plants. Methods We used a dataset of more than 70,000 vegetation survey plots to assess differences in biotic homogenisation with and without invasion using both occurrence- and abundance-based metrics of beta diversity. We estimated taxonomic biotic homogenisation by comparing beta diversity of invaded and uninvaded plots with both classes of metrics and investigated the characteristics of the non-native species pool that influenced the likelihood that these metrics disagree. Results In 78% of plot comparisons, occurrence- and abundance-based calculations agreed in direction, and the two metrics were generally well correlated. Our empirical results are consistent with previous theory. Discrepancies between the metrics were more likely when the same non-native species was at high cover at both plots compared for beta diversity, and when these plots were spatially distant. Main Conclusions In about 20% of cases, our calculations revealed differences in direction (homogenisation vs. differentiation) when comparing occurrence- and abundance-based metrics, indicating that the metrics are not interchangeable, especially when distances between plots are high and invader diversity is low. When data permit, combining the two approaches can offer insights into the role of invasions and extirpations in driving biotic homogenisation/differentiation.

Global Ecology and Biogeography

The effectiveness of non-native fish removal techniques in freshwater ecosystems: a systematic review

In aquatic systems, biological invasions can result in adverse ecological effects. Management techniques available for non-native fish removal programs (including eradication and population size control) vary widely, but include chemicals, harvest regimes, physical removal, or biological control. For management agencies, deciding on what non-native fish removal program to use has been challenging because there is little reliable information about the relative effectiveness of these measures in controlling or eradicating non-native fish. We conducted a systematic review, including a critical appraisal of study validity, to assess the effectiveness of different non-native fish removal methods, and to identify the factors that influence the overall success rate of each type of method. We found 95 relevant studies, generating 158 data sets. The evidence base was dominated by poorly documented studies with inadequate experimental designs (76% of removal projects). When the management goal was non-native fish eradication, chemical treatments were relatively successful (antimycin 75%; rotenone 89%) compared to other interventions. Electrofishing and passive removal measure studies indicated successful eradication was possible (58% each respectively) but required intensive effort and multiple treatments over a number of years. Of these studies with sufficient information, electrofishing had the highest success for population size control (56% of data sets). Overall, inadequate data quality and completeness severely limited our ability to make strong conclusions about the relationships between non-native fish abundance and different methods of eradication and population control, and the factors influencing the overall success rate of each method. Our review highlights that there is considerable scope for improving our evaluations of non-native fish removal methods. It is recommended that programs should have explicitly stated objectives, better data reporting, and study designs that (when possible and appropriate) incorporate replicated and controlled investigations with rigorous, long-term quantitative monitoring. Future research on the effectiveness of non-native fish removal methods should focus on: (1) the efficacy of existing or potentially new removal measures in larger, more complex environments; (2) a broader range of removal measures in general, and (3) phenotypic characteristics of individual fish within a population that fail to be eradicated or controlled.

Environmental Reviews

Integrating subsistence practice and species distribution modeling: assessing invasive elodea’s potential impact on Native Alaskan subsistence of Chinook salmon and whitefish

Alaska has one of the most rapidly changing climates on earth and is experiencing an accelerated rate of human disturbance, including resource extraction and transportation infrastructure development. Combined, these factors increase the state’s vulnerability to biological invasion, which can have acute negative impacts on ecological integrity and subsistence practices. Of growing concern is the spread of Alaska’s first documented freshwater aquatic invasive plant Elodea spp. (elodea). In this study, we modeled the suitable habitat of elodea using global and state-specific species occurrence records and environmental variables, in concert with an ensemble of model algorithms. Furthermore, we sought to incorporate local subsistence concerns by using Native Alaskan knowledge and available statewide subsistence harvest data to assess the potential threat posed by elodea to Chinook salmon ( Oncorhynchus tshawytscha ) and whitefish ( Coregonus nelsonii ) subsistence. State models were applied to future climate (2040–2059) using five general circulation models best suited for Alaska. Model evaluations indicated that our results had moderate to strong predictability, with area under the receiver-operating characteristic curve values above 0.80 and classification accuracies ranging from 66 to 89 %. State models provided a more robust assessment of elodea habitat suitability. These ensembles revealed different levels of management concern statewide, based on the interaction of fish subsistence patterns, known spawning and rearing sites, and elodea habitat suitability, thus highlighting regions with additional need for targeted monitoring. Our results suggest that this approach can hold great utility for invasion risk assessments and better facilitate the inclusion of local stakeholder concerns in conservation planning and management.

Alaska

Genetic reconstruction of a bullfrog invasion to elucidate vectors of introduction and secondary spread

Reconstructing historical colonization pathways of an invasive species is critical for uncovering factors that determine invasion success and for designing management strategies. The American bullfrog ( Lithobates catesbeianus ) is endemic to eastern North America, but now has a global distribution and is considered to be one of the worst invaders in the world. In Montana, several introduced populations have been reported, but little is known of their sources and vectors of introduction and secondary spread. We evaluated the genetic composition of introduced populations at local (Yellowstone River floodplain) and regional (Montana and Wyoming) scales in contrast to native range populations. Our objectives were to (1) estimate the number of introductions, (2) identify probable native sources, (3) evaluate genetic variation relative to sources, and (4) characterize properties of local- and regional-scale spread. We sequenced 937 bp of the mitochondrial cytochrome b locus in 395 tadpoles collected along 100 km of the Yellowstone River, from three additional sites in MT and a proximate site in WY. Pairwise Φ ST revealed high divergence among nonnative populations, suggesting at least four independent introductions into MT from diverse sources. Three cyt b haplotypes were identical to native haplotypes distributed across the Midwest and Great Lakes regions, and AMOVA confirmed the western native region as a likely source. While haplotype ( H d = 0.69) and nucleotide diversity ( π = 0.005) were low in introduced bullfrogs, the levels of diversity did not differ significantly from source populations. In the Yellowstone, two identified haplotypes implied few introduction vectors and a significant relationship between genetic and river distance was found. Evidence for multiple invasions and lack of subsequent regional spread emphasizes the importance of enforcing legislation prohibiting bullfrog importation and the need for continuing public education to prevent transport of bullfrogs in MT. More broadly, this study demonstrates how genetic approaches can reveal key properties of a biological invasion to inform management strategies.

Montana

Combining local, landscape, and regional geographies to assess plant community vulnerability to invasion impact

Invasive species science has focused heavily on the invasive agent. However, management to protect native species also requires a proactive approach focused on resident communities and the features affecting their vulnerability to invasion impacts. Vulnerability is likely the result of factors acting across spatial scales, from local to regional, and it is the combined effects of these factors that will determine the magnitude of vulnerability. Here, we introduce an analytical framework that quantifies the scale-dependent impact of biological invasions on native richness from the shape of the native species–area relationship (SAR). We leveraged newly available, biogeographically extensive vegetation data from the U.S. National Ecological Observatory Network to assess plant community vulnerability to invasion impact as a function of factors acting across scales. We analyzed more than 1000 SARs widely distributed across the USA along environmental gradients and under different levels of non-native plant cover. Decreases in native richness were consistently associated with non-native species cover, but native richness was compromised only at relatively high levels of non-native cover. After accounting for variation in baseline ecosystem diversity, net primary productivity, and human modification, ecoregions that were colder and wetter were most vulnerable to losses of native plant species at the local level, while warmer and wetter areas were most susceptible at the landscape level. We also document how the combined effects of cross-scale factors result in a heterogeneous spatial pattern of vulnerability. This pattern could not be predicted by analyses at any single scale, underscoring the importance of accounting for factors acting across scales. Simultaneously assessing differences in vulnerability between distinct plant communities at local, landscape, and regional scales provided outputs that can be used to inform policy and management aimed at reducing vulnerability to the impact of plant invasions.

Ecological Applications

Ten lessons for controlling invasive species: Wisdom from the long-standing sea lamprey control program on the Laurentian Great Lakes

Sea lamprey ( Petromyzon marinus ) control in the Laurentian Great Lakes of North America is among the largest and most successful control programs of an invasive species anywhere on the planet. The effort began more than 75 years ago; it unites multiple nations, states, and provinces with the common goal of controlling this invasive species and protecting a valuable fishery. The science-based control program is administered by the Great Lakes Fishery Commission (GLFC), a body arising from a treaty signed by the United States and Canada. In the present article, we share 10 lessons learned from decades of successful sea lamprey control with the hopes of informing ongoing and future control programs targeting biological invasions. The 10 lessons we identified are to act boldly in times of crisis, to maintain the social license, to invest in capacity building, to break down the silos, to support fundamental science, to diversify your portfolio of control measures, to strive for continuous improvement, to confront the trade-off between information and action, to keep your foot on the gas, and to keep your eyes on the prize. The GLFC has long fostered a framework that uses some military strategy and verbiage that extends across the lessons (e.g., know your enemy). Other lessons are more nascent as the GLFC reenvisions its relationship with Indigenous peoples and governments in a path to reconciliation where two-eyed seeing is being embraced. Through adaptive management, horizon scanning methods, and embracing implementation science, the lessons learned about sea lamprey control will continue to evolve, which is itself a lesson. We submit that the lessons shared in the present article will help guide invasive species control programs spanning taxa, ecosystems, and regions.

Laurentian Great Lakes

Arrival and expansion of the invasive foraminifera Trochammina hadai Uchio in Padilla Bay, Washington

Trochammina hadai Uchio, a benthic foraminifera native to Japanese estuaries, was first identified as an invasive in 1995 in San Francisco Bay and later in 16 other west coast estuaries. To investigate the timing of the arrival and expansion of this invasive species in Padilla Bay, Washington, we analyzed the distribution of foraminifera in two surface samples collected in 1971, in nine surface samples collected by Scott in 1972–1973, as well as in two cores (Padilla Flats 3 and Padilla V1/V2) obtained in 2004. Trochanimina hadai , originally identified as the native Trochammina pacifica Cushman in several early foraminiferal studies, dominates the assemblage of most of the surface samples. In the Padilla V1/V2 and Padilla Flats 3 cores, the species' abundance follows a pattern of absence, first appearance, rapid expansion commonly seen shortly after the arrival of a successful biological invasion, setback, and second expansion. Using Q-mode cluster analysis, pre-expansion and expansion assemblages were identified. Pb-210 dating of these cores proved unsuccessful. However, based on T. hadai's first appearance occurring stratigraphically well above sedimentological changes in the cores that reflect deposition of sediments in the bay due to previous diversions of the Skagit River, and its dominance in the early 1970s surface samples, we conclude that the species arrived in Padilla Bay somewhere between the late 1800s and 1971. Trochammina hadai may have been introduced into the bay in the 1930s when oyster culturing began there or, at a minimum, ten years prior to its appearance in San Francisco Bay.

Washington