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Review of concepts useful for maintaining quality of male reproductive field samples for laboratory study

Investigations into cellular and molecular characteristics of male gametes obtained from fish in natural ecosystems require careful sample handling and shipping in order to minimize artifacts. Maintaining sample integrity engenders confident assessments of ecosystem health, whereby animal condition is often reflected by gamete biomarkers - indicators that respond in measurable ways to changes. A number of our investigations have addressed the hypothesis that biomarkers from fish along a pollution gradient are reflective of site location. Species biology and the selected biological endpoints direct choice of parameters such as: temperature, buffer osmolality, time in transit, fixation, cryoprotectants, protease inhibition, and antibiotic inclusion in extender. This paper will highlight case studies, and outline parameters and thoughts on approaches for use by field and laboratory researchers.

Conference Paper

Acidification in the U.S. Southeast: Causes, potential consequences and the role of the Southeast Ocean and Coastal Acidification Network

Coastal acidification in southeastern U.S. estuaries and coastal waters is influenced by biological activity, run-off from the land, and increasing carbon dioxide in the atmosphere. Acidification can negatively impact coastal resources such as shellfish, finfish, and coral reefs, and the communities that rely on them. Organismal responses for species located in the U.S. Southeast document large negative impacts of acidification, especially in larval stages. For example, the toxicity of pesticides increases under acidified conditions and the combination of acidification and low oxygen has profoundly negative influences on genes regulating oxygen consumption. In corals, the rate of calcification decreases with acidification and processes such as wound recovery, reproduction, and recruitment are negatively impacted. Minimizing the changes in global ocean chemistry will ultimately depend on the reduction of carbon dioxide emissions, but adaptation to these changes and mitigation of the local stressors that exacerbate global acidification can be addressed locally. The evolution of our knowledge of acidification, from basic understanding of the problem to the emergence of applied research and monitoring, has been facilitated by the development of regional Coastal Acidification Networks (CANs) across the United States. This synthesis is a product of the Southeast Coastal and Ocean Acidification Network (SOCAN). SOCAN was established to better understand acidification in the coastal waters of the U.S. Southeast and to foster communication among scientists, resource managers, businesses, and governments in the region. Here we review acidification issues in the U.S. Southeast, including the regional mechanisms of acidification and their potential impacts on biological resources and coastal communities. We recommend research and monitoring priorities and discuss the role SOCAN has in advancing acidification research and mitigation of and adaptation to these changes.

Florida, Georgia, North Carolina, South Carolina

A multi-metric assessment of environmental contaminant exposure and effects in an urbanized reach of the Charles River near Watertown, Massachusetts

The Charles River Project provided an opportunity to simultaneously deploy a combination of biomonitoring techniques routinely used by the U.S. Geological Survey National Water Quality Assessment Program, the Biomonitoring of Environmental Status and Trends Project, and the Contaminant Biology Program at an urban site suspected to be contaminated with polycyclic aromatic hydrocarbons. In addition to these standardized methods, additional techniques were used to further elucidate contaminant exposure and potential impacts of exposure on biota. The purpose of the study was to generate a comprehensive, multi-metric data set to support assessment of contaminant exposure and effects at the site. Furthermore, the data set could be assessed to determine the relative performance of the standardized method suites typically used by the National Water Quality Assessment Program and the Biomonitoring of Environmental Status and Trends Project, as well as the additional biomonitoring methods used in the study to demonstrate ecological effects of contaminant exposure. The Contaminant Effects Workgroup, an advisory committee of the U.S. Geological Survey/Contaminant Biology Program, identified polycyclic aromatic hydrocarbons as the contaminant class of greatest concern in urban streams of all sizes. The reach of the Charles River near Watertown, Massachusetts, was selected as the site for this study based on the suspected presence of polycyclic aromatic hydrocarbon contamination and the presence of common carp ( Cyprinus carpio ), largemouth bass ( Micropterus salmoides ), and white sucker ( Catostomus commersoni ). All of these fish have extensive contaminant-exposure profiles related to polycyclic aromatic hydrocarbons and other environmental contaminants. This project represented a collaboration of universities, Department of the Interior bureaus including multiple components of the USGS (Biological Resources Discipline and Water Resources Discipline Science Centers, the Contaminant Biology Program, and the Status and Trends of Biological Resources Program), and the U.S. Fish and Wildlife Service. Samples for analyzing water chemistry, sediment chemistry and toxicity, fish community structure, tissue chemistry, and fish (20 carp, 20 bass, and 40 white sucker) and invertebrate pathology were collected in late August, 2005. This report provides results from the analyses of fish pathology, biomarkers of exposure and effects (reproductive, carcinogenic, genotoxic, and immunologic), sediment chemistry, toxicity, and fish and invertebrate community structure.

Massachusetts

Associations of stream geomorphic conditions and prevalence of alternative reproductive tactics among sockeye salmon populations

In many species, males may exhibit alternative life histories to circumvent the costs of intrasexual competition and female courtship. While the evolution and underlying genetic and physiological mechanisms behind alternative reproductive tactics are well studied, there has been less consideration of the ecological factors that regulate their prevalence. Here, we examine six decades of age composition records from thirty‐six populations of sockeye salmon ( Oncorhynchus nerka ) to quantify associations between spawning habitat characteristics and the prevalence of precocious sneakers known as ‘jacks’. Jack prevalence was independent of neutral genetic structure among stream populations, but varied among habitat types and as a function of continuous geomorphic characteristics. Jacks were more common in streams relative to beaches and rivers, and their prevalence was negatively associated with stream width, depth, elevation, slope and area, but positively related to bank cover. Behavioural observations showed that jacks made greater use of banks, wood and shallows than guard males, indicating that their reproductive success depends on the availability of such refuges. Our results emphasize the role of the physical habitat in shaping reproductive tactic frequencies among populations, likely through local adaptation in response to variable fitness expectations under different geomorphic conditions.

Journal of Evolutionary Biology

Biology and impacts of Pacific island invasive species. 2. Boiga irregularis , the Brown Tree Snake (Reptilia: Colubridae)

The Brown Tree Snake, Boiga irregularis (Merrem, 1802), was accidentally transported to the island of Guam shortly after World War II. Over the following two decades it spread throughout the island with little public or professional recognition of its extent or impacts. This secretive nocturnal arboreal snake occurs in all habitats on Guam, from grasslands to forests. Under the right conditions, it is capable of high rates of reproduction and population growth. The Brown Tree Snake caused the extirpation of 13 of Guam's 22 native breeding birds and contributed to the extirpation of several species of native bats and lizards. Guam's 12 forest birds were especially impacted, with 10 species eliminated and the other two severely reduced. In addition, the snake continues to substantially impact domestic poultry, pets, the island's electrical power infrastructure, and human health. To protect other vulnerable Pacific islands, the U.S. government annually spends several million dollars inspecting cargo outbound from Guam to exclude Brown Tree Snakes. Cargo destinations most at risk are in Micronesia, especially the Northern Mariana Islands, but Guam also has direct air transportation links to Hawai'i that will soon be supplemented with direct ship traffic. Ultimately, all Pacific islands are at risk but especially those obtaining cargo through Guam. ?? 2007 by University of Hawai'i Press. All rights reserved.

Pacific Science

Challenge to the model of lake charr evolution: Shallow- and deep-water morphs exist within a small postglacial lake

All examples of lake charr ( Salvelinus namaycush ) diversity occur within the largest, deepest lakes of North America (i.e. > 2000 km 2 ). We report here Rush Lake (1.3 km 2 ) as the first example of a small lake with two lake charr morphs (lean and huronicus). Morphology, diet, life history, and genetics were examined to demonstrate the existence of morphs and determine the potential influence of evolutionary processes that led to their formation or maintenance. Results showed that the huronicus morph, caught in deep-water, had a deeper body, smaller head and jaws, higher eye position, greater buoyancy, and deeper peduncle than the shallow-water lean morph. Huronicus grew slower to a smaller adult size, and had an older mean age than the lean morph. Genetic comparisons showed low genetic divergence between morphs, indicating incomplete reproductive isolation. Phenotypic plasticity and differences in habitat use between deep and shallow waters associated with variation in foraging opportunities seems to have been sufficient to maintain the two morphs, demonstrating their important roles in resource polymorphism. Rush Lake expands previous explanations for lake charr intraspecific diversity, from large to small lakes and from reproductive isolation to the presence of gene flow associated with strong ecological drivers.

Biological Journal of the Linnean Society

Comparative microhabitat characteristics at oviposition sites of the California red-legged frog ( Rana draytonii )

We studied the microhabitat characteristics of 747 egg masses of the federally-threatened Rana draytonii (California red-legged frog) at eight sites in California. our study showed that a broad range of aquatic habitats are utilized by ovipositing R. draytonii , including sites with perennial and ephemeral water sources, natural and constructed wetlands, lentic and lotic hydrology, and sites surrounded by protected lands and nested within modified urban areas. We recorded 45 different egg mass attachment types, although the use of only a few types was common at each site. These attachment types ranged from branches and roots of riparian trees, emergent and submergent wetland vegetation, flooded upland grassland/ruderal vegetation, and debris. eggs were deposited in relatively shallow water (mean 39.7 cm) when compared to maximum site depths. We found that most frogs in artificial pond, natural creek, and artificial channel habitats deposited egg masses within one meter of the shore, while egg masses in a seasonal marsh averaged 27.3 m from the shore due to extensive emergent vegetation. Rana draytonii appeared to delay breeding in lotic habitats and in more inland sites compared to lentic habitats and coastal sites. eggs occurred as early as mid-december at a coastal artificial pond and as late as mid-April in an inland natural creek. We speculate that this delay in breeding may represent a method of avoiding high-flow events and/or freezing temperatures. Understanding the factors related to the reproductive needs of this species can contribute to creating, managing, or preserving appropriate habitat, and promoting species recovery.

Herpetological Conservation and Biology

Assessing the impact of charr research past, present, and future

The 9th International Charr Symposium convened on 18–21 June 2018, in Duluth, Minnesota, USA to gather scientists with an interest in charr biology and management from the entire geographical range of the genus Salvelinus. The symposium was attended by 169 individuals from six countries, and included 99 oral and 32 poster presentations, 28 of which were published in the ensuing proceedings. Topic areas of presentations and publications included: (1) cultural anthropology; (2) genetics, evolution, taxonomy and systematics; (3) behavior, movement and habitat; (4) reproductive ecology, developmental ontogeny and physiology; (5) population ecology, dynamics and life history; (6) trophic ecology, parasites, predators, toxicology and pollution; and (7) management. Research reported in these proceedings has built upon the history of earlier symposiums and will be continued with the 10th symposium scheduled to convene in Nikko City, Tochigi Prefecture, Japan, in 2021.

Hydrobiologia

Regional differentiation in somatic growth and maturation attributes for loggerhead sea turtles (Caretta caretta) in the Northwest Atlantic

The ages and sizes at which organisms mature have significant implications for lifetime reproductive success. For species at risk of extinction, such as sea turtles, these attributes can ultimately impact probability of population persistence. Within the Northwest Atlantic Ocean, the broader loggerhead sea turtle ( Caretta caretta ) population comprises management units both along the US Gulf of America (formerly Gulf of Mexico) and Atlantic coasts. Although age, growth, and maturation have been more intensively studied along the US Atlantic, data specific to the Gulf of America have remained sparse. To address this data gap, we conducted skeletal growth mark analysis (skeletochronology) for 123 humerus bones collected from loggerheads found dead in the US Gulf of America from 1998 to 2021. We compared resulting age, growth, and maturation data with information from studies of US Atlantic coast loggerheads for similar size and year ranges, using the exact same skeletochronology methods, as well as with Gulf of America mark-recapture growth data. Results indicate that Gulf of America loggerheads exhibit significantly faster juvenile somatic growth. In addition, sizes at maturation were substantially smaller, corresponding with mean estimates of age at maturation 7.5 to 15 years earlier than US Atlantic counterparts. Finally, the maximum observed Gulf of America adult age estimate was 42.5 years, considerably less than the highest US Atlantic estimate of 77.0 years. These detailed data offer insights into regional variability in somatic growth dynamics and characteristics associated with maturation, which in turn can impact relative reproductive contributions and, ultimately, population trajectories.

Marine Biology

Adaptive management assists reintroduction as higher tides threaten an endangered salt marsh plant

In theory, extirpated plant species can be reintroduced and managed to restore sustainable populations. However, few reintroduced plants are known to persist for more than a few years. Our adaptive‐management case study illustrates how we restored the endangered hemiparasitic annual plant, Chloropyron maritimum subsp. maritimum (salt marsh bird's beak), to Sweetwater Marsh, San Diego Bay National Wildlife Refuge, California, United States, and used monitoring and experimentation to identify the factors limiting the reintroduced population. After extirpation in 1988, reintroduction starting that year led to a resilient, genetically diverse population in 2016 (a “boom” of approximately 14,000) that rebounded from a “bust” (62 in 2014). Multiple regressions attributed 82% of the variation in population counts to tidal amplitude, rainfall, and temperature. Populations of salt marsh bird's beak crashed when the diurnal tide range peaked during the 18.6‐year lunar nodal cycle (a rarely considered factor that periodically added approximately 12 cm to tidal ranges). We explain booms as follows: During smaller tidal amplitudes , above‐average rainfall could desalinize upper intertidal soils and stimulate salt marsh bird's beak germination. Then, moderate temperature in May favors growth to reproduction in June. In addition, salt marsh bird's beak needs a short and open canopy of native perennial plants, with roots to parasitize (not non‐native annual grass pseudohosts) and nearby upland soil for a preferred pollinator, ground‐burrowing bees. Although our reintroduced salt marsh bird's beak population is an exceptional case of persistence, this rare species‐specific environmental and biological requirement makes it vulnerable to rising sea levels and global warming.

California

A decade of death and other dynamics: Deepening perspectives on the diversity and distribution of sea stars and wasting

Mass mortality events provide valuable insight into biological extremes and also ecological interactions more generally. The sea star wasting epidemic that began in 2013 catalyzed study of the microbiome, genetics, population dynamics, and community ecology of several high-profile species inhabiting the northeastern Pacific but exposed a dearth of information on the diversity, distributions, and impacts of sea star wasting for many lesser-known sea stars and a need for integration across scales. Here, we combine datasets from single-site to coast-wide studies, across time lines from weeks to decades, for 65 species. We evaluated the impacts of abiotic characteristics hypothetically associated with sea star wasting (sea surface temperature, pelagic primary productivity, upwelling wind forcing, wave exposure, freshwater runoff) and species characteristics (depth distribution, developmental mode, diet, habitat, reproductive period). We find that the 2010s sea star wasting outbreak clearly affected a little over a dozen species, primarily intertidal and shallow subtidal taxa, causing instantaneous wasting prevalence rates of 5%–80%. Despite the collapse of some populations within weeks, environmental and species variation protracted the outbreak, which lasted 2–3 years from onset until declining to chronic background rates of ∼2% sea star wasting prevalence. Recruitment began immediately in many species, and in general, sea star assemblages trended toward recovery; however, recovery was heterogeneous, and a marine heatwave in 2019 raised concerns of a second decline. The abiotic stressors most associated with the 2010s sea star wasting outbreak were elevated sea surface temperature and low wave exposure, as well as freshwater discharge in the north. However, detailed data speaking directly to the biological, ecological, and environmental cause(s) and consequences of the sea star wasting outbreak remain limited in scope, unavoidably retrospective, and perhaps always indeterminate. Redressing this shortfall for the future will require a broad spectrum of monitoring studies not less than the taxonomically broad cross-scale framework we have modeled in this synthesis.

Biological Bulletin

Evidence of counter-gradient growth in western pond turtles ( Actinemys marmorata ) across thermal gradients

Counter-gradient growth, where growth per unit temperature increases as temperature decreases, can reduce the variation in ectothermic growth rates across environmental gradients. Understanding how ectothermic species respond to changing temperatures is essential to their conservation and management due to human-altered habitats and changing climates. Here, we use two contrasting populations of western pond turtles ( Actinemys marmorata ) to model the effect of artificial and variable temperature regimes on growth and age at reproductive maturity. The two populations occur on forks of the Trinity River in northern California, U.S.A. The South Fork Trinity River (South Fork) is unregulated, while the main stem of the Trinity River (Main Stem) is dammed and has peak seasonal temperatures that are approximately 10 °C colder than the South Fork. Consistent with other studies, we found reduced annual growth rates for turtles in the colder Main Stem compared to the warmer South Fork. The South Fork population matured approximately 9 year earlier, on average, and at a larger body size than the Main Stem population. When we normalised growth rates for the thermal opportunity for growth using water-growing degree-days (GDD), we found the reverse for growth rates and age at reproductive maturity. Main Stem turtles grew approximately twice as fast as South Fork turtles per GDD. Main Stem turtles also required approximately 50% fewer GDD to reach their smaller size at reproductive maturity compared to the larger South Fork turtles. We found we could accurately hindcast growth rates based on water temperatures estimated from the total volume of discharge from the dam into the Main Stem, providing a management tool for predicting the impacts of the dam on turtle growth rates. Given the importance of size and age at reproductive maturity to population dynamics, this information on counter-gradient growth will improve our ability to understand and predict the consequences of dam operations for downstream turtle populations.

California

Life history, genetics, range expansion and new frontiers of the lionfish (Pterois volitans, Perciformes: Pteroidae) in Latin America

Pterois volitans (lionfish) is a midsize predatory fish commonly found in waters of the western Pacific and Indian Ocean. The species was first documented in Dania Beach, Florida (northwestern Caribbean) in 1985. Since that time the species has expanded its range rapidly to the Northwestern Atlantic Ocean, Gulf of Mexico, and Caribbean Sea. Since its introduction P. volitans has changed community structure and biodiversity of Caribbean reef communities and other coastal tropical ecosystems. Continuous introductions (accidental or intentional), limited natural predators, naïve-range prey behavior, high predation rates on competitors, continuous reproduction, and an extended period of larval dispersal have been the keys for successful invasion and rapid range extension of P. volitans . This invasion has become so severe that it has been recognized as one of the world’s top conservation issues. Here, we review the life history, behavior, and historical and contemporary genetic patterns that facilitate expansion and the colonization process. A greater understanding of lionfish biology, ecology, and the changes related to its present condition as a super-invader could improve current and future management strategies and new detection and response methodologies. We also examine new invasion frontiers that this species has the potential to colonize such as the eastern Pacific. This information will provide managers, the scientific community, and the civil society better tools for eradication, control and management of future invasions of this and other invasive species.

Caribbean Sea, Latin America

Global phylogeographic limits of Hawaii's avian malaria

The introduction of avian malaria (Plasmodium relictum) to Hawaii has provided a model system for studying the influence of exotic disease on naive host populations. Little is known, however, about the origin or the genetic variation of Hawaii's malaria and traditional classification methods have confounded attempts to place the parasite within a global ecological and evolutionary context. Using fragments of the parasite mitochondrial gene cytochrome b and the nuclear gene dihydrofolate reductase-thymidylate synthase obtained from a global survey of greater than 13 000 avian samples, we show that Hawaii's avian malaria, which can cause high mortality and is a major limiting factor for many species of native passerines, represents just one of the numerous lineages composing the morphological parasite species. The single parasite lineage detected in Hawaii exhibits a broad host distribution worldwide and is dominant on several other remote oceanic islands, including Bermuda and Moorea, French Polynesia. The rarity of this lineage in the continental New World and the restriction of closely related lineages to the Old World suggest limitations to the transmission of reproductively isolated parasite groups within the morphological species. ?? 2006 The Royal Society.

Proceedings of the Royal Society B: Biological Sci

Biological consequences of the coaster brook trout restoration stocking program in Lake Superior tributaries with Pictured Rocks National Lakeshore

The coaster Brook Trout Salvelinus fontinalis is a Lake Superior ecotype representing intraspecific variation that has been impacted by habitat loss and overfishing. Hatchery strains of Brook Trout derived from populations in Lake Superior were stocked into streams within Pictured Rocks National Lakeshore, Michigan, as part of an effort to rehabilitate adfluvial coaster Brook Trout. Wild and hatchery Brook Trout from three streams (Mosquito River, Hurricane River, and Sevenmile Creek) were examined for movement behavior, size, physiology, and reproductive success. Behavior and size of the stocked fish were similar to those of wild fish, and less than 15% of the stocked, tagged Brook Trout emigrated from the river into which they were stocked. There was little evidence of successful reproduction by stocked Brook Trout. Similar to the results of other studies, our findings suggest that the stocking of nonlocal Brook Trout strains where a local population already exists results in limited natural reproduction and should be avoided, particularly if the mechanisms governing the ecotype of interest are poorly understood.

Michigan

An adaptive-management framework for optimal control of hiking near golden eagle nests in Denali National Park

Unintended effects of recreational activities in protected areas are of growing concern. We used an adaptive-management framework to develop guidelines for optimally managing hiking activities to maintain desired levels of territory occupancy and reproductive success of Golden Eagles (Aquila chrysaetos) in Denali National Park (Alaska, U.S.A.). The management decision was to restrict human access (hikers) to particular nesting territories to reduce disturbance. The management objective was to minimize restrictions on hikers while maintaining reproductive performance of eagles above some specified level. We based our decision analysis on predictive models of site occupancy of eagles developed using a combination of expert opinion and data collected from 93 eagle territories over 20 years. The best predictive model showed that restricting human access to eagle territories had little effect on occupancy dynamics. However, when considering important sources of uncertainty in the models, including environmental stochasticity, imperfect detection of hares on which eagles prey, and model uncertainty, restricting access of territories to hikers improved eagle reproduction substantially. An adaptive management framework such as ours may help reduce uncertainty of the effects of hiking activities on Golden Eagles

Alaska

The central arctic caribou herd

From the mid-1970s through the mid-1980s, use of calving and summer habitats by Central Arctic herd caribou ( Rangifer tarandus granti ) declined near petroleum development infrastructure on Alaska's arctic coastal plain (Cameron et al. 1979; Cameron and Whitten 1980, Smith and Cameron 1983. Whitten and Cameron 1983a, 1985: Dau and Cameron 1986). With surface development continuing to expand westward from the Prudhoe Bay petroleum development area (Fig. 4.1), concerns arose that the resultant cumulative losses of habitat would eventually reduce productivity of the caribou herd. Specifically, reduced access of adult females to preferred foraging areas might adversely affect growth and fattening (Elison et al. 1986. Clough et al. 1987), in turn depressing calf production (Dauphiné 1976, Thomas 1982, Reimers 1983, White 1983, Eloranta and Nieminen 1986. Lenvik et al. 1988, Thomas and Kiliaan 1991) and survival (Haukioja and Salovaara 1978, Rognmo et al. 1983, Skogland 1984, Eloranta and Nieminen 1986, Adamczewski et al. 1987). Those concerns, though justified in theory, lacked empirical support. With industrial development in arctic Alaska virtually unprecedented, there was little basis for predicting the extent and duration of habitat loss, much less the secondary short- and long-term effects on the well-being of a particular caribou herd. Furthermore, despite a general acceptance that body condition and fecundity of the females are functionally related for reindeer and caribou, it seemed unlikely that any single model would apply to all subspecies of Rangifer , and perhaps not even within a subspecies in different geographic regions. We therefore lacked a complete understanding of the behavioral responses of arctic caribou to industrial development, the manner in which access to habitats might be affected, and how changes in habitat use might translate into measurable effects on fecundity and herd growth rate. Our study addressed the following objectives: 1) estimate variation in the size and productivity of the Central Arctic herd; 2) estimate changes in the distribution and movements of Central Arctic herd caribou in relation to the oil field development; 3) estimate the relationships between body condition and reproductive performance of female Central Arctic herd caribou, and 4) compare the body condition, reproductive success, and offspring survival of females under disturbance-free conditions (i.e., east of the Sagavanirktok River) with the status of those exposed to petroleum-related development (i.e., west of the Sagavanirktok River).

Alaska, Northwest Territories, Yukon Territory

Observations of recruitment and colonization by tunicates and associated invertebrates using giant one-meter 2 recruitment plates at Woods Hole, Massachusetts

Large recruitment plates measuring 1 × 1 m were deployed over an 18-month period from September 2013 to March 2015 for the purpose of documenting recruitment and colonization processes of marine invertebrate species at Woods Hole, Massachusetts. Each side of two plates was subdivided into 16 subareas (25 × 25 cm), and an observational strategy was developed whereby, at approximately two-week intervals, a different subarea was cleaned. Using this approach, we were able to photographically document species recruitment and growth interactions. Water temperature records from the site show that steady warming and cooling between 3 and 20° C changed at a mean rate of 0.2 ° C d -1 . However, temperature changes during the coolest and warmest parts of the temperature cycle were highly variable. In 2014, between the first and last occurrence of 0° C, temperatures were ≤0° C 15 percent of the time, but in 2015 temperatures were ≤0° C 93 percent of the time. In 2014, between the first and last occurrence of 21° C, temperatures were ≥21° C 88 percent of the time, and this warm period correlated with the disappearance of the hydroid Ectopleura crocea , the solitary tunicates Ascidiella aspersa and Ciona intestinalis , and the 2013 generation of Botrylloides violaceus . In Woods Hole, large plates provided enough space to accommodate both fast- and slow-colonizing species, resulting in the establishment of a diverse assemblage that was observed over a long time period. The most successful colonizing species had relatively long reproductive and recruitment periods, grew rapidly, repelled settlement onto their surfaces by larvae of any species, defended themselves against overgrowth by any species, overwintered, and lived a long time. Of the three dominant species observed in this study, the colonial tunicates Didemnum vexillum and Botrylloides violaceus had these qualities; the encrusting colonial bryozoan Schizoporella unicornis had all but one, it grew more slowly than the others. Barnacles constituted the only biological substrate that was effectively colonized by other species, both by larval recruitment and overgrowth. In Woods Hole, after a substrate had become fully colonized, there was very little opportunity for new recruitment or colony growth until new substrate opened after the death of colonies and individuals and the disappearance of biogenic structures such as amphipod tubes. An understanding of colonization processes utilized by invasive species allows prediction of their potential effects on ecosystems in areas where they are not yet present.

Massachusetts