Emergence of Ixodes scapularis (Acari: Ixodidae) in a Small Mammal Population in a Coastal Oak-Pine Forest, Maine, USA.

In the United States, surveillance has been key to tracking spatiotemporal emergence of blacklegged ticks [Ixodes scapularis Say (Ixodida:Ixodidae)] and their pathogens such as Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner (Spirochaetales: Spirochaetaceae), the agent of Lyme disease. On the Holt Research Forest in midcoastal Maine, collection of feeding ticks from live-trapped small mammal hosts allowed us to track the emergence and establishment of I. scapularis, 1989-2019. From 1989-1995, we collected only I. angustus Neumann (Ixodida: Ixodidae)(vole tick), Dermacentor variabilis Say (Ixodida: Ixodidae) (American dog tick), and I. marxi Banks (Ixodida: Ixodidae) (squirrel tick) from seven species of small mammals. The most abundant tick host was the white-footed mouse [Peromyscus leucopus Rafinesque (Rodentia:Cricetidae)] followed by the red-backed vole (Myodes gapperi Vigors (Rodentia: Cricetidae)). Emergence of I. scapularis was signaled via the appearance of subadult I. scapularis in 1996. Emergence of B. burgdorferi was signaled through its appearance in I. scapularis feeding on mice in 2005. There was a substantial increase in I. scapularis prevalence (proportion of hosts parasitized) and burdens (ticks/host) on white-footed mice and red-backed voles in 2007. The ~11-yr time-to-establishment for I. scapularis was consistent with that seen in other studies. White-footed mice comprised 65.9% of all captures and hosted 94.1% of the total I. scapularis burden. The white-footed mouse population fluctuated interannually, but did not trend up as did I. scapularis prevalence and burdens. There were concurrent declines in I. angustus and D. variabilis. We discuss these results in the broader context of regional I. scapularis range expansion.

Frontiers of protected areas versus forest exploitation: Assessing habitat network functionality in 16 case study regions globally.

Exploitation of natural forests forms expanding frontiers. Simultaneously, protected area frontiers aim at maintaining functional habitat networks. To assess net effects of these frontiers, we examined 16 case study areas on five continents. We (1) mapped protected area instruments, (2) assessed their effectiveness, (3) mapped policy implementation tools, and (4) effects on protected areas originating from their surroundings. Results are given as follows: (1) conservation instruments covered 3-77%, (2) effectiveness of habitat networks depended on representativeness, habitat quality, functional connectivity, resource extraction in protected areas, time for landscape restoration, "paper parks", "fortress conservation", and data access, (3) regulatory policy instruments dominated over economic and informational, (4) negative matrix effects dominated over positive ones (protective forests, buffer zones, inaccessibility), which were restricted to former USSR and Costa Rica. Despite evidence-based knowledge about conservation targets, the importance of spatial segregation of conservation and use, and traditional knowledge, the trajectories for biodiversity conservation were generally negative.

A 2.5-million-year perspective on coarse-filter strategies for conserving nature's stage.

Climate change will require novel conservation strategies. One such tactic is a coarse-filter approach that focuses on conserving nature's stage (CNS) rather than the actors (individual species). However, there is a temporal mismatch between the long-term goals of conservation and the short-term nature of most ecological studies, which leaves many assumptions untested. Paleoecology provides a valuable perspective on coarse-filter strategies by marshaling the natural experiments of the past to contextualize extinction risk due to the emerging impacts of climate change and anthropogenic threats. We reviewed examples from the paleoecological record that highlight the strengths, opportunities, and caveats of a CNS approach. We focused on the near-time geological past of the Quaternary, during which species were subjected to widespread changes in climate and concomitant changes in the physical environment in general. Species experienced a range of individualistic responses to these changes, including community turnover and novel associations, extinction and speciation, range shifts, changes in local richness and evenness, and both equilibrium and disequilibrium responses. Due to the dynamic nature of species responses to Quaternary climate change, a coarse-filter strategy may be appropriate for many taxa because it can accommodate dynamic processes. However, conservationists should also consider that the persistence of landforms varies across space and time, which could have potential long-term consequences for geodiversity and thus biodiversity.

Incorporating geodiversity into conservation decisions.

In a rapidly changing climate, conservation practitioners could better use geodiversity in a broad range of conservation decisions. We explored selected avenues through which this integration might improve decision making and organized them within the adaptive management cycle of assessment, planning, implementation, and monitoring. Geodiversity is seldom referenced in predominant environmental law and policy. With most natural resource agencies mandated to conserve certain categories of species, agency personnel are challenged to find ways to practically implement new directives aimed at coping with climate change while retaining their species-centered mandate. Ecoregions and ecological classifications provide clear mechanisms to consider geodiversity in plans or decisions, the inclusion of which will help foster the resilience of conservation to climate change. Methods for biodiversity assessment, such as gap analysis, climate change vulnerability analysis, and ecological process modeling, can readily accommodate inclusion of a geophysical component. We adapted others' approaches for characterizing landscapes along a continuum of climate change vulnerability for the biota they support from resistant, to resilient, to susceptible, and to sensitive and then summarized options for integrating geodiversity into planning in each landscape type. In landscapes that are relatively resistant to climate change, options exist to fully represent geodiversity while ensuring that dynamic ecological processes can change over time. In more susceptible landscapes, strategies aiming to maintain or restore ecosystem resilience and connectivity are paramount. Implementing actions on the ground requires understanding of geophysical constraints on species and an increasingly nimble approach to establishing management and restoration goals. Because decisions that are implemented today will be revisited and amended into the future, increasingly sophisticated forms of monitoring and adaptation will be required to ensure that conservation efforts fully consider the value of geodiversity for supporting biodiversity in the face of a changing climate.

Why geodiversity matters in valuing nature's stage.

Geodiversity--the variability of Earth's surface materials, forms, and physical processes-is an integral part of nature and crucial for sustaining ecosystems and their services. It provides the substrates, landform mosaics, and dynamic physical processes for habitat development and maintenance. By determining the heterogeneity of the physical environment in conjunction with climate interactions, geodiversity has a crucial influence on biodiversity across a wide range of scales. From a literature review, we identified the diverse values of geodiversity; examined examples of the dependencies of biodiversity on geodiversity at a site-specific scale (for geosites <1 km(2) in area); and evaluated various human-induced threats to geosites and geodiversity. We found that geosites are important to biodiversity because they often support rare or unique biota adapted to distinctive environmental conditions or create a diversity of microenvironments that enhance species richness. Conservation of geodiversity in the face of a range of threats is critical both for effective management of nature's stage and for its own particular values. This requires approaches to nature conservation that integrate climate, biodiversity, and geodiversity at all spatial scales.

Improving management of small natural features on private lands by negotiating the science-policy boundary for Maine vernal pools.

Vernal pools are far more important for providing ecosystem services than one would predict based on their small size. However, prevailing resource-management strategies are not effectively conserving pools and other small natural features on private lands. Solutions are complicated by tensions between private property and societal rights, uncertainties over resource location and function, diverse stakeholders, and fragmented regulatory authority. The development and testing of new conservation approaches that link scientific knowledge, stakeholder decision-making, and conservation outcomes are important responses to this conservation dilemma. Drawing from a 15-y history of vernal pool conservation efforts in Maine, we describe the coevolution of pool conservation and research approaches, focusing on how research-based knowledge was produced and used in support of management decisions. As management shifted from reactive, top-down approaches to proactive and flexible approaches, research shifted from an ecology-focused program to an interdisciplinary program based on social-ecological systems. The most effective strategies for linking scientific knowledge with action changed as the decision-makers, knowledge needs, and context for vernal pool management advanced. Interactions among stakeholders increased the extent to which knowledge was coproduced and shifted the objective of stakeholder engagement from outreach to research collaboration and development of innovative conservation approaches. New conservation strategies were possible because of the flexible, solutions-oriented collaborations and trust between scientists and decision-makers (fostered over 15 y) and interdisciplinary, engaged research. Solutions to the dilemma of conserving small natural features on private lands, and analogous sustainability science challenges, will benefit from repeated negotiations of the science-policy boundary.

The complementary niches of anthropocentric and biocentric conservationists.

A divergence of values has become apparent in recent debates between conservationists who focus on ecosystem services that can improve human well-being and those who focus on avoiding the extinction of species. These divergent points of view fall along a continuum from anthropocentric to biocentric values, but most conservationists are relatively closer to each other than to the ends of the spectrum. We have some concerns with both positions but emphasize that conservation for both people and all other species will be most effective if conservationists focus on articulating the values they all share, being respectful of divergent values, and collaborating on common interests. The conservation arena is large enough to accommodate many people and organizations whose diverse values lead them to different niches that can, with good will and foresight, be far more complementary than competitive.

Clear-cutting affects habitat connectivity for a forest amphibian by decreasing permeability to juvenile movements.

Conservation of forest amphibians is dependent on finding the right balance between management for timber production and meeting species' habitat requirements. For many pond-breeding amphibians, successful dispersal of the juvenile stage is essential for long-term population persistence. We investigated the influence of timber-harvesting practices on the movements of juvenile wood frogs (Lithobates sylvaticus). We used a chronosequence of stands produced by clear-cutting to evaluate how stand age affects habitat permeability to movements. We conducted experimental releases of juveniles in 2008 (n = 350) and 2009 (n = 528) in unidirectional runways in four treatments: mature forest, recent clearcut, 11-year-old, and 20-year-old regeneration. The runways were 50 x 2.5-m enclosures extending into each treatment, perpendicular to a distinct edge, with four tracking stations at 10, 20, 30, and 40 m from the edge. We recorded the number of animals reaching each tracking station, and the proportion of animals changing their direction of movement at each distance. We found that the mature forest was 3.1 and 3.7 times more permeable than the 11-year-old regeneration and the recent clearcut, respectively. Animals actively avoided open-canopy habitats and sharp edges; significantly more animals returned toward the closed-canopy forest at 0 m and 10 m in the less permeable treatments. There were no significant differences in habitat permeability between the mature forest and the 20-year-old regeneration. Our study is the first to directly assess habitat permeability to juvenile amphibian movement in relation to various forestry practices. We argue that habitat permeability at this scale is largely driven by the behavior of animals in relation to habitat disturbance and that caution needs to be used when using spatial modeling and expert-derived permeability values to assess connectivity of amphibian populations. The effects of clear-cutting on the migratory success of juvenile L. sylvaticus are long-lasting. Forestry practices that involve canopy removal and conversion of natural forest to conifer plantations may affect regional population viability by hindering successful dispersal.

Terrestrial habitat selection and strong density-dependent mortality in recently metamorphosed amphibians.

To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.

Long-term effects of group-selection timber harvesting on abundance of forest birds.

Relatively few studies have examined the ecological effects of group-selection timber harvesting, and nearly all have been short-term and have lacked experimental manipulations that allow pre- and posttreatment comparisons. We have been documenting the effects of a group-selection timber harvest on bird abundance in a Maine forest for 24 years (preharvest, 1983-1987; postharvest, 1988-2006). Here we characterized the trends in bird abundance over the first 20 years of the study in the managed and control halves of the 40-ha study area. Species responses to the group-selection harvest were idiosyncratic, but in general the mature-forest bird community was retained and species dependent on early successional habitat temporarily (

Reversing the Fragmentation Perspective: Effects of Clearcut Size on Bird Species Richness in Maine.

The effects of clearcut size on bird species richness in a forest-dominated landscape were examined. Forty-five clearcuts, ranging in size from 2 to 112 ha and in age from 3 to 10 yr post-cut were selected as study sites in eastern Maine. At least 10% of each clearcut was censused using 1-16 fixed-radius (50-m) circular plots in May and June of 1989 and 1990. Species richness increased with clearcut size but the number of species present per plot did not differ significantly over the size range of cuts. There was some evidence of increased richness in clearcuts up to 20 ha. Of the 15 most common clearcut species in both years, 10 (1989) and 12 (1990) increased in abundance between 2 and 20 ha, beyond which no preference for clearcut size was found. Additionally bird species locations throughout clearcuts were largely independent of distance from the forest edge.