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Prevention, early detection and containment of invasive, nonnative plants in the Hawaiian Islands: current efforts and needs

Introduction: Invasive, non-native plants (or environmental weeds) have long been recognized as a major threat to the native biodiversity of oceanic islands (Cronk & Fuller, 1995; Denslow, 2003). Globally, several hundred non-native plant species have been reported to have major impacts on natural areas on oceanic islands (Kueffer et al ., 2009). In Hawaii, at least some 50 non-native plant species reach dominance in natural areas (Kueffer et al ., 2009) and many of them are known to impact ecosystem processes or biodiversity. One example is the invasive Australian tree fern ( Cyathea cooperi ), which has been shown to be very efficient at utilizing soil nitrogen and can grow six times as rapidly in height, maintain four times more fronds, and produce significantly more fertile fronds per month than the native Hawaiian endemic tree ferns, Cibotium spp. (Durand & Goldstein, 2001a, b). Additionally, while native tree ferns provide an ideal substrate for epiphytic growth of many understory ferns and flowering plants, the Australian tree fern has the effect of impoverishing the understory and failing to support an abundance of native epiphytes (Medeiros & Loope, 1993). Other notorious examples of invasive plant species problematic for biodiversity and ecosystem processes in Hawaii include miconia ( Miconia calvescens ), strawberry guava ( Psidium cattleianum ), albizia ( Falcataria moluccana ), firetree ( Morella faya ), clidemia ( Clidemia hirta ), kahili ginger ( Hedychium gardnerianum ), and fountain grass ( Pennisetum setaceum ), to name just a few. Fireweed ( Senecio madagascariensis ) is a recent example of a seriously problematic invasive species for Hawaii’s agriculture and is damaging certain high-elevations native ecosystems as well. The threat of invasive plants has long been recognized in Hawaii and is well documented (e.g. Cox, 1999; Loope & Kraus, 2009 in press; Loope et al ., 2004; Mooney & Drake, 1986; Stone & Scott, 1985; Stone et al. , 1992). In many respects, Hawaii may be near the forefront among national and international efforts to address the burgeoning threat of invasive plants, perhaps especially in the field of outreach and education (Holt, 1996; Van Driesche & Van Driesche, 2000). However, given the scale of the problem many challenges still need to be addressed and gaps in the existing management system need to be identified. In particular, it appears that new non-native plant species are still introduced to the Hawaiian Islands at a high rate with little or no regard for their potential invasiveness. In fact, a Pacific-wide and a global survey of non-native plants on oceanic islands have both shown that on Hawaii among all archipelagos by far the highest number of problematic invasive species known from other areas in the world is already present (Denslow et al . 2009, Kueffer et al . 2009). Hawaii lacks an effective mechanism for tracking what species are present or incoming. For instance, early detection nursery surveys conducted on Maui in 2008 found over 300 species of cultivated vascular plants that have not previously been recorded in Hawaii (Starr et al. , in prep.). In spite of an innovative Hawaii Biological Survey (e.g. Eldredge & Evenhuis, 2003), there is no mechanism for recording presence of a species until it becomes naturalized. Some of these new introductions may quickly become serious pests. Fireweed, first recorded in Hawaii on the Big Island in the early 1980s, is now considered one of the Kueffer & Loope 2009 5/48 worst weeds of pastures and is also invading natural areas from near sea level to above 10,000 feet. Although the cultivated and as yet non-invasive Cortaderia selloana has been present in Hawaii for 50 years or more, the morphologically similar Cortaderia jubata was simultaneously found to be present on Maui and invading on a large scale in 1989. It played an important role in inspiring the establishment of the Maui Invasive Species Committee (MISC) in 1997, and MISC now spends roughly $200,000 per year removing and containing C. jubata to keep it from becoming widespread in high elevation conservation lands of East and West Maui. The existence of many similar examples shows that to date regulatory action to prevent new invasive plant species from establishing and spreading in Hawaii has not yet been as successful as it needs to be. In particular, because some problematic invasive species known from other areas in the world (Kueffer et al ., 2009; Weber, 2003) have not yet been recorded from Hawaii, preventive measures against the introduction and spread of such likely invasive species is therefore an urgent need for Hawaii. Indeed, regulation of importation and early detection and eradication of introduced species before they become abundant and widespread are widely considered the most cost-efficient and often only effective measures against the threat of new invasive species (Kueffer & Hirsch Hadorn, 2008; Wittenberg & Cock, 2001). Timing seems favorable for Hawaii to achieve effective protection against the threat of new invasive species through prevention, early detection, and eradication/containment. Through the establishment and evolution of Invasive Species Committees (ISCs) on each major Hawaiian island, the institutional capacity has been built up for prevention, early detection, containment, and outreach at an island scale. Weed risk assessment (Daehler et al ., 2004) and early detection methodologies (Starr et al. , in review-a, b) have been developed and tested specifically for Hawaii. Containment strategies have been successful (e.g., Special Ecological Areas in Hawaii Volcanoes National Park), and so have eradications of particular species on an island scale (e.g. mullein ( Verbascum thapsus ) and other species on Maui, fireweed ( Senecio madagascariensis ) on Kauai). These successful management strategies may be further strengthened through recently developed novel approaches in research (e.g. remote sensing, species distribution modelling, and molecular genetics tools). Another major recent achievement is the gained support of the plant industry for preventive measures against invasive species (see p. 13ff). Last but not least, regulatory action is also moving forward. Passage of House Bill 2517 by the 2008 Hawaii House and Senate and prompt signing of the bill into law by the Governor provides hope that action to ban the sale of a meaningful suite of restricted weeds can quickly proceed through the rulemaking phase into the implementation phase. This report documents these achievements and experiences and provides a range of perspectives on how to further develop prevention, early detection and containment of invasive species in Hawaii. The report is based on a symposium and workshop held at the 2008 Hawaii Conservation Conference in Honolulu on 31 July 2008.

Hawaii

Demographic and temporal variations in immunity and condition of polar bears (Ursus maritimus) from the southern Beaufort Sea

Assessing the health and condition of animals in their natural environment can be problematic. Many physiological metrics, including immunity, are highly influenced by specific context and recent events to which researchers may be unaware. Thus, using a multifaceted physiological approach and a context-specific analysis encompassing multiple time scales can be highly informative. Ecoimmunological tools in particular can provide important indications to the health of animals in the wild. We collected blood and hair samples from free-ranging polar bears (Ursus maritimus) in the southern Beaufort Sea and examined the influence of sex, age, and reproductive status on metrics of immunity, stress, and body condition during 2013–2015. We examined metrics of innate immunity (bactericidal ability and lysis) and stress (hair cortisol, reactive oxygen species, and oxidative barrier), in relation to indices of body condition considered to be short term (urea to creatinine ratio; UC ratio) and long term (storage energy and body mass index). We found the factors of sex, age, and reproductive status of the bear were critical for interpreting different physiological metrics. Additionally, the metrics of body condition were important predictors for stress indicators. Finally, many of these metrics differed between years, illustrating the need to examine populations on a longer time scale. Taken together, this study demonstrates the complex relationship between multiple facets of physiology and how interpretation requires us to examine individuals within a specific context.

Alaska

Assessing the Feasibility of Reintroducing San Francisco Gartersnakes ( Thamnophis sirtalis tetrataenia ) to La Honda Creek Open Space Preserve, San Mateo County, California

Reintroductions are used worldwide to increase the viability of species and restore native ecological communities. The success of reintroductions is usually judged by the establishment of self-sustaining populations, restoration of naturally occurring ecological communities, and the species resuming its ecological function. Recovery for the endangered San Francisco gartersnake (SFGS, Thamnophis sirtalis tetrataenia ), a subspecies with a small range in San Mateo and Santa Cruz counties in California, will likely require reintroduction and establishment of new populations within its historical range. La Honda Creek Open Space Preserve (LHC), managed by the Midpeninsula Regional Open Space District (MROSD), is one potential site for the reintroduction of SFGS. The La Honda Creek Open Space Preserve is a preserve managed for wildlife, recreation, grazing, and agriculture located near extant populations of SFGS inhabiting other open space preserves managed by MROSD (Cloverdale Ranch Open Space Preserve [CR]; Russian Ridge Open Space Preserve [RR]). We compared the habitat and prey communities at LHC to nearby open space preserves that support extant SFGS populations. Based on pond surveys done annually since 2008, the occurrence of California red-legged frogs ( Rana draytonii ), Sierran chorus frogs ( Pseudacris sierra ), and Pacific newts ( Taricha spp.) at LHC indicates a similar prey community at this preserve to those at CR and RR. Likewise, the landscape at LHC is a similar mosaic of wetlands, open grassland, shrub-dominated scrub, and coast redwood ( Sequoia sempervirens ) and Douglas fir ( Pseudotsuga menziesii ) forest that meets the habitat requirements for the life history of SFGS at CR and RR. One difference between LHC and preserves with SFGS populations is the lack of vegetative cover immediately adjacent to some wetlands at LHC, which could affect the ability of SFGS to disperse from wetlands and find terrestrial refuges. To evaluate alternative reintroduction strategies, we simulated population viability for a fixed number of SFGS released at LHC into one to six subpopulations (where each wetland represents a subpopulation) over a period from 5 to 20 years. Population simulations indicated that the highest average viability (in other words, the lowest probability of quasi-extinction) occurred when all SFGS were released into a single subpopulation and releases continued annually for 15 to 20 years. Our results indicate that LHC is a good candidate for reintroducing SFGS with suitable habitat, climate, and prey for this snake subspecies. Supporting SFGS populations at LHC could require habitat management to provide sufficient vegetative cover in the terrestrial environment near wetlands. Maintaining genetic diversity in the reintroduced population will also be paramount to ensure negative effects of inbreeding and homozygosity do not affect population viability.

California

Behavior patterns and fates of adult steelhead, Chinook salmon, and coho salmon released into the upper Cowlitz River Basin, 2005–09 and 2012, Washington

A multiyear radiotelemetry evaluation was conducted to monitor adult steelhead ( Oncorhynchus mykiss ), Chinook salmon ( O. tshawytscha ), and coho salmon ( O. kisutch ) behavior and movement patterns in the upper Cowlitz River Basin. Volitional passage to this area was eliminated by dam construction in the mid-1960s, and a reintroduction program began in the mid-1990s. Fish are transported around the dams using a trap-and-haul program, and adult release sites are located in Lake Scanewa, the uppermost reservoir in the system, and in the Cowlitz and Cispus Rivers. Our goal was to estimate the proportion of tagged fish that fell back downstream of Cowlitz Falls Dam before the spawning period and to determine the proportion that were present in the Cowlitz and Cispus Rivers during the spawning period. Fallback is important because Cowlitz Falls Dam does not have upstream fish passage, so fish that pass the dam are unable to move back upstream and spawn. A total of 2,051 steelhead and salmon were tagged for the study, which was conducted during 2005–09 and 2012, and 173 (8.4 percent) of these regurgitated their transmitter prior to, or shortly after release. Once these fish were removed from the dataset, the final number of fish that was monitored totaled 1,878 fish, including 647 steelhead, 770 Chinook salmon, and 461 coho salmon. Hatchery-origin (HOR) and natural-origin (NOR) steelhead, Chinook salmon, and coho salmon behaved differently following release into Lake Scanewa. Detection records showed that the percentage of HOR fish that moved upstream and entered the Cowlitz River or Cispus River after release was relatively low (steelhead = 38 percent; Chinook salmon = 67 percent; coho salmon = 41 percent) compared to NOR fish (steelhead = 84 percent; Chinook salmon = 82 percent; coho salmon = 76 percent). The elapsed time from release to river entry was significantly lower for NOR fish than for HOR fish for all three species. Tagged fish entered the Cowlitz River in greater proportions than the Cispus River, regardless of origin. We found that 23–47 percent of the HOR fish entered the Cowlitz River and 12–38 percent entered the Cispus River. Similarly, 67–70 percent of the NOR fish entered the Cowlitz River and 38–66 percent entered the Cispus River. These behavioral differences translated into similar differences in fates during the spawning periods as higher percentages of tagged fish were assigned Cowlitz River fates than Cispus River fates. Fallback rates were affected by fish origin and release site. Overall, 12 percent of steelhead, 19 percent of Chinook salmon, and 8 percent of coho salmon fell back downstream of Cowlitz Falls Dam prior to spawning. Fallback rates were lower for fish that were released in the Cowlitz River or the Cispus River than for reservoir-released fish, but statistical comparisons were not robust because of small sample sizes at the river release sites. Fallback rates for fish released at the river release sites were 10 percent lower for steelhead, 4 percent lower for Chinook salmon, and 9 percent lower for coho salmon than for reservoir-released fish. However, fallback rates also were different between HOR and NOR fish. Fallback rates were significantly higher for HOR reservoir-released fish than for NOR reservoir-released fish. This study provided data that were insightful for understanding behavior and movement patterns in the upper Cowlitz River Basin and yielded estimates of fallback rates and fish fates that may be useful for fishery managers in the years to come. Studies from other systems have shown that factors such as prespawn mortality and fallback have resulted in substantial losses to spawning populations where trap-and-haul programs are being used as a restoration tool. Future research in the upper Cowlitz River Basin may use additional telemetry studies, genetic analyses, and spawning ground surveys to provide answers for new questions and to continue to monitor the progress of the reintroduction effort.

Washington

Abundance estimation and conservation biology

Abundance is the state variable of interest in most population–level ecological research and in most programs involving management and conservation of animal populations. Abundance is the single parameter of interest in capture–recapture models for closed populations (e.g., Darroch, 1958; Otis et al., 1978; Chao, 2001). The initial capture–recapture models developed for partially (Darroch, 1959) and completely (Jolly, 1965; Seber, 1965) open populations represented efforts to relax the restrictive assumption of population closure for the purpose of estimating abundance. Subsequent emphases in capture–recapture work were on survival rate estimation in the 1970’s and 1980’s (e.g., Burnham et al., 1987; Lebreton et al.,1992), and on movement estimation in the 1990’s (Brownie et al., 1993; Schwarz et al., 1993). However, from the mid–1990’s until the present time, capture–recapture investigators have expressed a renewed interest in abundance and related parameters (Pradel, 1996; Schwarz & Arnason, 1996; Schwarz, 2001). The focus of this session was abundance, and presentations covered topics ranging from estimation of abundance and rate of change in abundance, to inferences about the demographic processes underlying changes in abundance, to occupancy as a surrogate of abundance. The plenary paper by Link & Barker (2004) is provocative and very interesting, and it contains a number of important messages and suggestions. Link & Barker (2004) emphasize that the increasing complexity of capture–recapture models has resulted in large numbers of parameters and that a challenge to ecologists is to extract ecological signals from this complexity. They offer hierarchical models as a natural approach to inference in which traditional parameters are viewed as realizations of stochastic processes. These processes are governed by hyperparameters, and the inferential approach focuses on these hyperparameters. Link & Barker (2004) also suggest that our attention should be focused on relationships between demographic processes such as survival and recruitment, the two quantities responsible for changes in abundance, rather than simply on the magnitudes of these quantities. They describe a type of Jolly–Seber capture–recapture model that permits inference about the underlying relationship between per capita recruitment rates and survival rates (Link & Barker, this volume). Implementation used Bayesian Markov Chain Monte Carlo methods and appeared to work well, yielding inferences about the relationship between recruitment and survival that were robust to selection of prior distribution. We believe that readers will find their arguments compelling, and we expect to see increased use of hierarchical modeling approaches in capture–recapture and related fields. Otto (presentation without paper) also recommended use of hierarchical models in analysis of multiple data sources dealing with population dynamics of North American mallards. He integrated survival inferences from ringing data, abundance information from aerial survey data, and recruitment information based on age ratios from a harvest survey. He used a Leslie matrix population projection model as an integrating framework and obtained estimates of breeding population size using all data.Otto’s approach also permitted inference about biases in estimated quantities. As with the work of Link & Barker (2004), we find Otto’s recommendation to use hierarchical models to integrate data from multiple sources to be very compelling. Alisauskas et al. (2004) report results of an analysis of capture–recapture data for a askatchewan population of white–winged scoters. They used the approach of Pradel (1996) to estimate population growth rate (See the PDF) directly. Estimates for 1975–1985 were quite low, but estimates for the recent period, 2000–2003,increased to values > 1. Parameter estimates for seniority, survival and per capita recruitment (Pradel, 1996) led to the inference that increased recruitment was largely responsible for the improvements in population status and growth. However, various data sources also indicated that this increase in recruitment was likely a result of increased immigration rather than improved reproduction on the area. This latter inference is important from a conservation perspective in indicating the importance of birds in other locations to growth and health of the study population. Lukacs and Burnham presented material to be published elsewhere that dealt with the use of genetic markers in capture–recapture studies. The data sources for such studies are samples of hair or feces, which are then analyzed using molecular genetic techniques in order to determine individual genotypes with respect to a usually small number of loci. Two types of classification error can arise in such analyses. First, if only a small number of loci is examined, then there may be nonnegligible probabilities that multiple individual animals will have the same genotypes. The second type of error arises during the polymerase chain reaction (PCR) process and can result from failure of alleles to amplify (allelic dropout) or from PCR inhibitors in hair and feces that produce the appearance of false alleles or misprinting (Creel et al., 2003). Lukacs and Burnham developed models that formally incorporate possible misclassification of samples resulting from these errors. These models permit estimation of parameters such as abundance and survival in a manner that properly incorporates this uncertainty of individual identity. We anticipate that noninvasive sampling based on molecular genetic analyses of hair or feces will become extremely important for some species, and that the models of Lukacs and Burnham will become very popular for such analyses. MacKenzie & Nichols (2004) discuss the use of occupancy (proportion of patches or habitat area that is occupied) as a surrogate for abundance. In cases of territorial species and where birds occur at low densities, the number of occupied patches may provide a reasonable estimate of abundance. In other cases, occupancy can be viewed as providing information about one tail of the abundance distribution, P (N = 0). The motivation for considering occupancy as a surrogate for abundance is that occupancy is based on so–called presence–absence surveys that are frequently less expensive of time and effort than methods that estimate abundance directly. We describe one set of models that can be used to estimate occupancy for a single season and another that can be used to estimate parameters such as local probabilities of extinction and colonization that are associated with occupancy dynamics. We outline a possible hybrid approach that combines occupancy data with data on marked individuals in order to betterexplore the mechanisms underlying occupancy dynamics. These five presentations made for an interesting session containing useful information and recommendations for future work. A number of themes connecting these presentations could be emphasized. For example, two of the presentations considered alternatives to standard capture–recapture sampling that can be used to draw inferences about abundance, or a portion of the abundance distribution, with field methods that should be less expensive than usual capture–recapture approaches of handling animals. We believe that the most important theme of the session was the emphasis on the processes responsible for changes in abundance. In particular, we are excited by the potential for using hierarchical models as a means of investigating relationships among vital rates and as a means of combining multiple sources of data relevant to system dynamics. Indeed, we expect the importance of this session theme to be reflected in the content and presentations of the next EURING meeting.

Animal Biodiversity and Conservation

Graphic and algebraic solutions of the discordant lead-uranium age problem

Uranium-bearing minerals that give lead-uranium and lead—lead ages that are essentially in agreement, i.e. concordant, generally are considered to have had a relatively simple geologic history and to have been unaltered since their deposition. The concordant ages obtained on such materials are, therefore, assumed to approach closely the actual age of the minerals. Many uranium-bearing samples, particularly uranium ores, give the following discordant age sequences; Pb 206 U 238 &lt; Pb 207 U 235 &#x2AA1; Pb 207 Pb 206 "> Pb 206 U 238 <Pb 207 U 235 ⪡Pb 207 Pb 206 or, less frequently, Pb 207 Pb 206 &#x2AA1; Pb 207 U 235 &lt; Pb 206 U 238 "> Pb 207 Pb 206 ⪡Pb 207 U 235 <Pb 206 U 238 . These discordant age sequences have been attributed most often to uncertainties in the common lead correction, selective loss of radio-active daughter products, loss or gain of lead or uranium, or contamination by an older generation of radiogenic lead. The evaluation of discordant lead isotope age data may be separated into two operations. The first operation, with which this paper is concerned, is mechanical in nature and involves the calculation of the different possible concordant ages corresponding to the various processes assumed to have produced the discordant ages. The second operation is more difficult to define and requires, in part, some personal judgement. It includes a synthesis of the possible concordant age solutions with other independent geologic and isotopic evidence. The concordant age ultimately chosen as most acceptable should be consistent not only with the known events in the geologic history of the area, the age relations of the enclosing rocks, and the mineralogic and paragenetic evidence, but also with other independent age measurements and the isotopic data obtained on the lead in related or associated non-radioactive minerals. The calculation of the possible concordant ages from discordant age data has been greatly simplified by Wetherill's graphical method of plotting the mole ratios of radiogenic Pb 206 U 238 "> Pb 206 U 238 ( N 206 N 238 "> N 206 N 238 ) vs. radiogenic Pb 207 U 235 "> Pb 207 U 235 ( N 207 N 235 "> N 207 N 235 ) after correcting for the contaminating common Pb 206 and Pb 207 . The linear relationships noted in this graphical procedure have been extended to plots of the mole ratios of total Pb 206 U 238 "> Pb 206 U 238 ( t N 206 N 238 "> t N 206 N 238 ) vs. total Pb 207 U 235 "> Pb 207 U 235 ( t N 207 N 235 "> t N 207 N 235 ). This modification permits the calculation of concordant ages for unaltered samples using only the Pb 207 Pb 206 "> Pb 207 Pb 206 ratio of the contaminating common lead. If isotopic data are available for two samples of the same age, x and y , from the same or related deposits or outcrops, graphs of the normalized difference ratios [ ( N 206 N 204 )x &#x2212; ( N 206 N 204 )y ( N 238 N 204 )x &#x2212;( N 238 N 204 )y ] vs. [ ( N 207 N 204 )x &#x2212; ( N 207 N 204 )y ( N 235 N 204 )x &#x2212;( N 235 N 204 )y ] "> [(N 206 N 204 )x − (N 206 N 204 )y(N 238 N 204 )x −(N 238 N 204 )y] vs. [(N 207 N 204 )x − (N 207 N 204 )y(N 235 N 204 )x −(N 235 N 204 )y] can give concordant ages corrected for unknown amounts of a common lead with an unknown Pb 207 / Pb 206 ratio. (If thorium is absent the difference ratios may be normalized with the more abundant index isotope, Pb 208 .) Similar plots of tho normalized, difference ratios for three genetically related samples ( x − y ) and( x − z ), will give concordant ages corrected, in addition, for either one unknown period of past alteration or initial contamination by an older generation of radiogenic lead of unknown Pb 207 /Pb 206 ratio. Practical numerical solutions for many of tho concordant age calculations are not currently available. However, the algebraic equivalents of these new graphical methods give equations which may be programmed for computing machines. For geologically probable parameters the equations of higher order have two positive real roots that rapidly converge on the exact concordant ages corrected for original radiogenic lead and for loss or gain of lead or uranium. Modifications of these general age equations expanded only to the second degree have been derived for use with desk calculators. These graphical and algebraic methods clearly suggest both the type and minimum number of samples necessary for adequate mathematical analysis of discordant lead isotope age data. This mathematical treatment also makes it clear that discordant lead isotope data alone cannot provide the basis for the choice of one of the possible concordant age solutions. The new equations, in particular, provide an incentive to improve our physical constants, analytical techniques and sampling methods in order that we may derive all of the useful geologic information that is available in a comprehensive lead isotope age study.

Geochimica et Cosmochimica Acta

The geologic classification of the meteorites

The meteorite classes of Prior and Mason are assigned to three proposed genetic groups on the basis of a combination of compositional, mineralogical, and elemental characteristics: l) the calcium-poor, volatile-rich carbonaceous chondrites and achondrites; 2) the calcium-poor, volatile-poor chondrites (enstatite, bronzite, hypersthene, and pigeonite), achondrites (enstatite, hypersthene, and pigeonite), stonyirons (pallasites, siderophyre), and irons; and, 3) the calcium-rich (basaltic) achondrites. Chondrites are correlated with calcium-poor achondrites and the silicate phase of the pallasitic meteorites on Fe contents of olivine and pyroxene; and with metal of the stony-irons and irons on the basis of trace elements (Ga and Ge). Transitions in structure and texture between the chondrites and achondrites are recognized. The Van Schmus-Wood chemical-petrologic classification of the chondrites has been modified and expanded to a mineralogic-petrologic classification of the chondrites and calcium-poor achondrites. Chondrites apparently are the first rocks of the solar system. Paragenetic and textural relations in the Murray carbonaceous chondrite shed new light on the manner of accretion, and on the character of dispersed solid materials ('dust', and chondrules and metal) that existed in the solar system before accretion. Two pre-accretionary mineral assemblages (components) are recognized in the carbonaceous chondrites and in the unequilibrated volatile-poor chondrites. They are: 1) a 'low temperature' water-, rare gas-, and carbon-bearing component; and, 2) a high temperature anhydrous silicate and metal component. Paragenetic relations indicate that component 2 materials predate chondrite formation. An accretionary assemblage (component 3) also is recognized in the carbonaceous chondrites and in the unequilibrated volatile-poor chondrites. Component 3 consists of very fine grains of olivine and pyroxene, which occur as pervasive disseminations, as small irregular aggregates of grains, and as large subround to round, finely granular accretional chondrules. Evidence in Murray indicates that component 3 silicates precipitated abruptly and at low pressures, possibly from a high temperature gas, in an environment that contained dispersed component 1 and 2 materials. All component 3 aggregates in Murray contain component 1 material, most commonly as flakes, and locally as tiny granules and larger spherules, some of which are hollow and some of which were broken prior to their mechanical incorporation in accretionary chondrules. Accretion may have occurred as ices associated with dispersed water-bearing component 1 materials temporarily melted during the precipitation of component 3 silicates, and then abruptly refroze to form an icy cementing material. Group 1 materials may be cometary, and group 2 materials may be asteroidal. Schematic models are proposed. Evidence is reviewed for the lunar origin of the pyroxeneplagioclase achondrites. On the basis of natural remanent magnetism, it is suggested that the very scarce diopside-olivine achondrites may be samples from Mars. A classification of the meteorite breccias, including the calcium-poor and calcium-rich mesosiderites, and irons that contain silicate fragments, is proposed. A fragmentation history of the meteorites is outlined on the basis of evidence in the polymict breccias, and from gas retention ages in stones and exposure ages in irons. Cometal impacts appear to have caused the initial fragmentation, stud possibly the perturbation of orbits, of two inferred asteroidal bodies (enstatite and bronzite), one and possibly both events occurring before 2000 m.y. ago. Several impacts apparently occurred on the inferred hypersthene body in the interval 1000 to 2000 m.y. ago. Major breakups of the three bodies apparently occurred as the result of interasteroidal collisions at about 900 m.y. ago, and 600 to 700 m.y. ago. The breakups were followed by a number of fr

Open-File Report

Evaluation of fisher (Pekania pennanti) restoration in Olympic National Park and the Olympic Recovery Area: 2014 annual progress report

With the translocation and release of 90 fishers ( Pekania pennanti ) from British Columbia to Olympic National Park during 2008–2010, the National Park Service and Washington Department of Fish and Wildlife accomplished the first phase of fisher restoration in Washington State. Beginning in 2013, we initiated a new research project to determine the current status of fishers on Washington’s Olympic Peninsula 3–5 years after the releases and evaluate the short-term success of the restoration program. Objectives of the study are to determine the current distribution of fishers and proportion of the recovery area that is currently occupied by fishers, determine several genetic characteristics of the reintroduced population, and determine reproductive success of the founding animals through genetic studies. During 2014, we continued working with a broad coalition of cooperating agencies, tribes, and nongovernmental organizations (NGO) to collect data on fisher distribution and genetics using noninvasive sampling methods. The primary sampling frame consisted of 157 24-square-kilometer hexagons (hexes) distributed across all major land ownerships within the Olympic Peninsula target survey area. In 2014 we expanded the study by adding 58 more hexes to an expanded study area in response to incidental fisher observations outside of the target area obtained in 2013; 49 hexes were added south and 9 to the east of the target area. During 2014, federal, state, tribal and NGO biologists and volunteers established three baited motion-sensing camera stations, paired with hair snaring devices, in 80 hexes; 69 in the targeted area 11 in the expansion areas. Each paired camera/hair station was left in place for approximately 6 weeks, with three checks on 2-week intervals. We documented fisher presence in 5 of the 80 hexagons, and identified 5 different fishers through a combination of microsatellite DNA analyses and camera detections. All fisher detections were in the target area. These 5 individuals included 2 of the original founding population of 90, 1 of the 2 rescued and rehabilitated kits that were released in 2010, and 1 new recruit to the population (1 individual was not identified). Additionally, we identified more than 40 other species of wildlife at the baited camera stations. We also obtained eight incidental fisher observations through photographs and carcass retrieval. During 2015, we plan to sample 75 hexagons in the target area and 12 in the expansion area. We plan to sample all unsampled accessible hexes in the target area (26 hexes), and re-sample accessible hexes sampled in 2013 (49 hexes).

Washington

Geology and oil and gas assessment of the Fruitland Total Petroleum System, San Juan Basin, New Mexico and Colorado: Chapter 6 in Geology and Oil and Gas Assessment of the Fruitland Total Petroleum System, San Juan Basin, New Mexico and Colorado

The Fruitland Total Petroleum System (TPS) of the San Juan Basin Province includes all genetically related hydrocarbons generated from coal beds and organic-rich shales in the Cretaceous Fruitland Formation. Coal beds are considered to be the primary source of the hydrocarbons. Potential reservoir rocks in the Fruitland TPS consist of the Upper Cretaceous Pictured Cliffs Sandstone, Fruitland Formation (both sandstone and coal beds), and the Farmington Sandstone Member of the Kirtland Formation, and the Tertiary Ojo Alamo Sandstone, and Animas, Nacimiento, and San Jose Formations. Analysis of the geochemistry of Fruitland coal-bed gas and co-produced water suggests that hydrocarbons in Fruitland coal beds began to form early in the depositional history of the Fruitland Formation with the generation of early microbial gas. Source rocks in the Fruitland entered the oil generation zone in the late Eocene and continued to generate minor oil and large quantities of thermogenic gas into middle Miocene time. Near the end of the Miocene, thermogenic hydrocarbon generation and subsidence in the San Juan Basin ceased, and the basin was uplifted and differentially eroded. Late-stage (secondary) microbial gas has been documented in Fruitland coal-bed reservoirs and was formed by microbial reduction of carbon dioxide during introduction of groundwater in the late Pliocene and Pleistocene. Most of this late-stage microbial gas is found just downdip from the northern, western, and southern Fruitland outcrops. The northern part of the Fruitland Formation is overpressured as a result of artesian conditions established in the Pliocene or Pleistocene. South and east of the overpressured area, the Fruitland is either normally pressured or underpressured. Four assessment units (AU) were defined in the Fruitland TPS. Of the four AUs, one consists of conventional gas accumulations and the other three are continuous-type gas accumulations: Tertiary Conventional Gas AU, Pictured Cliffs Continuous Gas AU, Basin Fruitland Coalbed Gas (CBG) AU, and Fruitland Fairway CBG AU. No oil resources that have the potential for additions to reserves in the next 30 years were estimated for this TPS. Gas resources that have the potential for additions to reserves in the next 30 years are estimated at a mean of 29.3 trillion cubic feet of gas (TCFG). Of this amount, 23.58 TCFG will come from coal-bed gas accumulations and 83.1 percent of this total is estimated to come from the Basin Fruitland CBG AU. The remaining 5.72 TCFG is allocated to continuous-type gas accumulations (5.64 TCFG) and conventional gas accumulations (0.08 TCFG). Although the Fruitland Fairway CBG AU has produced the most significant amount of coal-bed gas to date, the area of the AU is limited. New potentially productive wells will come from infill drilling, and the number of these wells will be limited by effective drainage area. Total natural gas liquids (NGL) that have the potential for additions to reserves in the next 30 years are estimated at a mean of 17.76 million barrels. Of this amount, 16.92 million barrels will come from the Pictured Cliffs Continuous Gas AU and the remainder from the Tertiary Conventional Gas AU.

Colorado;New Mexico

Behavior and movement of adult winter steelhead ( Oncorhynchus mykiss ) in the upper Cowlitz River Basin, Washington, 2017–18

Executive Summary A 2-year radiotelemetry study was completed to monitor the movements of adult winter steelhead ( Oncorhynchus mykiss ) in the upper Cowlitz River Basin. A reintroduction program was established to restore healthy and harvestable populations of steelhead because volitional access to the area was blocked in the 1960s after construction of dams in the lower river. A trap-and-haul program is used to move adult steelhead and salmon ( Oncorhynchus spp.) upstream, around the dams and large reservoirs, and release them in the upper basin to spawn naturally. Fish are released into Lake Scanewa, the uppermost reservoir in the system, and into the Cowlitz and Cispus Rivers. The goal of this study was to describe the behavior, movement, and tributary use of adult steelhead in the upper Cowlitz River Basin to assist in evaluating the trap-and-haul program and reintroduction program. We were specifically interested in learning more about the locations where steelhead spawn. Individual fish were assigned one of four fates, based on their location during the spawning period: Cowlitz River, Cispus River, Lake Scanewa, or fallback below the dam that impounds the reservoir. A total of 215 steelhead were tagged for the 2017–18 study. Of these, 5 fish regurgitated their transmitters before or shortly after release, so 210 tagged fish were used for analyses, including 121 fish (57.6 percent) released into the Cispus River and 89 fish (42.4 percent) released into Lake Scanewa. The Cowlitz River release site was not evaluated. Hatchery-origin (HOR) and natural-origin (NOR) steelhead were included in the study. Within the first 10 days after release, most steelhead had moved at least 2 river kilometers away from their release site. Fish released into Lake Scanewa, however, moved from their release site significantly sooner than fish released into the Cispus River, regardless of origin or sex. Most radio-tagged steelhead (93–100 percent) made no more than one trip between the reservoir and one of the rivers prior to spawning. Steelhead were predominantly assigned fates in the river closest to where they were released, but origin also played a role. Steelhead released into the Cispus River were assigned Cispus River fates more than 80 percent of the time, including both origins and both years. About 13 percent of NOR fish had fates in the Cowlitz River, but the proportion was lower for HOR fish (0–2 percent). Fallback was the least common fate for Cispus-released steelhead (three HOR fish in 2017), followed by the reservoir fate, ranging from 5 to 9 percent in 2017, and zero in 2018. The steelhead released into Lake Scanewa were almost exclusively NOR fish, which had primarily Cowlitz River fates (55–57 percent). We released only six HOR steelhead into Lake Scanewa, and they all had Cowlitz River fates. The reservoir fate was uncommon for both release sites, and it was consistently lower in 2018 compared to 2017. Flow conditions were higher in 2017, which may have affected steelhead movement patterns or timing. The three HOR steelhead released in the Cispus River in 2017 were the only fish that fell back over Cowlitz Falls Dam, which represented 1.4 percent of the total study fish, 5.7 percent of the Cispus-released fish in 2017, and 9.4 percent of the Cispus-released HOR fish in 2017. About 62 percent of the steelhead released in Lake Scanewa spawned in the Cowlitz River, with the remaining 38 percent in the Cispus River. Within the Cowlitz River, close to one-half (about 47 percent) of the fish spawned in the upper reach, and about 16 percent spawned in the lower reach. Steelhead released into Lake Scanewa that spawned in the Cispus River were minimal in reach 5 (4 percent) and distributed in approximately equal proportions in the middle (reach 6; 16 percent) and upper (reach 7; 18 percent) reaches. Steelhead released into the Cispus River spawned almost exclusively in the Cispus River (94 percent). The upper Cispus River reach (reach 7) was used by more fish (72 percent) than the middle Cispus reach (reach 6; 16 percent). Taken together, the upper Cowlitz River reach (reach 4) and upper Cispus River reach (reach 7) accounted for more than 71 percent of the spawning locations for radio-tagged steelhead. When the middle Cispus River reach (reach 6) is included with the upper reaches, they account for 87 percent of our described spawning sites. We described 12 tributaries in the Cowlitz River and 8 tributaries in the Cispus River where steelhead spawned. After spawning, about 53–63 percent of steelhead moved downstream as kelts, either being collected at Cowlitz Falls Dam or being detected downstream from the dam. More fish were collected from both release sites in 2018 compared to 2017. Across release sites and years, more NOR fish and females moved downstream as kelts. This study added to the understanding of the behavior and movement of adult steelhead in the upper Cowlitz River Basin, supporting the findings of previous studies in the basin and describing spawning sites in the system’s two main rivers and their tributaries. Future research efforts in this system may use additional telemetry studies, genetic analyses, and spawning ground surveys to provide further insights into the progress of the reintroduction effort.

Washington

Connectivity of Mojave Desert tortoise populations—Management implications for maintaining a viable recovery network

Executive Summary The historic distribution of Mojave desert tortoises ( Gopherus agassizii ) was relatively continuous across the range, and the importance of tortoise habitat outside of designated tortoise conservation areas (TCAs) to recovery has long been recognized for its contributions to supporting gene flow between TCAs and to minimizing impacts and edge effects within TCAs. However, connectivity of Mojave desert tortoise populations has become a concern because of recent and proposed development of large tracts of desert tortoise habitat that cross, fragment, and surround designated conservation areas. This paper summarizes the underlying concepts and importance of connectivity for Mojave desert tortoise populations by reviewing current information on connectivity and providing information to managers for maintaining or enhancing desert tortoise population connectivity as they consider future proposals for development and management actions. Maintaining an ecological network for the Mojave desert tortoise, with a system of core habitats (TCAs) connected by linkages, is necessary to support demographically viable populations and long-term gene flow within and between TCAs. There are four points for wildlife and land-management agencies to consider when making decisions that could affect connectivity of Mojave desert tortoise populations (for example, in updating actions in resource management plans or amendments that could help maintain or restore functional connectivity in light of the latest information): Management of all desert tortoise habitat for persistence and connectivity . Desert tortoise populations continue to decline within most TCAs, and it is unlikely that trends are better in populations outside protected areas. Fragmentation exacerbates negative population trends by breaking large continuous populations into smaller isolated populations. Connectivity within large populations can enhance resilience to localized disturbances due to rescue by neighboring individuals. In contrast, smaller fragmented populations are resistant to rescue by their isolation and thus could suffer irreversible declines to extirpation from a variety of threats and stochastic events. Enhanced threat reduction to reverse declines within TCAs and to maintain occupied habitat in the surrounding matrix would help reduce the variability in population growth rates and improve the resilience of protected populations even while implementing efforts to improve connectivity. Each TCA has unique strengths and weaknesses regarding its ability to support minimum sustainable populations based on areal extent and its ability to support population increases based on landscape connection with adjacent populations. Considering how proposed projects (inside or outside of TCAs) affect connectivity and the ability of TCAs to support at least 5,000 adult tortoises (the numerical goal for each TCA) could help managers to maintain the resilience of TCAs to population declines. The same project, in an alternative location, could have very different impacts on local and regional populations. For example, within the habitat matrix surrounding TCAs, narrowly delineated corridors may not allow for natural population dynamics if they do not accommodate overlapping home ranges along most of their widths so that tortoises reside, grow, find mates, and produce offspring that can replace older tortoises. In addition, most habitat outside TCAs may receive more surface disturbance than habitat within TCAs. Therefore, managing the entire remaining matrix of desert tortoise habitat for permeability may be better than delineating fixed corridors. These concepts apply, especially given uncertainty about long-term condition of habitat, within and outside of TCAs under a changing climate. Ultimately, questions such as “ What are the critical linkages that need to be protected ?” could be better framed as “ How can we manage the remaining habitat matrix in ways that sustain ecological processes and habitat suitability for special status species ?” Land-management decisions made in the context of the latter question may be more conducive to maintenance of a functional ecological network. Limitations on landscape-level disturbance across habitat managed for the desert tortoise Clearly delineating habitat linkages and differentiating them from non-delineated areas by the uses that are permitted or prohibited within them by specific management guidelines can help achieve functional connectivity. Such guidelines would be most effective if they considered and accounted for all surface disturbances (for example, temporary disturbances such as fiberoptic lines or off-highway vehicle routes, right-of-ways, utility-scale solar development, urbanization) to the extent possible. A weighted framework that varies with the permanence or severity of the disturbance, and can be additive to quantify cumulative effects, could be useful (Xiong, 2020). For example, minor roads can alter tortoise movements independently of other features (Peaden and others, 2017; Hromada and others, 2020), but if the isolated dirt road is accompanied by a powerline that encourages raven predation (Xiong, 2020), then the two features together may be additive. Ignoring minor or temporary disturbance on the landscape could result in a cumulatively large impact that is not explicitly acknowledged (Goble, 2009); therefore, understanding and quantifying all surface disturbance on a given landscape is prudent. In California, the Bureau of Land Management established 0.1–1.0 percent caps on new surface-disturbance for TCAs and mapped linkages that address the issues described in number 1 of this list. Nevada, Utah, and Arizona currently do not have surface-disturbance limits. Limits comparable to those in the Desert Renewable Energy Conservation Plan (DRECP) would be 0.5 percent within TCAs and 1 percent within the linkages modeled by Averill-Murray and others (2013). Limits in some areas of California within the Desert Renewable Energy Conservation Plan, such as Ivanpah Valley, are more restrictive, at 0.1 percent. Continuity across the state line in Nevada could be achieved with comparable limits in the adjacent portion of Ivanpah Valley, as well as the Greater Trout Canyon Translocation Area and the Stump Springs Regional Augmentation Site. These more restrictive limits would help protect remaining habitat in the major interstate connectivity pathway through Ivanpah Valley and focal areas of population augmentation that provide additional population connectivity along the western flank of the Spring Mountains. In a recent study that analyzed 13 years of desert tortoise monitoring data, nearly all desert tortoise observations were at sites in which 5 percent or less of the surrounding landscape within 1 kilometer was disturbed (Carter and others, 2020a). To help maintain tortoise habitability and permeability across all other non-conservation-designated tortoise habitat, all surface disturbance could be limited to less than 5-percent development per square kilometer because the 5-percent threshold for development is the point at which tortoise occupation drops precipitously (Carter and others, 2020a). However, although individual desert tortoises were observed at development levels up to 5 percent, we do not know the fitness or reproductive characteristics of these individuals. This level of development also may not allow for long-term persistence of healthy populations that are of adequate size needed for demographic or functional connectivity; therefore, a conservative interpretation suggests that, ideally, development could be lower. Lower development levels would be particularly useful in areas within the upper 5th percentile of connectivity values modeled by Gray and others (2019). Reducing ancillary threats in places where connectivity is restricted to narrow strips of habitat, for example, narrow mountain passes or vegetated strips between solar development, could enhance the functionality of these vulnerable linkages. In such areas, maintaining multiple, redundant linkages could further enhance overall connectivity. Minimization of mortality from roads and maximization of passage under roads . Roads pose a significant threat to the long-term persistence of local tortoise populations, and roads of high traffic volume lead to severe population declines, which ultimately fragments populations farther away from the roads. Three points (a.–c.) pertain to reducing direct mortality of tortoises on the many paved roads that cross desert tortoise habitat and to maintaining a minimal level of permeability across these roads: Tortoise-exclusion fencing tied into culverts, underpasses, overpasses, or other passages below roads in desert tortoise habitat, would limit vehicular mortality of tortoises and provide opportunities for movement across the roads. Installation of shade structures on the habitat side of fences installed in areas with narrow population-depletion zones would limit overheating of tortoises that may pace the fence. Passages below highways could be maintained or retrofitted to ensure safe tortoise access, for example, by filling eroded drop-offs or modifying erosion-control features such as rip-rap to make them safer and more passable for tortoises. Wildlife management agencies could work with transportation departments to develop construction standards that are consistent with hydrologic/erosion management goals, while also incorporating a design and materials consistent with tortoise survival and passage and make the standards widely available. The process would be most effective if the status of passages was regularly monitored and built into management plans. Healthy tortoise populations along fenced highways could be supported by ensuring that land inside tortoise-exclusion fences is not so degraded that it leads to degradation of tortoise habitat outside the exclusion areas. For example, severe invasive plant infestations inside a highway exclusion could cause an increase of invasive plants outside the exclusion area and degrade habitat; therefore, invasive plants inside road rights of way could be mown or treated with herbicide to limit their spread into adjacent tortoise habitat and minimize the risk of these plants carrying wildfires into adjacent habitat. Adaptation of management based on new information . Future research will continue to build upon and refine models related to desert tortoise population connectivity and develop new ones. New models could consider landscape levels of development and be constructed such that they share common foundations to support future synthesis efforts. If model development was undertaken in partnership with entities that are responsible for management of desert tortoise habitat, it would facilitate incorporation of current and future modeling results into their land management decisions. There are specific topics that may be clarified with further evaluation: The effects of climate change on desert tortoise habitat, distribution, and population connectivity; The effects of large-scale fires, especially within repeatedly burned habitat, on desert tortoise distribution and population connectivity; The ability of solar energy facilities or similar developments to support tortoise movement and presence by leaving washes intact; leaving native vegetation intact whenever possible, or if not possible, mowing the site, allowing vegetation to re-sprout, and managing weeds; and allowing tortoises to occupy the sites; and The design and frequency of underpasses necessary to maintain functional demographic and genetic connectivity across linear features, like highways.

Arizona, California, Nevada