Wolves in the Great Lakes region: a phylogeographic puzzle

Empirical studies demonstrate that natural hybridization in animals is more common than thought so far ( Mallet 2005 ), particularly among species that originated recently through cycles of population contraction–expansion arising from climate changes over the last glacial period, the Pleistocene. In addition, the post‐glacial global growth of human populations has fostered anthropogenic hybridization events, mediated by habitat changes, the persecution of large predators and the introduction of alien species ( Allendorf et al. 2001 ). The Canis lineage shows cases of both natural and anthropogenic hybridization, exacerbating the controversy about the number of species that should be formally validated in the taxonomic lists, the evolutionary role of genetic introgression and the ways to manage hybrids with invading wild or domesticated populations. The study by Wheeldon et al. (2010) , published in this issue of Molecular Ecology, adds a new piece to the intricate puzzle of evolution and taxonomy of Canis in North America. They show that sympatric wolves (C. lupus) and coyotes (C. latrans) are not (extensively) hybridizing in the western North American Great Lakes region (GLR). Widespread hybridization between coyotes and a genetically distinct, but closely related, wolf‐like population (the eastern wolf) occurred in the northeastern regions of North America. In Wheeldon et al.’s (2010) opinion, these data should prove definitely that two different species of wolf (the western gray wolf C. lupus and the eastern wolf C. lycaon) and their hybrids are distributed across the GLR.     

Biography of a "Feathered Pig": The California Condor Conservation Controversy

In the early 20th century, after hundreds of years of gradual decline, the California condor emerged as an object of intensive scientific study, an important conservation target, and a cultural icon of the American wilderness preservation movement. Early condor researchers generally believed that the species' survival depended upon the preservation of its wilderness habitat. However, beginning in the 1970s, a new generation of scientists argued that no amount of wilderness could prevent the condor's decline and that only intensive scientific management - including captive breeding - could save the species from certain extinction. A bitter and highly politicized battle soon developed over how to best preserve the condor. For 5years the condor was extinct in the wild; however, by the time that officials released the first captive-bred birds the condor recovery program had garnered widespread public support, even among its former critics. Today, condor advocates from the scientific and activist communities work together to manage the species and protect its habitat. The condor's story illustrates some of the tensions, problems, and successes that have accompanied the rise of conservation biology as a scientific field and environmental movement in the United States.     

Molecules and beyond: assessing the distinctness of the Great Lakes wolf

The dog family, Canidae, is a widely distributed group of species that have evolved and radiated relatively recently into 16 genera and 36 recognized species ( Nowak 1999 ). Specific taxonomic designations for some canid taxa can be unclear due to frequent interspecific hybridization among species in both historical and contemporary times, and our imperfect molecular genetic approaches for determining among a series of hypotheses regarding hybridization and evolution. In this issue of Molecular Ecology , Koblmüller et al . tackle the difficult topic of Great Lakes wolf taxonomy and present data that suggest this taxon is currently genetically distinct despite a long history of human persecution and hybridization with related taxa.     

Genetic nature of eastern wolves: Past, present and future

Eastern North American wolves have long been recognized as morphologically distinct from both coyotes and gray wolves. This has led to questions regarding their origins and taxonomic status. Eastern wolves are mainly viewed as: (1) a smaller subspecies of gray wolf (Canis lupus lycaon), potentially the result of historical hybridization between gray wolves (C. lupus) and red wolves (C. rufus), (2) a hybrid, the result of gray wolf (C. lupus) and coyote (C. latrans) interbreeding, or (3) a distinct species, C. lycaon, closely related to the red wolf (C. rufus). Although debate persists, recent molecular studies suggest that the eastern wolf is not a gray wolf subspecies, nor the result of gray wolf/coyote hybridization. Eastern wolves were more likely a distinct species, C. lycaon, prior to the eastward spread of coyotes in the late 1800s. However, contemporary interbreeding exits between C. lycaon to both C. lupus and C. latrans over much of its present range complicating its present taxonomic characterization. While hybridization may be reducing the taxonomic distinctiveness of C. lycaon, it should not necessarily be viewed as negative influence. Hybridization may be enhancing the adaptive potential of eastern wolves, allowing them to more effectively exploit available resources in rapidly changing environments.     

Genetic identification of immigrants to the Scandinavian wolf population

Continued gene flow is fundamental to the survival of small, isolated populations. However, geography and human intervention can often act contrary to this requirement. The Scandinavian wolf population is threatened with a loss of genetic variation yet limited in the accessibility to new immigrants by the geographical distance of this peninsular population from its nearest neighbouring population and by human reluctance to allow wolves in the northern reindeer-breeding areas. In this study, we describe the identification of immigrants into this population using autosomal microsatellites, and maternally inherited mtDNA. Samples of 14 wolves collected in the “dispersal corridor” in northern Sweden in 2002–2005 were compared with 185 resident Scandinavian wolves and 79 wolves from the neighbouring Finnish population. We identified four immigrant wolves, suggesting some westward migration, although only one of these is likely to still survive. The integration of such immigrants into the breeding population is necessary to assure the long-term survival of this isolated and inbred population and highlights the importance of genetics techniques to the management of threatened populations.     

An efficient method for screening faecal DNA genotypes and detecting new individuals and hybrids in the red wolf (<Emphasis Type="Italic">Canis rufus</Emphasis>) experimental population area

Previously, sequencing of mitochondrial DNA (mtDNA) from non-invasively collected faecal material (scat) has been used to help manage hybridization in the wild red wolf (Canis rufus) population. This method is limited by the maternal inheritance of mtDNA and the inability to obtain individual identification. Here, we optimize the use of nuclear DNA microsatellite markers on red wolf scat DNA to distinguish between individuals and detect hybrids. We develop a data filtering method in which scat genotypes are compared to known blood genotypes to reduce the number of PCR amplifications needed. We apply our data filtering method and the more conservative maximum likelihood ratio method (MLR) of Miller et al. (2002 Genetics 160:357–366) to a scat dataset previously screened for hybrids by sequencing of mtDNA. Using seven microsatellite loci, we obtained genotypes for 105 scats, which were matched to 17 individuals. The PCR amplification success rate was 50% and genotyping error rates ranged from 6.6% to 52.1% per locus. Our data filtering method produced comparable results to the MLR method, and decreased the time and cost of analysis by 25%. Analysis of this dataset using our data filtering method verified that no hybrid individuals were present in the Alligator River National Wildlife Refuge, North Carolina in 2000. Our results demonstrate that nuclear DNA microsatellite analysis of red wolf scats provides an efficient and accurate approach to screen for new individuals and hybrids.     

Foliar regeneration of the endangered Red Vanda,Renanthera imschootiana Rolfe (Orchidaceae)

An in vitro method was developed to regenerate large numbers of phenotypically uniform plants from the basal parts of the leaves of flowering plants of Renanthera imschootiana Rolfe. Differentiation of up to 10 shoot buds free of callus and protocorm-like bodies occurred in 10–12 weeks from the base of a single leaf implanted in Mitra et al. (1976) medium supplemented with 2% sucrose, 2 g l^-1 peptone, 44.4 M benzyladenine (BA) and 10.7 M naphthaleneacetic acid (NAA). Subculture of the tissues in medium enriched with 10% coconut water and 35 g l^-1 ripe banana pulp resulted in the production of highest average number of 40 shoots in 12 weeks. No difference in the regeneration potential was observed among the three young leaves while mature leaves did not respond. All the leaves of the regenerated shoots were easily recultured to increase shoot multiplication. Shoots readily formed roots on transfer to a medium containing 4.4 M BA, 10.7 M NAA and 1% activated charcoal. All regenerated plants examined were normal diploids with 2n=38. Foliar meristem culture appears to have great potential for ex situ conservation and propagation of this extremely endangered orchid.Abbreviations BA benzyladenine - NAA naphthaleneacetic acid     

Shoot, shovel and shut up: cryptic poaching slows restoration of a large carnivore in Europe

Poaching is a widespread and well-appreciated problem for the conservation of many threatened species. Because poaching is illegal, there is strong incentive for poachers to conceal their activities, and consequently, little data on the effects of poaching on population dynamics are available. Quantifying poaching mortality should be a required knowledge when developing conservation plans for endangered species but is hampered by methodological challenges. We show that rigorous estimates of the effects of poaching relative to other sources of mortality can be obtained with a hierarchical state-space model combined with multiple sources of data. Using the Scandinavian wolf (Canis lupus) population as an illustrative example, we show that poaching accounted for approximately half of total mortality and more than two-thirds of total poaching remained undetected by conventional methods, a source of mortality we term as 'cryptic poaching'. Our simulations suggest that without poaching during the past decade, the population would have been almost four times as large in 2009. Such a severe impact of poaching on population recovery may be widespread among large carnivores. We believe that conservation strategies for large carnivores considering only observed data may not be adequate and should be revised by including and quantifying cryptic poaching.     

Experimental Extinction of Garlic Mustard (Alliaria petiolata) Populations: Implications for Weed Science and Conservation Biology

An assumption of weed science and conservation biology is that small populations are more vulnerable to elimination and extinction than large populations. We tested this with the invasive biennial garlic mustard (Alliaria petiolata). We compared 61 experimental populations from which every flowering plant was removed for 4 years, with 56 control populations. Whereas the majority of the control populations continued to expand in size over the 4 years, experimental populations showed a strong experimental effect, remaining stable in size, declining in size, or going extinct. Small populations were far more vulnerable to extinction than large populations: 43% of small experimental populations (initially fewer than 10 individuals) went extinct, but only 7% of large populations (initially more than 50 individuals). However, some small experimental populations persisted, and in a few cases, larger experimental populations continued to expand even though every flowering individual had been removed. These results and a simple population model suggest the importance of buried seeds in allowing this species to persist despite attempts to eradicate it.     

Poxviral Disease in Red Squirrels Sciurus vulgaris in the UK: Spatial and Temporal Trends of an Emerging Threat

The squirrel poxvirus (SQPV) is the probable mediator of apparent competition between the introduced invading gray squirrel (Sciurus carolinensis) and the red squirrel (Sciurus vulgaris) in the UK, and modeling studies have shown that this viral disease has had a significant impact on the decline of the red squirrel in the UK. However, given our limited understanding of the epidemiology of the disease, and more generally the effects of invasive species on parasite ecology, there is a need to investigate the transmission dynamics and the relative pathogenicity of the virus between species. We aimed to increase our knowledge of these processes through an empirical study in which we: (i) used pathological signs and transmission electron microscopy (TEM) to diagnose SQPV disease in red squirrels found dead during scanning surveillance between 1993 and 2005; (ii) detected antibody to SQPV using an enzyme-linked immunosorbent assay (ELISA) in the same animals; and (iii) mapped cases of the disease, and the gray squirrel distribution, using a geographical information system. We analyzed the distribution of cases of SQPV disease according to woodland type, a measure of squirrel density. SQPV disease occurred only in areas of England also inhabited by seropositive gray squirrels, and as the geographical range of gray squirrels expanded, SQPV disease occurred in these new gray squirrel habitats, supporting a role for the gray squirrel as a reservoir host of the virus. There was a delay between the establishment of invading gray squirrels and cases of the disease in red squirrels which implies gray squirrels must reach a threshold number or density before the virus is transmitted to red squirrels. The spatial and temporal trend in SQPV disease outbreaks suggested that SQPV disease will have a significant effect on Scottish populations of red squirrels within 25 years. The even spread of cases of disease across months suggested a direct rather than vector-borne transmission route is more likely. Eight juvenile and sub-adult free-living red squirrels apparently survived exposure to SQPV by mounting an immune response, the first evidence of immunity to SQPV in free-living red squirrels, which possibly suggests a changing host-parasite relationship and that the use of a vaccine may be an effective management tool to protect remnant red squirrel populations.     

Microsatellite variation within and among recently fragmented populations of the golden lion tamarin (Leontopithecus rosalia)

Four variable microsatellite loci were used toexamine the genetic diversity and differentiation of golden lion tamarins (Leontopithecus rosalia) in four populations recently isolated by habitat fragmentation. Using Rst estimates of genetic differentiation, a considerable genetic divergence was detected among these populations, with an averagedifferentiation of 31%. Significant differences in allele number among these populations were found. However, the heterozygosity among these populations was not statistically different. These results suggestthat loss in allele diversity was faster than loss in heterozygosity. Conservation implications, particularly for golden lion tamarins, are then discussed. Loss of allelic diversity might be as serious a concern to endangered species as heterozygosity or inbreeding.     

In vitro micropropagation of Primula scotica: a rare Scottish plant

A simple micropropagation method is reported for Primula scotica, a rare plant, endemic to the North of Scotland. The technique involves the clonal proliferation of seed-derived plantlets on either hormone free tissue culture medium or on medium containing benzyl amino purine and indole acetic acid. Average multiplication rates of 4–6 were obtained for plants grown on plant growth regulator supplemented medium. The micropropagation method was applied to four different clones of P. scotica and clonal differences were observed in relation to media type. Some plants from certain clones did display hyperhydricity, however, this was circumvented by using frequent sub-culturing intervals and transferring the plants to hormone-free medium. Plantlets rooted on both media types and displayed normal, true-to-type rosette morphology. The cultures did not callus and development proceeded via shoot and root production only. The in vitro-grown plants could be transferred to ex vitro conditions and a range of growth substrates were assessed for their efficacy in supporting ex vitro growth, with a view to developing longer-term strategies for the transfer and reintroduction of micropropagated P. scotica plants into natural habitats. The simple method described in this paper may offer the potential of being applied to other endangered Primula spp.     

Harvesting of the Kelp Ecklonia Maxima in South Africa Affects Its Three Obligate, Red Algal Epiphytes

In South Africa, more than 7000 t (f wt) of kelp (Ecklonia maxima) fronds are harvested annually to feed cultured abalone. Carpoblepharis flaccida, Gelidium vittatum and Polysiphonia virgata are conspicuous red algal epiphytes on older kelps and provide habitat and food for numerous animals. Over 4.5 y, we examined the effects of one destructive harvest of E. maxima on these 3 epiphytes. Two 20 × 20 m plots of kelp with similar epiphyte loads were demarcated. In one, all E. maxima sporophytes with stipes longer than 50 cm were harvested. The other plot served as a control. After 2.5 y the biomass of E. maxima in the harvested plot had recovered to control levels, but the epiphyte load (g epiphytes. Kg kelp⁻¹) was statistically lower in the harvested plot after 2.5 and 3.5 y, and only recovered after 4.5 y. While most commercial harvesters cut through the “heads” (primary blades) of the kelp, effectively killing them, a new, non-lethal method removes secondary blades 20–30 cm from their bases, leaving the meristems and primary blades intact. At 5 sites studied, G. vittatum and P. virgata were found almost entirely on stipes and primary blades, and harvesting only distal parts of secondary blades limited losses to about 50% of C. flaccida biomass. To protect epiphytes, non-lethal harvesting is recommended and permanent non-harvest zones have been established in addition to limiting kelp yields and disallowing harvesting in Marine Protected Areas.     

Habitat use of juvenile white sturgeon in the Kootenai River, Idaho and British Columbia

Ultrasonic telemetry was used to assess habitat features utilized by 36 endangered juvenile white sturgeon, Acipenser transmontanus, in the lower 120 km of the Kootenai River of Idaho, USA and British Columbia, Canada during the summer and early fall of 1999 and 2000. All 36 fish were initially captured in pools using gillnets and released there, but most of the subsequent telemetry contacts were in glides, indicating these fish moved freely between the two macro-habitats. The low electivity indices indicated little preference between glides and pools. Most contacts were in glides, in the outside bend of the river channel (50), and in or near a visually defined thalweg. Contacts were most often associated with sand substrates and no cover. Physical habitat characteristics (nose [bottom] water velocity, depth, substrate, and cover) were recorded at 168 contact locations. The combination of significantly greater velocities and depths at contact sites vs. non-contact sites (p < 0.01) indicated these fish actively found and used areas of higher velocity and greater depth within the Kootenai River. There was little cover found for fish in the river other than large sand dunes and depth. The combination of depth and nose velocity data supported the idea that large sand dunes are providing refugia in the form of velocity breaks.     

A new species of Acanthosyris (Santalaceae) from Bolivia and a key to the woody South American Santalaceae

Acanthosyris asipapote, a new species from the vicinity of Santa Cruz, Bolivia, is described and illustrated. It is most similar to A. paulo-alvinii Barroso, known only from the state of Bahia, Brazil, but differs by leaf characters, pilose filaments, and smaller seeds. A key to the five woody genera and nine species of Santalaceae in South America is provided. Cervantesia, Jodina, and Myoschilos are probably monotypic; Santalum (in South America) has a single species, and Acanthosyris has five species.     

The Chilean coastal range: a vanishing center of biodiversity and endemism in South American temperate rainforests

Temperate forests of southern South America are globally important because of their high level of endemism. I argue here that within southern South America, rainforests of the Chilean Coastal Range should be the primary target for new conservation efforts. Historically, most protected areas in southern South America have been designated to preserve forests above 600 m, mainly in the Andes. However, Coastal Range forests have higher species richness and are under greater threat than Andean forests at similar latitudes. Coastal forests are characterized by the presence of numerous narrow-range endemics, among them two monotypic plant families. The higher frequency of endemic species in Coastal Range forests is attributed to its more stable biogeographic history compared to the Andes, particularly during the glacial events of the Quaternary. Due to the extent of human impact, the remaining fragments of coastal forests are quite distant from one another, and are disconnected from the larger protected areas of Andean forests. Only 439000 ha of Valdivian coastal forest still remain, including some remnants of primary forest. New private or public reserves should be created to protect the last remnants of continuous forest remaining on the Coastal Range of the Valdivian region (40–42° S). A different conservation strategy should be applied north of 40° S, where protected areas are too small and fragments are too scattered to maintain viable populations of most vertebrates. In this latter area, I recommend expanding existing reserves, restoring native forests, and interconnecting remnant forests through a corridor network.     

Linking fisheries management and conservation in bioengineering species: the case of South American mussels (Mytilidae)

We examined a complete list of South American mussels (Mytilidae) to identify species with current or potential needs for management and conservation actions. Based on ecological/ecosystem (aggregations, beds or banks affecting ecosystem functioning) and socio-economic (artisanal fisheries or aquaculture systems) attributes species with high relevance were identified. At least 14 species exhibited large ecosystem level effects at local scales. Further, most of them also sustain important fisheries: nearly one/third of these fisheries showed characteristics that may contribute to their lack of sustainability and overexploitation, while half are either in the initial exploitation phase or in the stabilization of extraction and institutionalization phase. Invading species are modifying the structure of mussel habitats. Allocation of spatially explicit management tools, notably Territorial User Rights in Fisheries and Marine Reserves, together with co-management initiatives, are suggested as relevant tools to fulfill management and conservation objectives for these key bioengineering species.     

Patterns of allozyme variation in relation to population size of the threatened plantMegaleranthis saniculifolia (Ranunculaceae) in Korea

Patterns of variation at 27 allozyme loci were investigated in the endangered endemic plantMegaleranthis saniculifolia. Levels of allozyme variation (A = 1.47,P = 40%,He = 0.088) were also compared with other endemic plant species. Genetic divergence between populations was very high (G ^st = 0.271 ), with moderate to high interpopulation differentiation, which probably arose through historical bottlenecks in a landscape of habitat fragmentation and/or human influence. The percentage of polymorphic loci, heterozygosity, and mean number of alleles per locus were positively related to population size, probably due to the stochastic loss of rare alleles in the smaller populations. Individuals in the small and marginal populations (TB, KD, and CJ) showed higher proportions of fixed loci. These ecologically marginal populations were typically more distant from the nearest neighboring population and were more genetically distinct from one another. The genetic structure of the current population ofM. saniculifolia is probably the result of local extinctions of intervening populations. This, in turn, is due to the Pleistocene climatic change and increased habitat destruction. A positive association appears to exist between genetic diversity and population size. Although these small population sizes are more sensitive to stochastic events, securing a certain number of individuals from the three larger populations (SB, JB, and TG) could be accomplished as part of a conservation strategy. In addition, it is important to prioritize populations in different regions in order to limit population declines caused by large-scale environmental catastrophes.