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Paul Keim

Publications and source records attributed to Paul Keim.

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The innate immune response may be important for surviving plague in wild Gunnison's prairie dogs

Prairie dogs (Cynomys spp.) are highly susceptible to Yersinia pestis, with ≥99% mortality reported from multiple studies of plague epizootics. A colony of Gunnison's prairie dogs (Cynomys gunnisoni) in the Aubrey Valley (AV) of northern Arizona appears to have survived several regional epizootics of plague, whereas nearby colonies have been severely affected by Y. pestis. To examine potential mechanisms accounting for survival in the AV colony, we conducted a laboratory Y. pestis challenge experiment on 60 wild-caught prairie dogs from AV and from a nearby, large colony with frequent past outbreaks of plague, Espee (n = 30 per colony). Test animals were challenged subcutaneously with the fully virulent Y. pestis strain CO92 at three doses: 50, 5,000, and 50,000 colony-forming units (cfu); this range is lethal in black-tailed prairie dogs (Cynomys ludovicianus). Contrary to our expectations, only 40% of the animals died. Although mortality trended higher in the Espee colony (50%) compared with AV (30%), the differences among infectious doses were not statistically significant. Only 39% of the survivors developed moderate to high antibody levels to Y. pestis, indicating that mechanisms other than humoral immunity are important in resistance to plague. The ratio of neutrophils to lymphocytes was not correlated with plague survival in this study. However, several immune proteins with roles in innate immunity (VCAM-1, CXCL-1, and vWF) were upregulated during plague infection and warrant further inquiry into their role for protection against this disease. These results suggest plague resistance exists in wild populations of the Gunnison's prairie dog and provide important directions for future studies.

Arizona

Winter distribution of willow flycatcher subspecies

Documenting how different regions across a species' breeding and nonbreeding range are linked via migratory movements is the first step in understanding how events in one region can influence events in others and is critical to identifying conservation threats throughout a migratory animal's annual cycle. We combined two studies that evaluated migratory connectivity in the Willow Flycatcher ( Empidonax traillii ), one using mitochondrial DNA sequences from 172 flycatchers sampled throughout their winter range, and another which examined morphological characteristics of 68 museum specimens collected in the winter range. Our results indicate that the four subspecies occupy distinct but overlapping regions of the winter range. Connectivity between specific breeding and winter grounds appears to be moderate to strong, with distributions that suggest migration patterns of both the chain and leap-frog types connecting the breeding and nonbreeding grounds. The Pacific lowlands of Costa Rica appear to be a key winter location for the endangered Southwestern Willow Flycatcher ( E. t. extimus ), although other countries in Central America may also be important for the subspecies.

The Condor

Population differences in host immune factors may influence survival of Gunnison's prairie dogs ( Cynomys Gunnisoni ) during plague outbreaks

Over the past 40 yr, epizootics of plague ( Yersinia pestis ) in northern Arizona have reduced populations of the Gunnison’s prairie dog ( Cynomys gunnisoni ), with the exception of a large population found in the Aubrey Valley (AV). To examine potential mechanisms accounting for their survival, we collected prairie dog serum samples in 2005–2006 from AV and a neighboring population near Seligman (SE), Arizona. We quantified gene expression at 58 diverse immune proteins using a multiplexed enzyme-linked immunosorbent assay panel. We found a subset of proteins important in coagulation and inflammation (tissue factor [TF], calbindin [Cal], and thrombopoietin [TPO]) and T-cell responses (CD40L and CD40) that were present in AV at levels two to eight times greater than SE. These results suggest that AV and SE animals might differ in their ability to mount an immune response.

Arizona

Using Molecular Genetic Markers to Resolve a Subspecies Boundary: The Northern Boundary of the Southwestern Willow Flycatcher in the Four-Corner States

*Executive Summary* The northern boundary of the endangered Southwestern Willow Flycatcher (Empidonax traillii extimus) is currently approximated as running through southern Colorado and Utah, but the exact placement is uncertain because this subspecies shares a border with the more northern and non-endangered E. t. adastus. To help resolve this issue, we evaluated the geographic distribution of mitochondrial and nuclear DNA by sampling breeding sites across the four-corner states (Arizona, Colorado, New Mexico, and Utah). We found that breeding sites clustered into two major groups generally consistent with the currently designated boundary, with the exception of three sites situated along the current boundary. However, delineating a precise boundary that would separate the two subspecies is made difficult because (1) we found evidence for a region of intergradation along the boundary area, suggesting the boundary is not discreet, and (2) the boundary region is sparsely populated, with too few extant breeding populations to precisely locate a boundary. The boundary region encompasses an area where elevation changes markedly over relatively short distances, with low elevation deserts to the south and more mesic, higher elevation habitats to the north. We hypothesized that latitudinal and elevational differences and their concomitant ecological effects could form an ecological barrier that inhibited gene flow between the subspecies, forming the basis for the subspecies boundary. We modeled changes in geographic patterns of genetic markers as a function of latitude and elevation finding significant support for this relationship. The model was brought into a GIS environment to create multiple subspecies boundaries, with the strength of each predicted boundary evaluated on the basis of how much genetic variation it explained. The candidate boundary that accounted for the most genetic variation was situated generally near the currently recognized subspecies boundary, but should be more biologically meaningful because it incorporates the landscape features that may be driving separation of the subspecies. Even so, we caution that using any narrow boundary line as an indicator of subspecies identity could be misleading because biologically the boundary is a region of intergradation rather than a discrete line. Designating, a boundary ultimately becomes a regulatory and management decision based on how much of the genetic variation unique to a subspecies should be protected. We discuss how the results of this study can help guide this decision process by wildlife policy makers.

Open-File Report

Genetic variation in the endangered Southwestern Willow Flycatcher

The Southwestern Willow Flycatcher ( Empidonax traillii extimus ) is an endangered Neotropical migrant that breeds in isolated remnants of dense riparian habitat in the southwestern United States. We estimated genetic variation at 20 breeding sites of the Southwestern Willow Flycatcher (290 individuals) using 38 amplified fragment length polymorphisms (AFLPs). Our results suggest that considerable genetic diversity exists within the subspecies and within local breeding sites. Statistical analyses of genetic variation revealed only slight, although significant, differentiation among breeding sites (Mantel's r = 0.0705, P < 0.0005; θ = 0.0816, 95% CI = 0.0608 to 0.1034; Φ ST = 0.0458, P < 0.001). UPGMA cluster analysis of the AFLP markers indicates that extensive gene flow has occurred among breeding sites. No one site stood out as being genetically unique or isolated. Therefore, the small level of genetic structure that we detected may not be biologically significant. Ongoing field studies are consistent with this conclusion. Of the banded birds that were resighted or recaptured in Arizona during the 1996 to 1998 breeding seasons, one-third moved between breeding sites and two-thirds were philopatric. Low differentiation may be the result of historically high rangewide diversity followed by recent geographic isolation of breeding sites, although observational data indicate that gene flow is a current phenomenon. Our data suggest that breeding groups of E. t. extimus act as a metapopulation.

The Auk