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Decline of the meadow jumping mouse (Zapus hudsonius luteus) in two mountain ranges in New Mexico.

In North America, humid climates occur across the eastern and subarctic regions of the continent and decline toward the south and west to the dry climates of the interior west and southwestern regions. Distribution of the meadow jumping mouse (Zapus hudsonius) largely is coincident with the North American distribution of Koppen climatic type D (i.e., humid continental climates; Hall, 1981; Fig. 1). Here the species occupies a wide range of habitats and sometimes has been considered a habitat generalist, although it typically is associated with dense herbaceous vegetation (e.g., Quimby, 1951; Whitaker, 1972; Choate et al., 1991). However, at the southwestern edge of its range where the climate is dry (i.e., Koppen climatic type Bsk, semiarid continental climate), the species is associated with riparian zones (Morrison, 1990, 1992; Trainor et al., 2007x; P. Cryan, in litt.).

Increasing habitat specialization at the southwestern limit of the range can put populations at risk because wetland habitats in arid regions typically are small, isolated, and prone to modification through land-use practices by humans. For example, Preble's meadow jumping mouse (Z. h. preblei), which is associated with riparian habitats along foothills of the northeastern edge of the southern Rocky Mountains (Fig. 1), is listed as threatened under the United States Endangered Species Act due to declines associated with modification of riparian habitats (United States Fish and Wildlife Service, 1998). In comparison, the New Mexico meadow jumping mouse (Z. h. luteus) is a morphologically and genetically distinctive subspecies that occupies a more isolated and extreme environment in the arid American Southwest (Miller, 1911; Hafner et al., 1981; King et al., 2006). Distribution of Z. h. luteus primarily is associated with the Southern Intermontane Plateaus physiographic region, with records from several major mountain ranges, and the Rio Grande and adjacent Chama River valleys (Miller, 1911; Hafner et al., 1981; Hoffmeister, 1986; Morrison, 1990, 1992; Zwank et al., 1997; Jones, 1999; Fig. 1). This disjunct distribution is considered relictual from a more widespread distribution during past glacial maxima (Hafner, 1993). Currently, Z. h. luteus is listed as a candidate for protection under the United States Endangered Species Act, endangered in New Mexico, threatened in Arizona, sensitive by the United States Forest Service and United States Bureau of Land Management, and it has a Natural Heritage conservation status of imperiled in Arizona and critically imperiled in Colorado and New Mexico. Identified threats include negative impacts to wetland habitats from development, grazing, water diversions, and conversion of riparian habitat to agricultural crops (Hafner and Yensen, 1998; New Mexico Department of Game and Fish, 2004).


During the mid-1980s to early 1990s, Joan Morrison conducted the first intensive surveys for Z. h. luteus, which were detailed in Morrison (1992) and reports to agencies, and were documented by specimens deposited in the Museum of Southwestern Biology, University of New Mexico. Surveys in New Mexico during the 1980s revealed that Z. h. luteus persisted at all historical localities sampled (although not all historical localities were resurveyed) and was discovered at new localities, which eased conservation concerns (New Mexico Department of Game and Fish, 1998). However, subsequent surveys in the White Mountains of Arizona during 1991 failed to document Z. h. luteus at 19 of 24 (79%) locations surveyed, which included 2 of 4 historical localities and many areas with seemingly suitable habitat U. L. Morrison, in litt.). Morrison concluded that the population in the White Mountains was declining and endangered as a result of habitat degradation due to livestock grazing and recreation.

The historical status of Z. h. luteus is best documented in the Jemez and Sacramento mountains, New Mexico, where it occupied a broad distribution (Morrison, 1992). However, the most recent captures of Z. h. luteus in the Jemez and Sacramento mountains were in 1989 and 1994, respectively, despite regular mammalogy fieldwork in both ranges. New Mexico Department of Game and Fish (2004) considered threats to habitat of Z. h. luteus likely most severe in montane areas and anecdotal observations in both ranges suggested declines in distribution and quality of riparian habitat. Consequently, the purpose of this study was to assess current status and habitat of Z. h. luteus in the Jemez and Sacramento mountains, New Mexico.

MATERIALS AND METHODS--Survey--We compiled historical locality records of Z. h. luteus in the Jemez and Sacramento mountains from museums, published literature, unpublished reports, and interviews with knowledgeable biologists. Field surveys at historical and potential new localities occurred 27 June-12 August 2005 in the Jemez and Sacramento mountains, and 14-19 August 2006 in the Jemez Mountains. Land ownership prevented surveying some historical localities. Survey locations were examined for potentially suitable habitat of Z. h. luteus, which we defined as presence of flowing surface water with riparian vegetation that provided adequate ground cover to conceal a traveling mouse (e.g., vertical cover >30 cm; Morrison, 1990).

We attempted to capture Z. h. luteus if potentially suitable habitat was present. Sherman (H. B. Sherman Trap, Tallahassee, Florida) live-traps were used because they allowed animals to be released unharmed and they were effective in previous inventories of Z. h. luteus (e.g., J. L. Morrison, in lift.; J. G. Koloszar and M. F. Ingraldi, in litt.). Traps were baited with commercial horse sweet feed (i.e., three or four types of grain mixed with molasses) and typically spaced ca. 3 m apart in transects of 40-80 traps situated within 1 m of water in the best-developed riparian vegetation. Sample sites within <1 km were considered a single location. Trapping effort at each locality was generally 200-600 trap-nights (i.e., 1 trap-night = one trap set for one night) over two nights, except where small available patches of riparian habitat were saturated with traps. Trapping ceased at a locality when Z. h. luteus was captured. Zapus h. luteus was considered functionally absent at a locality if either potentially suitable riparian habitat was absent or the species was not captured. We collected tissue samples from all captured Z. h. luteus, and retained at least one Z. h. luteus from each locality as a voucher specimen. In addition, to establish relationships between conditions of riparian habitat and communities of small mammals, we sampled some localities lacking potentially suitable habitat for Z. h. luteus generally using ca. 100 trap-nights. We calculated relative abundance as number of individuals captured/ 100 trap-nights. Trapping and euthanasia protocols followed American Society of Mammalogists guidelines (Animal Care and Use Committee, 1998).

Habitat--We collected habitat data at each trap where Z. h. luteuswas captured and at a random point within the best-developed riparian habitat at sites where Z. h. luteus was not captured. At each point where habitat data were collected, we measured distance to surface water, estimated slope with a compass, and measured canopy cover with a densitometer in the four cardinal directions. We obtained an index of soil moisture ranging from 1 (dry) to 10 (saturated) using a soil-moisture probe inserted into the ground ca. 40 mm. We measured vertical cover with a Robel pole (Robel et al., 1970) from 4 m distance at 1 m eye level with measurements taken at three random azimuths away from and three random azimuths towards the habitat point. Vertical cover was reported for the three measurements taken at the habitat point, the three measurements taken 4 m away from the habitat point, and averaged across all six measurements.

We established four 4-m perpendicular transects at a random azimuth from the habitat point. At each 1-m interval along a transect, we used a Daubenmire (1959) frame to assess the percent of ground covered by open water, sedges and rushes, forbs, grasses, litter, rocks, gravel, bare ground, and riparian shrubs (i.e., alder Ulnus, willow Salix). Cover classes were 1 for 0-5% cover, 2 for 5-25% cover, 3 for 25-50% cover, 4 for 50-75% cover, 5 for 75-95% cover, and 6 for 95-100% cover. Within each frame, we measured soil moisture, depth of litter, and two measures of height of stubble. We measured laid-over height of stubble as height of vegetation as it naturally lay and vertical height of stubble as height of a representative blade of graminoid vegetation that was fully extended vertically from the ground. Finally, we recorded identity and number of each tree and shrub within 1 m of each transect. At some habitat points where Z. h. luteus was not captured, we measured habitat along two transects because preliminary analyses indicated no significant difference (P > 0.05) in measurements whether based on two or four transects.

We calculated statistics using SPSS 10.0 for Windows (SPSS, Inc., 1999). We assessed all variables for normality with a one-sample Kolmogorov-Smimov test (SPSS, Inc., 1999). To test for differences in habitat variables at points where Z. h. luteus was captured or not captured, we used two-tailed tests and Mann-Whitney Utests for parametric and non-parametric data, respectively. We used Chis-quare tests to test for relationships between presence of a livestock exclosure with presence of potentially suitable riparian habitat and capture of Z. h. luteus.

RESULTS--Historical Localities--We identified 13 and 18 historical capture localities for Z. h. luteus in the Jemez and Sacramento mountains, respectively (Appendix 1). These included several records overlooked during previous studies. In the Jemez Mountains, this included one from the Redondo Creek drainage on the Valles Caldera National Preserve (locality J-1), while in the Sacramento Mountains this included two localities on the Rio Tularosa (localities S-1, S-2), one from Wills Canyon (locality S-12), and one from lower Aqua Chiquita Creek (locality S-19). The series of specimens from the Rio Tularosa represent the first records in the Sacramento Mountains from outside the Rio Penasco watershed.


Field Surveys--Our survey effort involved 22 localities using 3,513 trap-nights in the Jemez Mountains and 18 localities using 2,375 trap-nights in the Sacramento Mountains, which exceeded effort during prior survey efforts in these ranges (Fig. 2; J. L. Morrison, in litt.). We captured 14 Z. h. luteus, which had an overall relative abundance of 0.24 captures/100 trap-nights. At localities where it occurred, relative abundance had a mean of 0.84 and range was 0.13-1.45. Lowest relative abundances were at three small, isolated localities, including Silver Springs Creek (0.13; locality S-4) and Agua Chiquita Creek (0.28; locality S-16) in the Sacramento Mountains, and San Antonio Creek (0.48; locality J-3) in the Jemez Mountains. In contrast, those from four localities on the Rio Cebolla in the Jemez Mountains were >1.00.

In the Jemez Mountains, we captured Z. h. luteus at three historical and two new localities (Table 1, Fig. 2, Appendix 1). Four localities (localities J-6, J-8, J-9, and J-10) were along a ca. 12-km reach of the Rio Cebolla, although riparian habitat within this segment was fragmented. Three of the four localities were entirely within livestock exclosures, while the fourth (locality J-9) included captures both within and outside a livestock exclosure. No Z. h. luteus and only poorly developed riparian habitat was found above or below this reach, although historical records attested to its former occurrence in those areas. The remaining locality was an isolated area of suitable habitat in a public campground along a 0.8-km reach of lower San Antonio Creek. Via drainages, this locality was isolated from those along the Rio Cebolla by ca. 48 km of mostly unsuitable habitat. No Z. h. luteus and only a few small, isolated patches of potentially suitable habitat were found within the Rio de las Vacas watershed, which accounted for 27% of historical localities in the Jemez Mountains.

In the Sacramento Mountains, we captured Z. h. luteus at one historical and one new locality (Table 1, Fig. 2, Appendix 1). The historical locality (locality S-4) was on a 1.8-km reach of Silver Springs Creek within a fenced livestock exclosure. A second 1.6-km reach with potentially suitable habitat was ca. 1.3 km upstream, but was not surveyed due to land ownership. On Agua Chiquita Creek, we captured Z. h. luteus in the second of three livestock exclosures located below a major headwater spring. Total length of the reach occupied by the three exclosures was ca. 2.6 km. Two livestock watering access points that were devoid of riparian vegetation fragmented this reach. The largest gap (between the second and third exclosure) was ca. 0.2 km, which might serve as a significant barrier to the upper two exclosures, which together were ca. 1.8 km. Although most drainages in the Rio Penasco watershed were surveyed visually, the only other area that appeared suitable for Z. h. luteus was a livestock exclosure on the upper Rio Penasco (locality S-8), but no Z. h. luteus was captured in 600 trap-nights.

Habitat--Sites where Z. h. luteus was captured had significantly greater vertical cover and vertical height of stubble (Table 2). There was a highly significant relationship between presence of a livestock exclosure with presence of potentially suitable riparian habitat ([chi square] = 12.108, df = 1, P = 0.001) and capture of Z. h. luteus ([chi square] = 15.010, df = 1, P = 0.001).

DISCUSSION--Newly discovered historical records for Z. h. luteus indicate that it had a broader distribution in the Jemez and Sacramento mountains than previously recognized. However, field surveys at these localities revealed a dramatic decline in distribution of Z. h. luteus and its habitat since the late 1980s. Zapus h. luteus or its habitat was not found at 73% of historical localities surveyed in the Jemez Mountains and at 94% of historical localities surveyed in the Sacramento Mountains. In both ranges, current distribution of Z. h. luteus was restricted to small areas of well-developed riparian habitat along isolated stream reaches. We consider the population in the Sacramento Mountains to be at serious risk of extinction because it is known to persist only within grazing exclosures along <5 km of streams. Given similar declines documented for Z. h. luteus in the White Mountains and for Z. h. preblei in the southern Rocky Mountains, these results suggest that declines might be expected for other southwestern peripheral populations of Z. hudsonius (United States Fish and Wildlife Service, 1998; J. L. Morrison, in litt.). Further, because the primary reason for decline of this species was loss of herbaceous riparian habitat, other species associated with these habitats (e.g., montane vole Microtus montanus, ermine Mustela erminea) also might have experienced similar declines in distribution and abundance in the American Southwest.

Sites where Zapus h. luteus was captured were in ungrazed, herbaceous, emergent, riparian habitats dominated by tall, dense stands of sedges on saturated soils. However, we did not find Z. h. luteus in sedge-dominated habitats where standing water was deep (i.e., [greater than or equal to] 2 cm). In contrast, Morrison (1990, 1992) described Z. h. luteus as occupying riparian habitat dominated by grasses and forbs. However, this difference in habitat description likely was due to different methods of data collection. Data Morrison collected were from 10 plots along the stream where traps were set and at 10 locations ca. 15 m away from the stream; hence her data describe general features of the survey area (J. L. Morrison, in litt.). In contrast, our data were collected at 16 plots within a 4m radius of locations of captures and, hence, describe specific habitat used by Z. h. luteus. Our assessment of habitat should be considered conservative because it remains a possibility that Z. h. luteus was present but not captured at some survey sites, particularly in instances where multiple sites existed within an area excluded from grazing. Consequently, Z. h. luteus appears to be more highly specialized on wetland associations as compared with other peripheral subspecies of Z. hudsonius, including Z. h. preblei, Z. h. campestris, and Z. h. pallidus, which use a broader array of riparian and mesic grassland habitats (e.g., Choate et al., 1991; United States Fish and Wildlife Service, 1998, 2002; Trainor et al., 2007x; P. Cryan, in litt.). This specialization might reflect the more arid climate occupied by Z. h. luteus in the American Southwest.

Reason for decline of Z. h. luteus was loss of tall, dense, herbaceous riparian vegetation. Our data indicated that the primary proximate cause of this habitat loss was livestock grazing. Other studies also have demonstrated a negative response of jumping mice to grazed habitats (e.g., Hanley and Page, 1982; Giuliano and Homyack, 2004; Morrison, 1990; Schulz and Leininger, 1991). Presence of a livestock exclosure was a highly significant factor related to presence of potentially suitable riparian vegetation and presence of Z. h. luteus.

Morrison (1990, in litt.) noted that Z. h. luteus occasionally was captured where livestock occurred, but those findings cannot be interpreted given the paucity of data concerning the specific circumstances. Our only captures of Z. h. luteus outside of fenced livestock exclosures was on the lower Rio Cebolla in the Jemez Mountains (locality J-9). This locality was unique because Z. h. luteus was captured in both a livestock exclosure and the adjacent grazed area. Zapus hudsonius is a relatively vagile and long-lived rodent (Quimby, 1951). Thus, it is possible that captures in the grazed area represented transient individuals or individuals persisting from pre-grazing periods. However, a more likely explanation was that the location was in a broad valley that had an extensive complex of dams of the American beaver (Castor canadensis), which created a diverse network of emergent wetland habitat. Although cattle were present in adjacent uplands and grazing was observed along the periphery of the wetland, we did not observe evidence of cattle within the wetland associated with dams of American beavers. Further, habitat at capture sites in the wetland was similar to localities within livestock exclosures. Thus, wetland associated with dams of American beavers likely served to naturally inhibit cattle; perhaps, due to reticence of cattle to walk in saturated mud and presence of ample upland forage. Zapus h. luteus was not captured at other areas along the lower Rio Cebolla where wetlands associated with American beavers or livestock exclosures were absent. Because American beavers can create extensive areas of suitable habitat for Z. h. luteus, the historical decline and current absence of American beavers from many areas likely had a negative impact on Z. hudsonius (Huey, 1956; pers. observ.).

Livestock grazing can exert many influences on riparian habitats (Elmore and Kauffman, 1994; Ohmart, 1996; Cartron et al., 2000). For example, we determined that habitat within livestock exclosures had significantly higher soil moisture (z = -3.001, P = 0.003), vertical cover (t = -7.703, P < 0.001), vertical height of stubble (t = -7.028, P < 0.001), sedge-rush ground cover (t = -4.404, P < 0.001), and depth of litter (z = -3.810, P < 0.001), but significantly less bare ground cover (z = -3.716, P < 0.001) and distance to water (z = -2.503, P = 0.012). Livestock exclosures can promote stream flow, which is required by Z. h. luteus (Morrison, 1990), by reducing soil compaction and evaporation. Second, livestock exclosures allow development of tall, dense, herbaceous cover by reducing grazing and trampling (Wyman et al., 2006). This and other studies have concluded that adequate herbaceous cover is required to maintain populations of Z. hudsonius (Whitaker, 1963).

Climatic variation likely exerts an important influence on distribution and abundance of Z. h. luteus. Wet periods result in increased herbaceous growth in uplands and wetlands due to increased availability of surface water (Trainor et al., 2007b). Relative to Z. h. luteus, dense herbaceous vegetation might function to decrease risk of predation and competition with aggressively dominant voles (Microtus), and increase food resources, exploratory behavior, and quality of adjacent upland habitat used for hibernation (Boonstra and Hoyle, 1986; Meaney et al., 2003; Trainor et al., 2007x, 2007b). Movement, dispersal, and gene flow in riparian jumping mice are largely determined by habitat connectivity with most movements along riparian corridors (Vignieri, 2005J. L. Morrison, in litt.). Thus, wet periods might provide longer, more continuous stretches of suitable riparian habitat allowing Z. h. luteus to expand its distribution, including marginal localities that might be suitable only during wet periods. In contrast, drought might shrink and isolate suitable habitat, possibly resulting in local extirpations (Bessinger, 2000). This might explain the apparent absence of Z. h. luteus from the grazing exclosure on the upper Rio Penasco in the Sacramento Mountains (locality S-8). Finally, Z. h. luteus might be better able to co-exist with livestock during wet periods due to increased availability of wetland habitat and upland forage. However, livestock grazing should be more carefully controlled during droughts. Given projected climatic warming (Intergovernmental Panel on Climate Change, 2001), we expect that drought will become an increasing problem for Z. h. luteus.

Based on Palmer drought data, previous surveys for Z. h. luteus in the Jemez Mountains during 1985-1986 and 1989, and in the Sacramento Mountains during 1988 occurred during or immediately following periods of above-average moisture (National Climatic Data Center, drought/xmgr.html). In contrast, surveys in the White Mountains, during 1991, occurred during a period of near-normal conditions following 1.5 years of severe-to-extreme drought. Our study was conducted during unusually moist conditions, but most of the preceding 5 years were during severe-to-extreme drought. Thus, the restricted distribution and rarity of Z. h. luteus during this and the previous survey in the White Mountains might have been influenced by drought.

We observed localities where habitat of Z. h. luteus was lost due to development, recreation, and forest fires. In the Sacramento Mountains, we observed many springs that were capped and diverted for development of residential and city infrastructure and streams that were diverted for irrigation. In the Jemez Mountains, we observed loss of riparian habitat due to off-road vehicles, camping, and human-social trails, which result in soil compaction, erosion, and destruction of vegetation. Finally, we observed areas (e.g., locality S-15) where riparian habitat had been eliminated due to erosion or aggradation resulting from flooding following forest fire.

Recommendations--Conservation and management of montane populations of Z. h. luteus must focus on maintaining and enhancing the distribution of undisturbed, herbaceous riparian habitat. Although appropriate grazing management can maintain the ecological integrity of some riparian ecosystems (Baker et al., 2001), presence of cattle in riparian zones during the growing season might negatively impact habitat of Z. h. luteus through reductions in vegetative cover due to grazing and trampling. Maintenance of undisturbed riparian habitat might only be possible through creation of refugial areas by complete exclusion of livestock from the riparian zone, particularly in areas where there is passive-continuous grazing during the growing season (Elmore and Kauffman, 1994; Ohmart, 1996; Wyman et al., 2006). Exclusion fencing can be the most practical means for achieving rapid recovery of riparian habitat, and loss of forage on degraded streams may be negligible (Wyman et al., 2006). Giuliano and Homyack (2004) demonstrated that small mammals, including Z. hudsonius, respond quickly to establishment of livestock exclosures. We recommend establishment of refugial areas that are large enough to sustain a local population throughout the longest anticipated drought cycle and spatially arranged to maintain viable metapopulations throughout the historical range. Zapus hudsonius is unwilling to cross even small areas with little or no cover (Quimby, 1951). Because unmanaged grazing by livestock results in excessive use of riparian areas with concomitant degradation, grazing management should include measures such as growing-season rest to enhance riparian vegetation outside refugia to promote dispersal among refugia (Ohmart, 1996). Given the recent and rapid decline in distribution of Z. h. luteus, it is important to conduct additional inventory work to identify any additional populations that have persisted, to initiate long-term monitoring of populations and riparian habitat, and to restore riparian habitat and repatriate key populations of Z. h. luteus. Finally, additional studies are needed on relationships among livestock grazing, habitat, and Z. h. luteus, including potentially important factors such as activity by American beavers, conditions of upland habitat, and climate.

Editor was Michael L. Kennedy.
APPENDIX 1--Localities where Zapus hudsonius luteus was captured
in the Jemez Mountains, Sandoval County, and the Sacramento
Mountains, Otero County, New Mexico. For museum records, locality
data are from specimen tags and museum records; brackets include
other locality information. Specimen catalog numbers are from
University of Kansas Museum of Natural History (KU), the Museum
of Southwestern Biology, University of New Mexico (MSB), New
Mexico Department of Health Zoonoses Program (NMDH), United
States National Museum (USNM), and Academy of Natural Sciences of
Philadelphia (ANSP).

 Results of
Locality current
 survey Drainage

Jemez Mountains

 J-1 Not surveyed San Antonio
 J-2 No suitable San Antonio
 riparian Creek
 J-3 Present at San Antonio
 new Creek
 J-4 No suitable Guadalupe
 riparian River
 J-5 Species Rio Cebolla
 J-6 Present at Rio Cebolla
 J-7 Not surveyed Rio Cebolla
 J-8 Present at Rio Cebolla
 J-9 Present at Rio Cebolla
 J-10 Present at Rio Cebolla
 J-11 Species Rio Cebolla
 J-12 Species Rio de las
 absent Vacas
 J-13 No suitable Rio de las
 riparian Vacas
 J-14 No suitable Rio de las
 riparian Vacas
 J-15 No suitable Rio de las
 riparian Vacas
 -- habitat Unknown

Sacramento Mountains

 S-1 Not surveyed Rio Tularosa
 S-2 Not surveyed Rio Tularosa
 S-3 No suitable Silver Springs
 S-4 Present at Silver Springs
 S-5 No suitable Silver Springs
 S-6 No suitable James Canyon
 S-7 No suitable James Canyon
 S-8 Species Upper Rio
 absent Penasco
 S-9 No suitable Upper Rio
 riparian Penasco
 S-10 No suitable Upper Rio
 riparian Penasco
 S-11 No suitable Upper Rio
 riparian Penasco
 S-12 No suitable Upper Rio
 riparian Penasco
 S-13 Species Agua Chiquita
 S-14 No suitable Agua Chiquita
 S-15 No suitable Agua Chiquita
 S-16 Present at Agua Chiquita
 S-17 No suitable Agua Chiquita
 S-18 No suitable Agua Chiquita
 S-19 No suitable Agua Chiquita

Locality Locality

Jemez Mountains

 J-1 Beaver pond in geothermal well area on a
 gentle northwest slope of Redondo Peak,
 elevation ca. 8,500 feet
 J-2 T20N, R3E, south central Sec 20 San
 Antonio Creek
 J-3 Jemez Mountains, San Antonio Creek,
 south end San Antonio Campground, 1.2
 mile N, 0.5 mile W junction NM Hwy 4 and
 NM Hwy 126; T19N, R3E, NEl/4 of NW 1/4
 of NW 1/4 Sec 17
 J-4 Virgin Canyon, T18N, R2E [lower Virgin
 Canyon, below old cabins near spotted
 owl site; J. L. Morrison, in litt.]
 J-5 Rio Cebolla, T20N, R2E, Sec 24 near Hay
 J-6 T20N, R2E, NW 1/4 Sec 35 Seven Springs
 Fish Hatchery
 Jemez Mountains, Seven Springs State
 Fish Hatchery; T20N, R2E, SE 1/4 of NE
 39817 of NE 1/4 Sec 34
 J-7 1 mile S Seven Springs Hatchery
 J-8 Fenton Lake, Jemez Mountains
 T19N, R2E, SW 1/4 Sec. 10 Fenton Lake,
 marsh E of lake, W of Route 126
 Jemez Mountains, Fenton Lake State Park,
 marsh at upper end of lake along Rio
 Cebolla above NM Hwy 126; T19N, R2E,
 Swl/4 of NW 1/4 of SW 1/4 of SW 1/4 Sec
 Fenton Lake-creek that runs into lake
 from south, T19N, R2E, NW 1/4 Sec 15
 Jemez Mountains, Fenton Lake State Park,
 Lake Fork Day Use Area, mouth of small
 tributary that flows W along NM Hwy 126
 and entering S side Fenton Lake; T19N,
 R2E, SE 1/4 of NE 1/4 of NE 1/4 Sec 16
 J-9 Rio Cebolla at intersection of Route 376
 & Lake Fork Creek, T19N, R2E, NE 1/4 Sec
 Jemez Mountains, Rio Cebolla at junction
 with Lake Fork Canyon, above Forest
 Service Road 376 bridge; T19N, R2E, SW
 39817 of SW 1/4 of SW 1/4 Sec 20 and SE
 Rio Cebolla, 0.6 mile (by Forest Road
 376) southwest of Forest Road 376 bridge
 over Rio Cebolla, which is located at
 the junction of Lake Fork Canyon, 9.5 km
 N, 6.5 km W Jemez Springs, T19N, R2E, W
 39815 of NE 1/4 Sec. 30 30 39817 of SE
 J-10 Jemez Mountains, Rio Cebolla, 1.7 N, 0.4
 mile W jct Rio Cebolla and Rio de las
 Vacas; T19N, R1E, SE 1/4 of SE 1/4 Sec
 25 1/4 of SE 1/4 Sec 19
 J-11 Rio Cebolla, T19N, R1E, 1 mile up from
 Rio de las Vacas
 J-12 T20N, R2E NE 1/4 Sec 3 Rito Penas Negras
 8,360 feet
 J-13 17 km SE Cuba, T20N, R1E, S 12,
 elevation 2,600 m
 J-14 Rito Penas Negras, T20N, R1E, Sec 13,
 int. Rio de las Vacas [Rito Penas Negras
 at the junction of Forest Road 126; J.
 L. Morrison, in lift.]
 J-15 T20N, R1E west central Sec 25 Rio
 de las Vacas x Turkey Creek
 -- Unknown

Sacramento Mountains

 S-1 Bank of Tularosa Creek,
 1 mile above Mescalero
 S-2 Bank of Tularosa Creek,
 2 mile above Mescalero
 S-3 Silver Springs Canyon, intersection
 Forest Road 24 & 1 [Silver Springs,
 T15S, R13E, Sec 29, 8,400 feet; J. L.
 Morrison, in litt.]
 S-4 Silver Creek, 8 miles NE Cloudcroft
 8 miles E Cloudcroft
 Silver Springs Canyon, boundary
 Mescalero Apache Reservation
 T15S R13E Sec 22
 Sacramento Mountains, Silver Springs
 Creek at jct Turkey Pen Canyon and
 Forest Service Road 405 (=County Road
 C7), 2.9 miles N, 4.6 miles E
 Cloudcroft; T15S, R13E, SE 1/4 of NW
 39817 Sec 22 Sec 22
 S-5 10 miles NE Cloudcroft [10 miles NE
 Cloudcroft, 8,500 feet;
 Bailey, 1931]
 S-6 Int. Pumphouse Canyon, Route 82
 [Pumphouse Canyon, T16S, R12E, Sec 3,
 8,300 feet; J. L. Morrison, in litt.]
 3.2 miles (by road) E Cloudcroft
 S-7 12 miles E Cloudcroft, 7,500 feet
 [Penasco Creek, 12 miles E Cloudcroft,
 7,500 feet; Bailey, 1931]
 S-8 Rio Penasco, Int. Route 164 & Route 64
 [Rio Penasco, T17S, RUE, Sec 11, 8,600
 feet; J. L. Morrison, in litt.]
 S-9 Water Canyon [Water Canyon, T17S, R1IF,
 Sec 24, 8,600 feet; J. L. Morrison, in
 S-10 Rio Penasco [Rio Penasco, T17S, R12E,
 Sec 10, 8,000 feet; J. L. Morrison, in
 S-11 Rio Penasco, T17S, R13E, Int. with Route
 541 [Rio Penasco at Cox, T17S, R13E,
 Sec 3, 7,200 feet; J. L. Morrison, in
 S-12 Wills Canyon, UTM E4331, N36311
 S-13 Hay Canyon, int. 257, 541, T17S, R12E,
 Sec. 19 [Masterson Springs, T17S, R12E,
 Sec 19, 8,000 feet; J. L. Morrison, in
 S-14 Spring Canyon [Spring Canyon, T17S,
 R12E, Sec 36, 8,400 feet; J. L.
 Morrison, in litt.]
 S-15 Potato Canyon T18S R13E Sec 5 [Potato
 Canyon, T18S, R13E, Sec 5, 8,200 feet;
 J. K. Morrison, in lift.] [Potato Creek;
 Morrison, 1992]
 S-16 Sacramento Mountains, Agua Chiquita
 Creek, 5.75 miles S, 6.5 miles W
 Sacramento; T18S, R12E, SE 1/4 of
 NE 1/4 of SW 1/4 Sec 25
 S-17 Agua Chiquita [upper Aqua Chiquita,
 T18S, R13E, Sec 19, 8,000 feet; J. L.
 Morrison, in lift.]
 S-18 Agua Chiquita [lower Agua Chiquita,
 T18S, R13E, Sec 17, 8,000 feet; J. L.
 Morrison, in lift.]
 S-19 Weed, east of Cloudcroft,
 Sacramento Mountains

 Results of
Locality current
 survey Reference Date

Jemez Mountains

 J-1 Not surveyed W. Whitford, pers. Mid-1970s
 J-2 No suitable MSB 56991-56992 5 September 1985
 J-3 Present at Current study 28 June 2005
 J-4 No suitable MSB 62096 2 August 1989
 J-5 Species MSB 62101 4 August 1989
 J-6 Present at MSB 56993-56994 23, 27 August 1985
 locality Current study 1 July 2005
 J-7 Not surveyed NMDH 17 September 1969
 J-8 Present at MSB 41055 5 August 1979
 historical MSB 56979-56983 23, 27-28 August
 locality 7 June 1905
 Current study 29 June 2005
 MSB 56984 27 August 1985
 Current study 29 June 2005

 J-9 Present at MSB 56985 30 August 1985
 historical Current study 5 July 2005
 locality Current study 15 August 2006
 J-10 Present at Current study 4 July 2005
 J-11 Species MSB 62097-62098 24 August 1989
 J-12 Species MSB 56987-56990 5-6 September 1985
 J-13 No suitable MSB 67525 12 July 1985
 J-14 No suitable MSB 62102 3 August 1989
 J-15 No suitable MSB 56986 6 September 1985
 -- habitat MSB 56995-56997 unknown

Sacramento Mountains

 S-1 Not surveyed ANSP 15771-15772 18 June 1932
 S-2 Not surveyed ANSP 15573-15579 27 June, 1 July, 2
 1 August 1932
 S-3 No suitable MSB 61700-61702 22 July 1988
 S-4 Present at MSB 36142 21 July 1977
 historical MSB 37154-37155 21 July 1977
 locality MSB 61703-61704 22 July 1977
 Current study 22 July 2005
 S-5 No suitable USNM 118798 10 September 1902
 S-6 No suitable MSB 61684 15 July 1988
 riparian MSB 37323-37326, 3 September 1978,
 habitat 41058-41066 17-18 August 1979
 S-7 No suitable USNM 119032- 7 September 1902
 riparian 119033
 S-8 Species MSB 61678-61680, 13, 16 July 1988
 absent 61687
 S-9 No suitable MSB 61690, 62095 17 July 1988, 18 Jul
 riparian 11 June 1905
 S-10 No suitable MSB 61686 16 July 1988
 S-11 No suitable MSB 61696 18 July 1988
 S-12 No suitable P. Ward, pers. 11 July 1992,
 riparian comm. 11 July 1993,
 habitat 13-14 July 1994
 S-13 Species MSB 61712 31 July 1988
 S-14 No suitable MSB 61693 19 July 1988
 S-15 No suitable MSB 61688-61689 17 July 1988
 S-16 Present at Current study 19 July 2005
 S-17 No suitable MSB 61692 18 July 1988
 S-18 No suitable MSB 61691 18 July 1988
 S-19 No suitable ANSP 14779 1 July 1931

 Results of
Locality current
 survey Notes

Jemez Mountains

 J-1 Not surveyed
 J-2 No suitable
 J-3 Present at 1 captured
 J-4 No suitable
 J-5 Species
 J-6 Present at
 locality 2 captured
 J-7 Not surveyed
 J-8 Present at
 locality 2 captured
 1 captured
 J-9 Present at
 locality 2 captured
 3 captured
 J-10 Present at 1 captured
 J-11 Species
 J-12 Species
 J-13 No suitable
 J-14 No suitable
 J-15 No suitable
 -- habitat Collected by J.
 Morrison; not a
 new locality.
Sacramento Mountains

 S-1 Not surveyed
 S-2 Not surveyed
 S-3 No suitable
 S-4 Present at
 locality 1 captured
 S-5 No suitable Locality is in Silver
 riparian Springs
 habitat Canyon near
 junction with
 Poison Spring
 and Indian Joe
 canyons (J. K.
 Frey et al., in
 S-6 No suitable
 S-7 No suitable Locality is in
 riparian James Canyon
 habitat near junction
 of Eightmile
 Canyon (J. K
 Frey et al., in
 S-8 Species
 S-9 No suitable
 S-10 No suitable
 S-11 No suitable
 S-12 No suitable 4 captures
 S-13 Species Locality is likely
 absent Prestridge
 Spring, T17S,
 R13E, Sec. 20
 S-14 No suitable
 S-15 No suitable
 S-16 Present at 1 captured
 S-17 No suitable
 S-18 No suitable
 S-19 No suitable

We thank S. Cary, T. C. Frey, the T. Frey family, J. Malaney, C. Pety, and personnel from Fenton Lake State Park, Lincoln National Forest, Santa Fe National Forest, and Seven Springs State Fish Hatchery for help with various aspects of this study. For providing information or loans of specimens, we thank R. Fisher and C. Ludwig from the United States National Museum, N. Gilmore from the Academy of Natural Sciences of Philadelphia, R. Baker and H. Garner from Museum of Texas Tech University. We thank Z. Schwenke for assistance in the field during 2006, W. Whitford and P. Ward for providing information on their historical captures, A. Hope, D. Hafner, and two anonymous reviewers for helpful comments on a previous version of this paper, and M. Ryan for the Spanish translation of the abstract. New Mexico Department of Game and Fish, Jemez Ranger District of the Santa Fe National Forest, and New Mexico State Parks provided partial funding for this study.

Submitted 29 January 2008. Accepted 16 June 2008.


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Frey Biological Research, P.O. Box 294, Radium Springs, NM 88054 (JKF) Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131 (JLM)

* Correspondent:
TABLE 1--Results of surveys for Zapus hudsonius luteus in
the Jemez and Sacramento mountains, New Mexico,
during 2005 and 2006.

 Historical localities
 Percent of
Mountain Not where
range surveyed Present Absent absent

Jemez 2 3 8 73
Sacramento 2 1 15 94

Mountain New
range localities Total

Jemez 2 15
Sacramento 1 19

TABLE 2--Habitat characteristics at sites where Zapus
hudsonius luteus was captured (n = 14) or not captured
(n = 30) in the Jemez and Sacramento mountains in 2005
and 2006.

 Not captured Captured

 x SD x SD

Slope ([degrees]) 1.8 2.78 1.1 2.89
Soil moisture (1-10) 8.1 2.95 9.0 1.16
Distance to water (m) 62.3 295.40 1.6 2.78
Canopy cover (%) 9.8 18.59 11.5 18.50
Vertical cover (cm)
 4 m from trap or
 random location 49.1 31.24 82.1 35.14
 At trap or
 random location 53.3 33.67 83.6 34.90
 Mean 51.2 30.62 82.9 33.95
Height of stubble (cm)
 Vertical 62.6 39.16 89.8 30.83
 Laid-over 52.2 34.61 70.5 28.61
Depth of litter (cm) 5.7 10.82 7.6 9.09
Ground-cover class (1-6)
 Sedge/rush 2.9 1.64 3.1 1.33
 Forb 1.8 0.87 2.0 0.97
 Grass 2.0 0.90 1.9 1.39
 Alder/willow 1.0 0.17 1.1 0.29
 Litter 1.3 0.41 1.1 0.18
 Rock 1.0 0.08 1.0 0.00
 Gravel 1.0 0.07 1.0 0.00
 Bare ground 1.3 0.62 1.1 0.12
 Open water 1.2 0.40 1.4 0.36
Number of trees-
 shrubs/transect 3.4 17.59 0.1 0.23

 statistic P

Slope ([degrees]) z = -1.167 0.243
Soil moisture (1-10) z = -0.280 0.779
Distance to water (m) z = -0.771 0.440
Canopy cover (%) z = -0.279 0.780
Vertical cover (cm)
 4 m from trap or
 random location t = -3.145 0.003
 At trap or
 random location t = -2.748 0.009
 Mean t = -3.089 0.004
Height of stubble (cm)
 Vertical t = -2.286 0.027
 Laid-over t = -1.717 0.093
Depth of litter (cm) z = -1.285 0.199
Ground-cover class (1-6)
 Sedge/rush t = 0.427 0.672
 Forb t = -0.645 0.522
 Grass z = -1.417 0.157
 Alder/willow z = -1.285 0.199
 Litter z = -0.358 0.720
 Rock z = -0.977 0.328
 Gravel z = -1.211 0.226
 Bare ground z = -0.119 0.905
 Open water z = -1.481 0.139
Number of trees-
 shrubs/transect z = -0.206 0.837
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Author:Frey, Jennifer K.; Malaney, Jason L.
Publication:Southwestern Naturalist
Article Type:Report
Geographic Code:1U8NM
Date:Mar 1, 2009
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