Distribution and Abundance of Fallow Deer Leks at Point Reyes National Seashore, California

By Gary M. Fellers and Michael Osbourn

Scientific Investigations Report 2006-5149, published 2006.

U.S. Department of the Interior

Dirk Kempthorne, Secretary

U.S. Geological Survey

P. Patrick Leahy, Acting Director
The use of firm, trade, and brand names in this report is for identification purposes only and does not constitute endorsement by the U.S. Geological Survey.

Abbreviations

cm
centimeter
ESRI
Environmental Systems Research Institute
GIS
Geographic Information System
ha
hectare
km
kilometer
m
meter
m2
square meter
PDA
Personal Digital Assistant
UTM
Universal Transverse Mercator. A metric coordinate system commonly used for mapping; UTM coordinates are marked on the sides of most USGS topographic maps.

Introduction

Only two species of ungulates (hoofed mammals) are native to Marin County, tule elk (Cervis elaphus nannodes) and Columbian black-tailed deer (Odocoileus hemionus columbianus). In the 1940s, European fallow deer (Dama dama) obtained from the San Francisco Zoo were released at Point Reyes. When Point Reyes National Seashore was established in 1962, fallow deer were well established within the boundaries of the National Seashore. The fallow deer population was estimated to be 500 in 1973 (Wehausen, 1973) and that number increased to 860 by 2005 (National Park Service, unpubl. data).
Fallow deer have an unusual mating system. During the fall mating season (or rut), male fallow deer establish areas known as leks where they display to potential mates (Hirth, 1997). This behavior is unique among deer and elk, but it is similar to breeding systems used by grouse and a few other birds and mammals. Formation of leks in ungulates decreases the number of aggressive encounters in which dominant males are involved when the local male density becomes too high, because the spatial stability of territories in leks reduces the number of aggressive encounters between males (Hovi et al., 1996; Pélabon et al., 1999).
A fallow deer lek is typically an area of about 100–150 m2 and typically includes two to five males. Using their hooves and antlers, each male clears away most or all of the vegetation and digs a rutting pit that he defends throughout the breeding season.
Stenström et al. (2000) described rutting pits in a Swedish population of fallow deer:
"Pits are large patches of bare soil found at the center of mating stands where most of the rutting activities take place. . . . Scrapes are small patches of bare soil found throughout the areas of deer activity. Only bucks showed any interest in scrapes. Within a 10 day period half the scrapes were rescraped at least once. Larger scrapes were more frequently rescraped than smaller ones. Frayings, i.e., removal of bark and subsequent scent marking on bushes and small trees close to scrapes, also had a positive effect on the frequency of rescraping. . . . fallow deer bucks in our study do not seem to mark territorial boundaries, rather the intensity of markings tends to decrease with distance from the rutting pit suggesting that scraping may instead be used in male status signaling."
Establishing and defending a rutting pit is energetically expensive. Apollonio et al. (1989) concluded that: "Body condition appears to be an important determinant of male copulatory success, because only males in superior condition could defend a lek territory for up to 2 weeks. Males do not feed while defending lek territories. Foraging ability during the year probably determines condition at the onset of the rut. Females appear to choose mates at least partially on the basis of location, preferring males located near traditional routes. Females may ultimately select mates in the best body condition."
In the fall of 2005, we initiated a study of the leks in two study sites at Point Reyes National Seashore. The goal of this work was to determine the distribution and size of fallow deer leks, and to evaluate the impact of both the leks and the associated rutting pits on the soil and vegetation.

Study Areas

Our study was conducted in two areas of Point Reyes National Seashore, the northern portion of the Olema Valley and an area around the Estero trailhead (fig. 1). The two study areas were selected to represent areas of high fallow deer density, and medium density.
Figure 1Distribution of high and medium/low density areas for fallow deer at Point Reyes National Seashore, California.
Triangle symbols (see figures 2 and 3) mark the location of leks within the Olema Valley and Estero trailhead study areas.
Distribution of high and medium/low density areas for fallow deer at Point Reyes National Seashore, California.
The Olema Valley was selected because it was typical of areas within the Seashore that have a high density of fallow deer, based on combined aerial and ground counts conducted in January 2001 (Gates, 2001). Initially, we intended to survey the entire high density area, but it quickly became obvious that the number of leks and the time required to document each lek site would preclude a complete survey. Hence we restricted our work to an area that was bounded by features that were readily discernable in the field (e.g., Olema Creek, Bear Valley Road, fence lines). The areas surveyed included Divide Meadow, Bear Valley, and portions of the pasture and riparian habitat along Olema Creek. The Olema Valley study area was 147.2 ha (363.7 acres) in size (fig. 2). The predominant vegetation was coast live oak (Quercus agrifolia) and California bay (Umbellularia californica) woodlands; red alder (Alnus rubra) and willow (Salix sp.) riparian zones; and grassy meadows and pastures.
Figure 2Fallow deer lek sites in the Olema Valley study area, Point Reyes National Seashore, California.
Fallow deer lek sites in the Olema Valley study area, Point Reyes National Seashore, California.
Once field work was completed in the Olema Valley, a similar size area was delineated near the Estero trailhead for surveys in the moderate density area. As with the Olema Valley study area, the Estero trailhead area was defined by using a combination of fence lines, roads, and natural features. The Estero trailhead area was selected as representative of an area with low-to-moderate densities of fallow deer (Gates, 2001). The Estero study area was 151.6 ha (374.6 acres) in size (fig. 3), roughly equal to the Olema Valley study area. The primary vegetation was coastal scrub dominated by coyote brush (Baccharis pilularis), riparian zones with red alder, and pasture grasslands.
Figure 3Fallow deer lek sites in the Estero trailhead study area, Point Reyes National Seashore, California.
Fallow deer lek sites in the Estero trailhead study area, Point Reyes National Seashore, California.

Methods

Leks were located by conducting visual surveys for fallow deer during, and shortly after, the fall lekking season (Oct. 3–Dec. 6, 2005). A total of 202 hours were spent conducting surveys. Fallow deer surveys were carried out at dawn from hilltop vantage points. Treelines and grasslands were scanned with binoculars to locate congregations of fallow deer, damaged vegetation, or bare ground. These areas were subsequently investigated to determine whether or not a lek was present. By repeatedly surveying the entire study area, we were confident that few, if any, leks were overlooked.
Leks were documented by noting the predominant vegetation, recording the condition of the soil and vegetation, measuring the length and width of the lek, and measuring each individual rutting pit. Leks and rutting pits were photographed and data were recorded with a PDA (Personal Digital Assistant) for later input into a Microsoft Access database.

Delineating Leks

Only areas with disturbed ground or rutting pits were recorded as leks. Pits ranged from shallow depressions where the vegetation had been scrapped away to trenches over 50 cm deep. Areas that showed only vegetation damage and lacked ground disturbance were not scored as leks. Areas with obvious cattle impacts or sites with no discernable deer signs (hoof prints, feces, urine stains, antler scrapes, or deer present) were not counted as leks, resulting in a conservative estimate of lek density.
Leks tended to be concentrated linearly along the treelines at margins of fields. Since many of these areas had a nearly continuous band of disturbance, it was difficult to define individual leks. We marked the end of a lek when there was at least 20 m of undamaged ground between lek areas. The length and width of the disturbed ground was measured with a digital range finder (Bushnell Yardage Pro). These measurements were used to calculate the area for each lek.

Measuring Rutting Pits

Within a lek, fallow deer bucks excavate numerous rutting pits. The length and width of each pit was measured with a fiberglass tape. Pit depth was measured with a ruler. These measurements were used to determine the total area of excavated ground and the depth of the pits.

Vegetation Condition

In addition to recording the predominant vegetation associated with each lek site, vegetation condition was evaluated. Disturbed ground, damaged foliage, damaged bark, and exposed roots were all noted. Disturbed ground occurred where the herbaceous cover had been scraped away, resulting in bare soil or excavated rutting pits. Damaged foliage included leaves, twigs, or branches that had been shredded or broken. Damaged bark was noted where fallow deer had used their antlers to break, tear, or scrape off the bark of trees or shrubs. Exposed roots were found in some of the deeper pits where the roots of trees were damaged and exposed to the air.
Damaged vegetation that was not closely associated with a lek was ignored, even though it might have been caused by fallow deer. Native black-tailed deer will feed on buds, twigs, sprouts, leaves, fruit, and flowers of woody plants. We never observed these deer in or around fallow deer leks, probably because fallow deer tend to be behaviorally dominant. Beef cattle were present in all of the Estero trailhead study area and part of the Olema Valley study area. While cattle will sometimes rub on vegetation and cause discernable vegetation damage, we did not observe this behavior at fallow deer lek sites. Additionally, most of the vegetation damage we observed was in riparian areas where cattle had been excluded by fencing.
During our initial surveys, we did not quantify vegetation condition at each lek, so we revisited 22 randomly selected leks in the Olema Valley to record the condition of both the soil and vegetation in more detail. We used a scoring system (Cole 1989a, 1989b) to evaluate the impacts of fallow deer (table 1). At each lek site, we scored ground surface disturbance, percent vegetative cover, damage to live trees, and presence of exposed roots. No attempt was made to quantify damage to individual trees or to assess likelihood of tree death resulting from the lek damage.
Table 1 (View this table on a separate page.) Codes used to evaluate ground surface disturbance, vegetation cover, tree damage, and root exposure at fallow deer leks in the Olema Valley, Point Reyes National Seashore, California.
 Surface disturbance
0No exposed roots seen
1Little disturbance to ground cover, whether cover is vegetation or litter
2Noticeable disturbance to litter or vegetation
3Moderate disturbance and bare ground
4Ground surface highly disturbed, extensive areas of bare ground
  
 Vegetation cover
150–100 percent
215–50 percent
35–15 percent
40–5 percent
  
 Live tree damage
0No tree damage
1A few small broken branches
2Several broken branches and/or scraping of bark
3Extensive bark gouging or scraping; broken branches may be present
  
 Root exposure
0No exposed roots seen
1One to two roots are up to 10 percent exposed (in cross–section) for <1 m
2Some roots 10–50 percent exposed for up to 1 m
3Some roots >50 percent exposed and/or exposed 10–50 percent for > 1 m

Photo documentation

A digital camera was used to document leks, rutting pits, and damaged vegetation. Bucks displaying and actively digging, or shredding foliage were also photographed, as were groups of does. Photographs were also taken of deer feces, hoof prints, and urine stains as evidence of deer use.

Mapping

UTM coordinates were recorded using a Garmin XL GPS unit. ArcView 3.3 software (ESRI, http://www.esri.com/software/arcgis/arcinfo/) was used to plot lek locations and to determine the size of the two study areas.

Results

Leks and Rutting Pits

A total of 159 fallow deer leks were located within the 298.8 ha (738.3 acres) surveyed at Point Reyes National Seashore (table 2, appendix 1). In the Olema Valley, there were 116 lek sites compared with 43 leks in the Estero trailhead area. The mean dimensions of a lek site were 13 × 7 m with an area of 115 m2 (SD = 132). The total area of the leks in the Olema Valley was 16,188 m2, while the area at the Estero trailhead was 2,136 m2 for a combined total of 18,324 m2 (4.5 acres). This was 0.6 percent of the 298.8 ha surveyed. There was a notably higher proportion of the Olema Valley study site that was part of a lek, 1.1 percent compared to 0.1 percent at Estero trailhead. In the Olema Valley, there were 0.8 leks per ha, while the Estero trailhead area had 0.3 leks per ha (table 2).
Table 2 (View this table on a separate page.) Number of fallow deer leks and rutting pits in two study areas at Point Reyes National Seashore, California.
[Numbers in parenthesis are Standard Deviations]
 Olema ValleyEstero TrailheadCombined
Study area size (ha)147.2151.6298.8
Number of leks11643159
Leks per ha0.80.30.5
Mean lek area (m²)140 (±142)50 (±60)115 (±132)
Total lek area (m²)16,1882,13618,324
Percent lek area (m²)1.1 percent0.1 percent0.6 percent
    
Number of rutting pits598107705
Total pit area (m²)1,4633581,821
Percent study area as pits 0.1 percent0.02 percent0.6 percent
Mean number pits/lek5.1 (±5.1)2.5 (±1.9)4.4 (±4.6)
Mean pit depth (cm)10 (±9)6 (±5)9 (±9)
Maximum pit depth (cm)601560
A total of 705 rutting pits were found in the two study areas, with an average size of 1 × 2 m, and an area of 2.6 m (SD = 3.0) for each pit. The mean number of pits per lek was 5.1 in the Olema Valley and 2.5 for Estero trailhead. The total combined area of excavated ground in rutting pits was 1,821 m2, or 0.6 percent of the total area surveyed. Eighty-five percent (598) of pits were found in the Olema Valley study area. Though fewer in number, the pits at the Estero trailhead study area were larger (3.3 m2) than the pits in Olema Valley (2.4 m2).

Vegetation Condition

There was vegetation damage at 110 (69.2 percent) of the lek sites (appendix 2). Damaged foliage was present at 102 (64 percent) of the lek sites. During initial surveys for leks, sites were often located by broken live oak or California bay branches that were visible from considerable distances. Low branches and bark adjacent to rutting pits were often heavily damaged. On several occasions, bucks were observed thrashing vegetation with their antlers, digging in the rutting pits, and displaying at their lek. Bark damage was recorded at 72 (45 percent) of the leks. Exposed roots were documented for 30 (19 percent) of the lek sites. In addition to having nearly three times as many leks, the Olema Valley study area had a higher percentage of sites with damaged foliage and a higher percentage of sites with damaged bark, but the result was not statistically significant when using an α = 0.05 for evaluating level of significance (x2 = 3.16, df = 2, p = 0.206).

Riparian Impacts

Vegetation damage was greater in riparian areas (compared with non-riparian) for both the damaged foliage and damaged bark categories (table 3). In riparian areas, willows and alders were the trees most often observed with damage. Several alders were completely girdled. Less commonly, there were exposed roots, especially in the Estero trailhead area (table 3). Overall, there was more damage in riparian areas, but the result was not statistically significant (x2 = 5.74, df = 2, p = 0.057). A larger sample size would quite likely result in a statistically significant difference.
Table 3 (View this table on a separate page.) Vegetation damage at fallow deer lek sites at Point Reyes National Seashore, California.
[The numbers under each study area are the number of leks in each category; the percents are the proportion of leks in that category]
 Olema ValleyEstero TrailheadCombined 
Riparian1715 percent819 percent2516 percent
Damaged foliage1588 percent788 percent2288 percent
Damaged bark1482 percent8100 percent2288 percent
Exposed roots16 percent338 percent416 percent
       
Non-riparian10086 percent3479 percent13484 percent
Damaged foliage7171 percent824 percent7959 percent
Damaged bark4545 percent618 percent5138 percent
Exposed roots2020 percent618 percent2619 percent

Vegetation and Soil Condition Scores

In our quantitative evaluation of the vegetation and soil at 22 random sites, surface disturbance ranged from 0 (no disturbance) to 4 (ground surface highly disturbed with extensive areas of bare ground) with a mean score of 1.6 and a median of 2 (table 4). The mean score corresponds to a damage level between "Little disturbance to ground cover" and "Noticeable disturbance to litter or vegetation." Both vegetation cover and live tree damage had a mean and median score of 1 (50–100 percent cover, and a few small broken branches). There were no roots exposed in the sample plots.
Table 4 (View this table on a separate page.) Vegetation data from randomly selected fallow deer leks in the Olema Valley, Point Reyes National Seashore, California.
[See Table 1 for a description of the codes]
LekSurface disturbanceVegetative coverLive tree damageRoot exposureSpecies composition
224210Oak, grass, hedge nettle, geranium
252100Bay, forget-me-not, nettles, cow parsnip, currant, sword fern
270000Oak, blackberry, nettles, Australian fireweed, hazelnut, grass, sedge
292120Bay, grass which is mostly purple velvet, oak, coffeeberry
332220Oak and littler, bay seedlings, grass, Italian thistle, nettles
342120Oak, litter, grass
402110Oak, grass, Italian thistle, mustard
423110Oak, grass, Italian thistle, mustard
430000Not recorded
461100Oak, bay, litter, hemlock, grass
522110Oak and litter, grass, hemlock
600000Oak, grasses
613220Oak, with little vegetation under the canopy.
642120Oak and oak litter, blackberry, fern, hazelnut, Italian thistle, poison oak
660000Oak and litter, grass (mostly Italian rye and purple velvet), bracken fern, blackberry
670000Oak, forbs (didn't record what)
822010Oak, grass, thistles, radish, poison oak
850020Oak, willow, grass, blackberry, poison oak, dock
902120Oak, grass, small forbs, blackberry, hemlock, mint, dock
971310Oak and oak litter. Nearby are blackberry, some grass.
1003210Oak, bay, poison oak, blackberry, Italian thistle, hazelnut.
Mean1.6 (±1.2)110 
Median2110 

Photograph documentation

Typically, leks were found at the edge of a woodland or at the edge of the low-hanging part of the canopy of isolated trees. Figure 4 shows a large area of bare ground at the edge of a coast live oak. Some rutting pits were more than 50 cm deep (fig. 5), often surrounded by an even larger area cleared of all vegetation. Other leks had only a modest depression and were identified by the lack of vegetation along with associated fecal material, hoof prints, and damage to woody vegetation (fig. 6). Rutting pits in close association with bushes and trees were often associated with significant damage to the woody vegetation, including broken branches, stripped bark, and sometimes, girdled trees (fig. 7). Fallow deer were observed using their antlers to clear vegetation, rub the trunk of trees, break limbs, and dig pits. Vegetation was sometimes caught in their antlers (fig. 8).
Figure 4. Figure 4.
Typical location for a fallow deer rutting pit, at the interface between a tree and the adjacent grassland or the edge of the low-hanging canopy, Olema Valley, Point Reyes National Seashore, California.
Figure 5. Figure 5.
Rutting pit and bare ground associated with a fallow deer lek in the Olema Valley, Point Reyes National Seashore, California.
Figure 6. Figure 6.
Bare ground in a fallow deer lek at edge of a coast live oak tree, Olema Valley, Point Reyes National Seashore, California.
Figure 7. Figure 7.
Broken willow branches at a fallow deer lek in the Olema Valley, Point Reyes National Seashore, California.
Figure 8. Figure 8.
Fallow deer buck with vegetation caught in his antlers, Olema Valley, Point Reyes National Seashore, California.

Discussion

Fallow deer are one of the few mammals that use a lek breeding system where males gather in groups and display to potential mates. As part of the display, male deer dig rutting pits with their hooves and antlers, and scrape bushes and trees adjacent to the pits. This results in soil disturbance, loss of vegetation, and occasionally damage to the trunks and limbs of nearby vegetation. The extent of the impacts was related to the density of fallow deer. This density was higher in the Olema Valley (Gates, 2001) and lek impacts were greater in that area, as would be expected. The Olema Valley had a greater mean lek area, total lek area, percentage of total area as leks, mean number of rutting pits per lek, percentage of total area as rutting pits, and mean pit depth (table 2). In the Olema Valley study area, more than 1 percent of the total land area surveyed was impacted by lek damage, with riparian areas being disproportionately affected. For example, in riparian areas of the two study areas, there was bark damage at 88 percent of the leks and 84 percent of the leks had damaged foliage (table 3).
The primary habitat impact of leks at Point Reyes was caused by the digging of rutting pits that resulted in a loss of soil and vegetation. At the peak of the rut, pits were commonly found in Olema Valley, especially at the woodland/grassland interface where fallow deer tend to congregate. The density of rutting pits was less in the Estero trailhead study area due to the smaller fallow deer population there.
Our research shows that fallow deer are having a measurable impact on the soil and vegetation at Point Reyes. In the Olema Valley, there were 0.8 leks per ha, while the Estero trailhead area had 0.3 leks per ha (table 2). The total number of rutting pits was 705 with a total area of 1,821 m2. Pélabon et al. (1999) have shown that the formation of leks in ungulates is "a mating tactic that aims at decreasing the number of aggressive encounters in which dominant males are involved when the local male density becomes too high." If the fallow deer population continues to increase, the number of leks, the number of rutting pits, and the associated habitat damage will likely increase.

Acknowledgments

Wende Rehlaender conducted the detailed soil and vegetation evaluations and provided those data to us for analysis. Patrick Kleeman provided assistance in data management. Kan Dhillon assisted with the initial surveys for fallow deer leks and with general advice on the location of fallow deer within the seashore. Dave Schriokaurer and Amelia Ryan provided GIS mapping assistance. Natalie Gates helped offered advice throughout the development and implementation of the project. Rick Golightly, Dale McCullough, Kurt Jenkins, Patrick Kleeman, and Joan Fellers provided comments on an earlier draft of this report. We greatly appreciate the assistance of all of the above.

Appendix

Table A1 (View this table on a separate page.) Locations of fallow deer leks at Point Reyes National Seashore, California.
Olema Valley SitesLek siteUTMEUTMN
    
2Cluster of Douglas Firs on Knob ~200 m SSW of Morgan Ranch house, Bear Valley5174964209939
3Lone pit at base of small bay5175354209914
4SE edge of grove of live oaks, Douglas Fir, and bay along Woodpecker and Bear Valley Trails5176254209915
5Within grove of live oaks, Douglas Fir, and bay along Woodpecker and Bear Valley Trails5176204209967
7Top of Knob at edge of field, along Bear Valley Trail5174604209845
8At base of Douglas Fir next to meadow behind Morgan Horse Ranch5172734210372
9At forest edge adjacent to Woodpecker Trail, 15m from Morgan Trail5173904209948
10Near large bay in meadow at Bear Valley between Earthquake Trail and Vadanta5179344209851
11Near base of live oak in meadow at Bear Valley between Earthquake Trail and Vadanta5179244209878
12Grove of bays in meadow in Bear Valley, south of Earthquake Trail5178944209936
13In groove of bays ~100m SE of SE corner of Bear Valley parking lot5178074209980
14Edge of grove of bays ~100 SE of Bear Valley parking lot5178424210007
1510 m S of Earthquake Trail, adjacent to bay tree5178574210053
16East edge of Divide Meadow5177764207778
17SE corner of Divide Meadow5177594207823
18East edge of Divide Meadow, next to live oaks5177854207705
19Small open area east of tree line, SE corner of Divide Meadow5178294207822
20At edge of live oaks along edge of Divide Meadow5178724207694
21Base of live oak at NE corner of Divide Meadow5177504207615
22Edge of NE corner of Divide Meadow, live oak5178004207587
23Under live oak at tree line of edge of Divide Meadow5177814207548
24Inside forest edge, under bay trees, NE corner of Divide Meadow5177694207516
25Forest edge under bay and alders, Divide Meadow5177064207462
26Under alder at N end of Divide Meadow5177084207459
27Under live oak, west of Bear Valley Trail, across from Divide Meadow5175714207660
2820m N of Earthquake Trail, at edge of meadow and bays in Bear Valley5179164210024
29Base of bay between Earthquake Trail and Rift Zone Trail5180124209969
3020 m from Rift Zone Trail in clearing at edge of Bear Valley meadow5180874209925
3120 m N of Rift Zone Trail on knob SE of Bear Valley, under live oaks5181744209983
32On knob 75 m N of Rift Zone Trail, under live oak5182174209989
33Adjacent to live oak, on knob, 75 m N of Rift Zone Trail5182074210015
34Under live oak, ~100 m N of Rift Zone Trail on W side of knob5182064210053
35Under live oak, ~125 m from Rift Zone Trail on knob5182194210070
36Under live oak, 20 m N of Rift Zone Trail5181784210049
37Under live oak, 30 m N of Rift Zone Trail on knob5181504209982
38At large bay, ~40 m N of Rift Zone Trail on knob5181384210048
39Under bay tree, ~50 m N of Rift Zone Trail on knob5181114210048
40Under lone live oak in meadow near top of knob N of Rift Zone Trail5180864210124
41Area at base of large oak and bay trees, on knob near Rift Zone Tail and Earthquake Trail5180624210116
42Near top of knob N of Rift Zone Trail at forest edge, ~20 m E of Earthquake Trail5180614210150
43At forest edge on knob N of Rift Zone Trail and E of Earthquake Trail5180874210094
44On slope 40 m NE of Earthquake Trail and 150 m SE of Red Barn parking lot5179704210251
45Area around small live oak on top of knob, NE of Earthquake Trail5180254210213
46Field at base of knob between Rift Zone Trail and Bear Valley Rd., ~100 m SW of BV Road5181834210311
47In grove of live oaks at base of knob ~100 m SW of Bear Valley Rd5181464210301
48Grove of live oaks on East side of top of knob, 250 m SW of BV Rd. and 220 m N of Rift Zone Trail5181784210152
49Area surrounding live oak in field on top of knob, 0.5 km W of Olema5181584210140
50Edge of field at grove of live oaks, ~0.5 km E of Red Barn and 0.5 km W of Olema5181634210218
51Grove of live oaks and bays on NE corner of knob, ~0.5 km W of Olema5181544210245
52Grove of oaks near base of NE corner of knob, 0.5 km W of Olema5182204210221
53Edge of meadow in grove of oaks5175444211545
54At edge of live oak grove on SW edge of hills between BV Rd. and Hwy 15175224211568
55In grove of live oaks in pasture between BV Rd. and Hwy 15175124211590
56Grove of willows and live oaks at base of east-facing slope between BV Rd. and Hwy 15175924211546
57Edge of pasture and treeline, 390 m N of Headquarters5176844210871
58Edge of pasture 410 m N of Headquarters5176824210887
59Near live oak on SW-facing slope of hill NE of BV Rd., 1.15 km NNW of Headquarters5173104211503
60Stewart's pasture at base of two live oaks, 1.4 km NNW of Headquarters5174384211544
61Under live oak, 1.0 km NNW of Headquarters5174204211507
62Base of live oak in Stewart's pasture NE of Bear Valley Rd., 1.17 km NNW of Headquarters5174874211564
63Under live oaks in grove at NE edge of Stewart's pasture, 1.2 km NNW of Headquarters5174924211601
64Under live oaks on NE side of hill in Stewart's Pasture NE of Bear Valley Road, 1.22 km NNW of Headquarters5174524211635
65Underneath live oak tree, 550 m N of Headquarters5177214211065
66Grove of live oaks, 550 m NNE of Headquarters5178034211003
67Under live oaks at base of hill on east facing slope, 550 m NNE of Headquarters5177964211020
68Stand of willows at east-facing slope of hill, 510 m NE of Headquarters5178444210995
69Under alders along creek, 510 m NE of Headquarters5179034210968
70Alders along Olema Creek, 600 m NNE of Headquarters5178874211061
71Adjacent to Olema Creek on gravel bar in stand of alders, 560 m NNE of Headquarters5178864211111
72Edge of pasture and stand of willow and alder, along Olema Creek, 580 m NNE of Headquarters5178504211143
73Edge of pasture along fence, creek, and stand of willow and alder, 610 m NNE of Headquarters5178394211170
74Edge of pasture along fence, creek, and willows, 860 m N of Headquarters5177854211309
75Adjacent to creek in stand of willows and alders, 900 m N of Headquarters5177684211361
76Pasture, 910 m N of Headquarters5177394211369
77Border of pasture and riparian, 940 m N of Headquarters5177434211393
78Pasture, 1.0 km N of Headquarters5177064211447
79Pasture, 1.01 km N of Headquarters5176914211476
80Under live oaks at base of E slope of hill, 1.02 km N of Headquarters5176184211496
81Grove of willows, 1.1 km N of Headquarters5176144211527
82Top of hill by lone fallen live oak, 1.25 km NW of Headquarters5173274211647
83Edge of forest under live oak, near top of hill, 1.22 km NW of Headquarters5172674211628
84Grove of oaks and small clearing at forest edge, 1.3 km NW of Headquarters5171854211672
85Edge of treeline in pasture NE of Bear Valley Rd., 1.1 km NW of Headquarters5172684211494
86Treeline 30 m NE of Bear Valley Rd. at pasture edge, 970 m NW of Headquarters5172974211458
87Treeline at edge of pasture NE of Bear Valley Rd., 930 m NW of Headquarters5174054211330
88Under oak in clearing 15m NE of Bear Valley Rd., 840 m NW of Headquarters5174424211244
89Grove of oak and bay NE of Bear Valley Rd., 600 m NNW of Headquarters5175704211039
90Grove of oaks along Bear Valley Road, 580 m NNW of Headquarters5175944211046
91Grove of oaks bordering Bear Valley Rd., 520 m NNW of Headquarters5175954210987
92Grove of oaks bordering Bear Valley Rd., 480 m NNW of Headquarters5176204210966
93Under lone tree at SE end of pasture, 320m NNE of headquarters5178764210789
94Grove of live oaks adjacent to Olema Creek, 480m NE of headquarters5179544210882
95Grove of oak and bay at East edge of pasture, along Olema Creek, 500 m NE of headquarters5179454210911
96Grove of oak and bay along Olema Creek, 500 m NE of headquarters5179214210928
97At base of live oak, along treeline at edge of pasture 1.36 km NW of Headquarters5171574211873
98Forest edge of pasture, 1.30 km NW of Headquarters5171884211892
99Forest edge of pasture, 1.40 km NW of Headquarters5170954211738
100Edge of pasture and forest, under oak and bay 1.42 km NW of Headquarters5170794211764
101Under lone live oak in pasture, 1.6 km NW of Headquarters5169834211907
102Open pasture, 1.7 km NW of Headquarters5169744212069
103In open pasture on top of knob, 1.73 km NW of Headquarters5170164212074
104Open pasture on top of knob, 1.7 km NW of Headquarters5170454212027
105Small cluster of willow in field east of Olema Marsh5170044212337
106Patch of invasive plants in field east of Olema Marsh5169744212376
107Edge of willows and field along Olema Creek, east of Olema Marsh5169824212418
108Edge of willows and field along Olema Creek, east of Olema Marsh5170044212415
109Under willows along Olema Creek, east of Olema Marsh5170344212360
110Edge of riparian willows along Olema Creek, 400 m east of Olema Marsh5171364212262
111Next to willow, 400m east of Olema Marsh5171184212252
112Edge of oak grove and pasture 1.45 km NW of Headquarters5173044211868
113Under live oak at edge of pasture near Olema Creek, 1.37 km NNW of Headquarters5174504211826
114Base of live oak in pasture, 1.27 km NNW of Headquarters5174464211695
115Next to bay tree at NE end of Bear Valley picnic area5177324210281
116Next to bay at edge of Bear Valley picnic area5178094210229
117Edge of treeline at SE end of Bear Valley picnic area5178034210182
118Grassy area at edge of tree line along road, 100 m South of Headquarters5176794210336
    
Estero trailhead sites   
119Open pasture, near Home Ranch, 200 m E of Estero Trail parking lot5078284214722
120Open pasture, near Home Ranch, 200 m E of Estero Trail parking lot5078354214696
121Open pasture, near Home Ranch, 250 m ENE of Estero Trail parking lot5078614214769
122Open pasture, near Home Ranch, 300 m ENE of Estero Trail parking lot5078974214811
123Patch of thistle, near Home Ranch, 260 m ENE of Estero Trail parking lot5078704214797
124Open pasture, near Home Ranch, 220 m ENE of Estero Trail parking lot5078004214790
125Open grassland on ridge top, near Home Ranch, 750 m WNW of Estero Trail parking lot5068854214833
126Open grassland on ridge top, near Home Ranch, 650 m WNW of Estero Trail parking lot5070244214838
127Top of point West of Home Bay, 1.5 km SW of Estero Trail parking lot5064704213642
128Ridge top 1.1 km W of Estero Trail parking lot5065094214534
129Ridge top 1.0 km WSW of Estero Trail parking lot5066924214484
130Open grassland on ridge top 700 m NW of Estero Trail parking lot5072754215275
131Open pasture, near Home Ranch, 200 m ENE of Estero Trail parking lot5077984214783
132Top of Knob, near Home Ranch, 500 m NE of Estero Trail parking lot5079614215090
133Top of Knob, near Home Ranch, 450 m NE of Estero Trail parking lot5079374215053
134Top of Knob, near Home Ranch, 550 m NE of Estero Trail parking lot5079414215133
135Side of Knob, near Home Ranch, 650 m NE of Estero Trail parking lot5080974215160
136Side of Knob, near Home Ranch, 600 m NE of Estero Trail parking lot5080854215122
137Side of Knob, near Home Ranch, 400 m NE of Estero Trail parking lot5079454214969
138Side of Knob, near Home Ranch, 370 m NE of Estero Trail parking lot5079274214944
139Pasture on knob 275 m NE of Estero Trail parking lot5078344214889
140Pasture on NW side of knob 500 m NE of Estero Trail parking lot5079274215134
141Pasture on knob 600 m NNE of Estero Trail parking lot5079504215280
142Pasture 900 m NE of Estero Trail parking lot5082054215368
143Bare ground adjacent to coyote brush 800 m NNE of Estero Trail parking lot5079414215429
144Pasture 500 m NNE of Estero Trail parking lot5077584215188
145Pasture 150 m N of Estero Trail parking lot5076674214877
146Edge of pasture along unnamed stream, 450 m SE of Estero Trail parking lot5079654214373
147Edge of pasture along unnamed stream, 500 m ESE of Estero Trail parking lot5081304214576
148Edge of pasture along unnamed stream, 475 m ESE of Estero Trail parking lot5081444214601
149Edge of pasture along unnamed stream, 510 m ESE of Estero Trail parking lot5081504214625
150Edge of pasture along unnamed stream, 525 m E of Estero Trail parking lot5081904214700
151Edge of pasture along unnamed stream, 550 m E of Estero Trail parking lot5082144214747
152Pasture in saddle 850 m NE of Estero Trail parking lot5082974215253
153Edge of pasture along unnamed stream, 650 m ENE of Estero Trail parking lot5082874214922
154Pasture 600 m N of Estero Trail parking lot5075984215337
155Pasture 650 m N of Estero Trail parking lot5076224215353
156Pasture 400 m W of Estero Trail parking lot5072654214667
157Grove of willows along unnamed creek, 350 m W of Estero Trail parking lot5072904214685
158Pasture on ridge top, 800 m NW of Estero Trail parking lot5068884215054
159Scrub 750 m WSW of Estero Trail parking lot5068844214571
160Scrub 700 m WSW of Estero Trail parking lot5069994214505
161Edge of willows along unnamed creek, 600 m WSW of Estero Trail parking lot5071164214443
Table A2 (View this table on a separate page.) Description of leks and associated ground and vegetation damage at the two study sites at Point Reyes National Seashore, California.
Olema Valley sitesLength (m)Width (m) Area (m2)Number rutting pitsGround disturbanceDamaged foliageDamaged barkExposed roots
220.016.0320.05XX-X
37.02.014.01XXX-
425.015.0375.05XX--
58.02.016.03X---
78.02.016.02XX--
83.01.54.51XX-X
925.05.0125.04XX-X
102.01.02.01X---
113.01.54.51X---
1213.010.0130.06XX-X
132.51.23.01X--X
1420.020.0400.08XX-X
152.01.42.81X---
1625.010.0250.04X---
1710.02.020.02X-XX
1820.015.0300.04XXXX
1910.010.0100.04XXXX
204.51.04.52XX-X
2115.010.0150.05X---
222.01.12.23X---
2316.05.080.05X---
2420.010.0200.06X--X
2515.02.030.04X---
264.01.04.01X---
272.51.12.751X---
283.01.23.62XXXX
294.51.56.753XX--
3017.02.034.02XX--
3120.010.0200.04XX-X
325.04.020.01X---
3317.015.0255.08XX--
3416.010.0160.04XX--
355.02.010.02XX--
3610.03.030.03X---
3720.017.0340.016XX-X
3820.018.0360.07XX-X
395.05.025.01XX-X
4020.017.0340.02X---
4110.05.050.05XXXX
4216.08.0128.09XX-X
4310.05.050.03XXXX
4410.05.050.01XXXX
458.05.040.01XX--
4617.05.085.06X---
4715.05.075.04XX--
4817.013.0221.09XXX-
4914.010.0140.011XXX-
5010.05.050.03X-X-
5133.012.0396.012XXX-
5220.010.0200.08XXX-
5320.010.0200.08XXX-
5418.010.0180.08XXX-
5510.02.020.02X---
5625.010.0250.04XXX-
5725.08.0200.08X---
5820.05.0100.04X---
5915.05.075.02XX--
6035.024.0840.022XXX-
6120.010.0200.06XXX-
6218.010.0180.06XXX-
6310.010.0100.02XX--
645.04.020.02X---
6513.05.065.09XXX-
6624.010.0240.09XXX-
6720.010.0200.015XXX-
6810.05.050.03XXX-
6910.08.080.01XXX-
7010.05.050.01XXX-
7115.05.075.05XXX-
725.05.025.01XXX-
734.04.016.01XXX-
7420.04.080.09XXX-
7510.05.050.03XXX-
768.05.040.01X---
7727.08.0216.012XXXX
7810.010.0100.02XX--
791.01.01.01X---
8034.08.0272.05XXX-
8120.020.0400.04XXX-
8215.015.0225.06X---
838.05.040.02XX--
8420.015.0300.09XXX-
8510.05.050.02XXX-
868.05.040.01XXX-
8727.010.0270.018 XXX-
8816.07.0112.03XX--
8920.010.0200.05XX--
9015.012.0180.011XXX-
9132.020.0640.024XXX-
9220.012.0240.07XXX-
9314.010.0140.08XXX-
9430.017.0510.030XXX-
9520.015.0300.05XXX-
9625.010.0250.07XXX-
9715.013.0195.02XXX-
9815.010.0150.05XX--
9915.010.0150.02XXX-
1008.05.040.04XXX-
10110.05.050.01XXX-
10210.05.050.01X---
1035.03.015.01X---
10425.010.0250.05X---
10517.05.085.03XXX-
10620.010.0200.03XX--
10715.010.0150.03XXX-
10810.05.050.02X---
10910.04.040.04XXX-
11015.05.075.03XXX-
1118.03.024.01XXX-
11210.03.030.01XXX-
11321.05.0105.07XXX-
11410.04.040.05XXX-
11530.05.0150.08XXX-
1165.05.025.01XXX-
11732.015.0480.024XXX-
11817.06.0102.06XX--
         
Estero trailhead sites        
11912.07.084.05X---
1203.03.09.02X---
12120.07.0140.04X---
12215.07.0105.05X---
12313.010.0130.04X---
12410.05.050.03X---
1253.02.06.01X---
1262.02.04.01XXX-
12710.010.0100.06X---
12810.04.0401XX-X
12910.03.030.03X---
13010.04.040.05XX--
13120.010.0200.04X---
1322.02.04.01X---
13310.02.525.02X---
1342.02.04.01X---
13510.03.030.02XX-X
1363.02.06.01X---
1375.02.010.03XXXX
1382.02.04.01XXX-
1397.04.028.02X---
1408.03.024.03X---
1413.02.06.01XX-X
14225.010.0250.09X---
1435.05.025.01XXXX
14415.02.030.02X--X
1452.02.04.01XX--
14610.08.080.02XXX-
14718.08.0144.06X-X-
14810.04.040.03X-X-
14915.010.0150.05XXXX
15015.05.075.06XXX-
1517.05.035.01XXX-
1523.02.06.01X---
1535.02.010.01X--X
15412.04.048.03X---
1555.01.57.52X---
15610.03.030.01XXX-
15715.010.0150.02XXXX
1584.04.016.01X---
1592.01.02.01X---
1605.01.758.752X---
16110.03.030.01XXX-
         
Totals--18,3247051591027230
Mean13.06.71154.4    
Percentage----10063.545.318.2

References

Apollonio, Marco, Festa-Bianchet, Marco, and Mari, Franco, 1989, Correlates of copulatory success in a fallow deer lek: Behavioral Ecology and Sociobiology, 25, no. 2, p. 89–97.

Cole, D.N., 1989a, Area of Vegetation Loss: A New Index of Campsite Impact: U.S. Forest Service, Intermountain Research Station. Research Note INT, v. 389, 5 p.

Cole, D.N., 1989b, Wilderness Campsite Monitoring Methods: A Sourcebook: U.S. Forest Service, Intermountain Research Station. General Technical Report INT, v. 259, 60 p.

Gates, Natalie, 2001, Aerial and ground censuses of non-native deer, November 2000 and January 2001, Point Reyes National Seashore: NPS unpublished report, Point Reyes National Seashore, Point Reyes, Calif., 21 p.

Hirth, D.H., 1997, Lek breeding in a Texas population of fallow deer (Dama dama): American Midland Naturalist, v. 138, p. 276–289.

Hovi, M., Alatalo, R.V., and Rintamäki, P.T., 1996, Habitat differences and variability in the lek mating system of black grouse: Behaviour, v. 133, p. 561–578.

Pélabon, C., Komers, P.E., and Höglund, J., 1999, Do leks limit the frequency of aggressive encounters in fallow deer? Linking local male density and lek occurrence: Can. J. Zool. v. 77, p. 667–670.

Stenström, D., Dahlblom, S., Jones Fur, C. and Höglund, J., 2000, Rutting pit distribution and the significance of fallow deer Dama dama scrapes during the rut: Wildlife Biology, v. 6, p. 23–29.

Wehausen, J.D., 1973, Some aspects of the natural history and ecology of fallow deer on Point Reyes peninsula: University of California, Davis, M.S. thesis, 68 p.