Crossosoma 33(2R), Fall-Winter 2007 45
PLANT SUCCESSION IN THE EASTERN MOJAVE DESERT: AN EXAMPLE
FROM LAKE MEAD NATIONAL RECREATION AREA, SOUTHERN NEVADA
Scott R. Abella
Public Lands Institute and School of Life Sciences
University of Nevada Las Vegas
4505 S. Maryland Parkway
Las Vegas, NV 89154-2040
[email protected]
Alice C. Newton
National Park Service
Lake Mead National Recreation Area
601 Nevada Way
Boulder City, NV 89005
-and-
Dianne N. Bangle
Public Lands Institute
University of Nevada Las Vegas
4505 S. Maryland Parkway
Las Vegas, NV 89154-2040
ABSTRACT: Plant succession remains a poorly understood process in the Mojave
Desert, yet knowledge is needed in this area where increasing human populations may
amplify disturbance frequencies and intensities. In a retrospective study, we examined
plant communities on two pipeline corridors, one cleared in 1998 and one in 1968 to
supply water to metropolitan Las Vegas, Nevada. We also evaluated the effectiveness of
restoration treatments (raking soil surfaces, spreading artificial desert varnish, and
planting four species of native shrubs) applied by the National Park Service on the 1998
corridor to enhance recovery of Larrea tridentata communities. Plant cover was sparse (<
5%) on the untreated 1998 corridor eight years after clearing, with a mean shrub density
of only 99/ha. On the restoration-treated area, however, L. tridentata established at a
density of 300/ha, 36% of the density of an adjacent control area. Restoration treatments
also made the corridor less visually distinct from surrounding L. tridentata communities.
Even 38 years after clearing, the older corridor was dominated by species such as
Stephanomeria pauciflora and Encelia farinosa, which are classified as early colonizers
in the Mojave Desert. Our findings concur with long recovery estimates after vegetation-
removing disturbances given in the literature, but suggest that ecological restoration has
potential for manipulating the speed and trajectory of plant succession in the Mojave
Desert.
KEYWORDS: disturbance, ecological restoration, Encelia farinosa, Hymenoclea
salsola, Larrea tridentata, Stephanomeria pauciflora, vegetation.
46 Crossosoma 33(2R), Fall-Winter 2007
INTRODUCTION
Mining, military activities, off-road vehicles, agriculture, livestock grazing, and land
clearing for linear corridors (e.g., roads, power lines) are some of the many types of
human disturbances impacting Mojave Desert ecosystems (Lovich and Bainbridge 1999).
Plant succession (rate and species composition) following these disturbances can vary
with disturbance type (Webb et al. 1987), disturbance size (Hunter et al. 1987),
precipitation (Brum et al. 1983), time since disturbance (Carpenter et al. 1986), and also
with other less-documented factors such as soil type (Lathrop and Archbold 1980).
Larrea tridentata (DC.) Cov. communities, which are a dominant vegetation type in the
Mojave Desert, have generally taken decades to more than centuries to approximate pre-
disturbance plant composition (Lovich and Bainbridge 1999).
Vasek (1979/1980) documented plant succession in the southeastern Mojave Desert nine
years after land clearing for a highway borrow pit in southern California. He found that
early colonizers included Ambrosia dumosa (A. Gray) Payne, Encelia frutescens (A.
Gray) A. Gray, Stephanomeria pauciflora (Torrey) Nelson, and Porophyllum gracile
Benth. These species exhibited 19-177 times greater densities in the disturbed pit bottom
than in adjacent Larrea tridentata communities. Vasek (1983) further classified Mojave
Desert perennial species into three main successional categories: early colonizers that
respond strongly and positively to disturbance and have short individual life spans (e.g.,
Hymenoclea salsola A. Gray, S. pauciflora, Encelia spp.), long-lived opportunistic
species important in older communities but also exhibiting pioneering ability (e.g., A.
dumosa, Opuntia bigelovii Engelm.), and long-lived perennials that recover slowly from
disturbance (e.g., L. tridentata). Vasek (1983) also noted that many early colonizers after
human disturbance are abundant in frequently disturbed “natural” habitats such as
washes, and that annual plants occur in both early and late-successional communities.
Specific questions about succession, such as factors affecting its rate and trajectory,
remain poorly understood in the Mojave Desert, and in deserts in general (Bolling and
Walker 2000). This hinders ecological management in deserts, where increasing human
populations may intensify disturbance levels (Kemp and Brooks 1998; Lovich and
Bainbridge 1999). In a retrospective study in the eastern Mojave Desert, we assessed
plant community and soil characteristics on two water pipeline corridors cleared of
surface soil and vegetation eight (1998) and thirty-eight (1968) years before this study.
The National Park Service also applied restoration treatments designed to speed recovery
of Larrea tridentata communities on part of the 1998 corridor. Both corridors cross
National Park Service land (Lake Mead National Recreation Area [LMNRA]) and were
constructed to supply water to the metropolitan Las Vegas Valley. Since further water
developments are planned to occur within LMNRA, this study was intended to evaluate
potential for natural succession and ecological restoration of these disturbances.
Additionally, our study adds site-specific successional data needed to build general
theories of succession for the Mojave Desert. We sought to answer the following
questions at this site: (1) What is species composition, shrub density, and species richness
on corridors cleared in 1998 and 1968 relative to adjacent L. tridentata communities? (2)
On the 1998 corridor, do soil properties differ among treatments and below L. tridentata
compared to interspaces between shrubs? (3) How does species composition on this site
Crossosoma 33(2R), Fall-Winter 2007 47
after disturbance compare with other successional sequences described for the Mojave
Desert?
METHODS
Study Area and Pipeline Treatments
This study was conducted in LMNRA, Clark County, Nevada, 30 km east of the city of
Las Vegas at an elevation of 400 m (UTM 696000 m E, 3993000 m N; zone 11;
NAD83). The study area consisted of a 0.21-ha area in each of four adjacent locations: a
1998 corridor receiving no restoration treatments (hereafter untreated 1998 corridor), an
adjacent section of the same corridor that received restoration treatments (hereafter
treated 1998 corridor), an adjacent Larrea tridentata community off the corridor that
served as a control, and a corridor cleared in 1968 adjacent to the 1998 corridor (Fig. 1).
This study of a pre-existing disturbance and an operational management activity is
limited by a lack of replication; however, the study area comprises one landform (an
alluvial fan) and one soil association (Carrizo-Carrizo-Riverbend, primarily consisting of
Typic Torriorthents; Lato 2006). This supports an assumption that potential differences
among the four areas primarily result from their successional age or the restoration
treatments, rather than from pre-existing environmental differences. Both the treated and
the untreated 1998 corridor were cleared by blading with heavy equipment, with the
upper 20 cm of soil stockpiled and reapplied after construction. The 20-cm depth may
have varied slightly depending on rockiness or other factors. The 1968 corridor also was
cleared by mechanical blading, but topsoil was not replaced (David Connally, Southern
Nevada Water Authority, personal communication).
Restoration treatments applied by the National Park Service in January-February 1999 to
the 1998 corridor included hand-raking the soil surface after soil replacement to re-spread
rocks, applying artificial desert varnish (product name = permeon, Soil-Tech, Inc., Las
Vegas, NV) evenly to the soil surface for color restoration, and planting Larrea tridentata
(96 plants), Ambrosia dumosa (12 plants), Opuntia basilaris Engelm. & J. Bigelow (9
plants), and Acacia greggii A. Gray (2 plants). Desert varnish is a brown-black coat given
its color by iron and manganese oxides. This varnish commonly forms on stable surfaces
on volcanic rock, a process that is hastened by the chemical application of artificial desert
varnish (Moore and Elvidge 1982). The planting treatment is detailed in Newton (2001).
By 2001, no planted A. greggii or A. dumosa were alive, but survival was 92% for L.
tridentata and 100% for O. basilaris (Newton 2001).
Annual precipitation from 1999-2005 after clearing of the 1998 corridor averaged 105%
of the long-term (32 yr) mean (14 cm/yr), measured at Willow Beach, AZ, 26 km south
of the study site (Western Regional Climate Center, Reno, NV). As is typical of the
Mojave Desert, however, precipitation ranged widely from 27% (2002) to 205% (2004)
of the long-term mean among years.
Field Sampling
Between 31 August and 25 October 2006, we delineated a 30 × 70 m section in the
centers of each of the four areas. Within these sections, we randomly established a 10 ×
70 m transect divided into seven 10 × 10 m (0.01 ha) plots. Using simple random
sampling, we selected three plots in each section for sampling. Within each plot, we
48 Crossosoma 33(2R), Fall-Winter 2007
Fig. 1 (a) Fig. 1 (b)
Fig. 1 (c) Fig. 1 (d)
Figure 1. Views of (a) bladed 1998 water pipeline corridor that received no
restoration except for soil replacement; (b) the same corridor that received the
restoration treatments of raking the soil surface, applying artificial desert
varnish, and planting four species of native shrubs in addition to soil
replacement; (c) 1968 water pipeline corridor; and (d) control area (note the
intact desert pavement) adjacent to the corridors, Lake Mead National
Recreation Area, southern Nevada. Photos by S.R. Abella, 8 October 2007.
established six 1 × 1 m subplots per plot located at the plot corners and at the midpoints
(5 m) of southern and northern plot lines. We visually estimated areal percent cover of
each plant species rooted in subplots using a 1-m2 frame divided into 25, 0.04 m2
compartments. We also surveyed whole plots on a presence/absence basis for species not
occurring in subplots. Our sampling time in late summer and fall was not designed to
capture live annuals, but we recorded standing dead annuals in subplot and plot sampling.
Live shrubs, including seedlings, were counted on each plot. Nomenclature and
native/exotic species classifications follow Baldwin et al. (2002). To compare soils
among the control and the treated and untreated 1998 corridor, we collected a 0-10 cm
soil sample in an interspace (≥ 1 m away from any shrub) at the northwestern and
southeastern corners of each plot and composited these samples on a plot basis. We also
selected a dominant Larrea tridentata on each plot on the control and on the treated 1998
corridor (the untreated corridor contained no L. tridentata) and collected four, 0-10 cm
soil samples (composited on a plot basis) halfway between the main stem and the canopy
edge.
Crossosoma 33(2R), Fall-Winter 2007 49
Laboratory and Data Analysis
Air dried < 2 mm fractions of soil samples were analyzed for pH (saturated paste), total P
and K (Olsen NaHCO3 method), total C and N (Leco C/N analyzer), and texture
(hydrometer method). We compared mean (n = 3 for each area) species richness and total
mean shrub density among the control, treated and untreated 1998 corridor, and the 1968
corridor using one-way analysis of variance and Tukey’s test in JMP (SAS Institute
2004). Analysis of variance also was used to compare interspace soils among the control
and 1998 corridor areas. For the control and the treated 1998 corridor, we used paired t
tests to compare soil properties between interspaces and below Larrea tridentata.
Statistical results should not be extrapolated to other sites since treatments were not
replicated, but mean comparisons are presented as interpretational aids.
RESULTS
1998 Corridor
Exotic species richness/m2 was similar between treated and untreated areas in the 1998
corridor but was greater than in the control (Fig. 2). Total species richness/100m2 was
similar among treatments, ranging from 8-9.3 species. The exotic annual grasses
Schismus spp. exhibited the highest relative cover on the 1998 corridor compared to the
control, but total absolute cover for all species on the corridor was only 5-6% (Fig. 3).
Relative cover of the native annual Plantago ovata Forsskal was highest in the control,
intermediate in the treated corridor, and lowest in the untreated corridor. Perennial forbs
and grasses were sparse or absent from all treatments. Shrub density was eight times
higher in the control than in the untreated corridor, which contained no Larrea tridentata
(Fig. 4). Ambrosia dumosa and Encelia farinosa Torrey & A. Gray were the only shrubs
inhabiting the untreated corridor, and these species did not occupy plots on the treated
corridor or on the control. Larrea tridentata exhibited a density of 300/ha on the treated
corridor, which was 36% of the density on the control.
In interspaces, soil properties were similar among the treated, untreated, and control areas
except for K, which was significantly greater on the untreated corridor than on the control
(Table 1). Sand concentration was 10% higher and silt 9% lower on the untreated
corridor compared to the control, but all soils were still sandy loams. P and K both tended
to be greater below Larrea tridentata than in interspaces for the treated corridor and the
control, but the only difference that was statistically significant (p < 0.05) was for P for
the control.
1968 Corridor
Exotic species richness was lowest in the 1968 corridor relative to the control or the 1998
corridor, and total richness/100 m2 was comparable to both the 1998 corridor and the
control (Fig. 2). Similar to the 1998 corridor, Plantago ovata was a major contributor to
relative cover, although Stephanomeria pauciflora exhibited the highest relative cover.
Total shrub density averaged 3134/ha, 31 times more than the untreated 1998 corridor,
nine times more than the treated 1998 corridor, and four times more than the control.
Stephanomeria pauciflora and Hymenoclea salsola contributed 76% of the total shrub
density.
50 Crossosoma 33(2R), Fall-Winter 2007
Figure 2. Mean plant species richness at (a) 1 m2 and (b) 100 m2 scales among water
pipeline corridors and a control area, Lake Mead National Recreation Area, southern
Nevada. Error bars are 1 SD for total mean richness. In comparisons within native or
exotic categories, only exotic species per m2 and per 100 m2 differed significantly (p <
0.05) among the four areas.
Figure 3. Relative cover of dominant plant species and genera among water pipeline
corridors and a control area, Lake Mead National Recreation Area, southern Nevada.
CHASPP = Chamaesyce spp., CRYSPP = Cryptantha spp., ERIDEF = Eriogonum
deflexum, LARTRI = Larrea tridentata, PLAOVA = Plantago ovata, SCHSPP =
Schismus spp., and STEPAU = Stephanomeria pauciflora. Numbers at the top of each bar
represent total mean absolute % cover.
Crossosoma 33(2R), Fall-Winter 2007 51
Figure 4. Shrub densities among water pipeline corridors and a control area, Lake Mead
National Recreation Area, southern Nevada. Error bars are 1 SD for total mean density.
Means without shared letters differ at p < 0.05 for total density.
Table 1. Comparison of 0-10 cm soil properties among treatments and between
interspaces and below Larrea tridentata within treatments on an eight-year-old (1998)
water pipeline corridor, Lake Mead National Recreation Area, southern Nevada.
_________________________________________________________________
Untreated corridor Treated corridor1 Control
___________________________________________________
Property Interspace Larrea Interspace Larrea Interspace Larrea
________________________________________________________________
pH 8.1±0.12 ––3 8.1±0.2 8.1±0.1 8.1±0.1 7.9±0.1
P (mg/kg) 4.0±1.2 –– 3.5±0.7 3.7±0.8 4.1±1.6 11.5±2.9
K (mg/kg) 555±13a –– 491±62ab 575±204 400±61b 552±42
C (mg/kg) 942±35 –– 716±118 954±107 686±180 736±90
N (mg/kg) 27±5 –– 37±14 43±8 57±27 50±15
Sand (% wt.) 70±2a –– 65±3b 67±4 60±1b 66±6
Silt (% wt.) 24±1b –– 29±3a 27±3 33±2a 28±5
Clay (% wt.) 6±2 –– 6±0 6±1 7±1 6±2
________________________________________________________________
1
Restoration treatments included raking the soil surface, applying artificial desert varnish, and
planting four species of native shrubs.
2
Vales are mean ± SD (n = 3 within each treatment and canopy combination). Letters within a
row compare means among treatments for interspaces only. Values in bold denote significant
differences at p < 0.05 between interspaces and below Larrea tridentata within treatments.
3
Not measured because L. tridentata did not occur in this treatment.
52 Crossosoma 33(2R), Fall-Winter 2007
DISCUSSION
Although this assessment of an existing disturbance and an unreplicated operational
management activity supported only limited statistical inference, our findings represent a
case study of succession after land clearing in the eastern Mojave Desert, and how a
particular set of restoration treatments may influence succession. Effects of individual
restoration treatments cannot be discerned in this study, but the set of treatments
including surface raking, applying artificial desert varnish, and planting of shrubs,
appeared to make shrub composition on the treated 1998 corridor converge with that of
the control (Fig. 4). Although our study was not designed to track survival of individual
plants in the planting, we found that Larrea tridentata established on the treated 1998
corridor at a density 36% of that of the control. No L. tridentata established on the
untreated 1998 corridor. Previous studies of L. tridentata outplanting have produced
widely differing results, ranging from complete mortality (Graves et al. 1978) or < 2%
survival (Brum et al. 1983), to > 90% survival (Wallace et al. 1980; Clary and Slayback
1984; Newton 2001). In our view, the restoration treatments also made the 1998 corridor
appear more similar to surrounding L. tridentata communities, an important consideration
on National Park Service lands (Fig. 1). This visual blending resulted from the L.
tridentata establishment and also probably from the artificial desert varnish. Potential
ecological effects of this darkening varnish remain unclear, but it could result in warmer
soil temperatures or other effects. One of the largest aesthetic differences between the
treated and untreated 1998 corridor and the control was that the control contained desert
pavement. Pavement can require millennia to form (Elvidge and Iverson 1983), and it is
unclear whether the raking portion of the restoration suite had an effect or will have an
effect on surface layers such as desert pavement.
The approximately 20 cm of upper soil was salvaged, stockpiled, and reapplied after
blading on both the treated and untreated 1998 corridor. Although salvage operations add
logistical challenges and expense to projects, ecological effects of soil salvage are not
well known in the Mojave Desert and require further study. Soil salvage effects cannot be
evaluated in this study of an operational project because this would have required areas
on the 1998 corridor that were bladed but did not have soil replaced. The effects of soil
salvage and replacement after disturbance may depend on several factors, such as soil
type, depth of salvage, and length of time soil is stored (Bainbridge et al. 1998). Effects
also hinge on whether or not nutrients and seed banks are diluted upon reapplication by
mixing upper and lower soil layers (Nelson and Chew 1977). Based on soil seed bank
sampling in the northern Mojave Desert, Guo et al. (1998) reported that 91% of the total
seeds were in the upper 2 cm of soil and only 9% occurred from 2-10 cm. Using these
data and assuming that the 20 cm of salvaged soil in our study was evenly mixed during
salvage operations, the upper 2 cm (likely the germination zone) of the salvaged soil
would contain only 10% of the original seeds. Further reductions in viable seeds may
occur during topsoil handling or storage. However, it is possible that soil salvage could
result in nutrient retention. For example, Rundel and Gibson (1996) reported that total N
below shrubs in the northern Mojave Desert was approximately two or more times more
concentrated in the upper 5-9 cm of soil than in deeper layers.
Shrub species composition on the 38-year-old 1968 corridor is largely consistent with
Vasek’s (1983) classification of the successional status of species on an abandoned
Crossosoma 33(2R), Fall-Winter 2007 53
borrow pit in the Sacramento Mountains in the southeastern Mojave Desert. Three
(Stephanomeria pauciflora, Hymenoclea salsola, and Encelia farinosa) of the five shrub
species on plots within the 1968 corridor in our study were classified as “pioneer
perennials” by Vasek (1983). A fourth species, Ambrosia dumosa, was classified by
Vasek (1983) as a long-lived opportunistic species that can be both an early and late-
successional species. The last species, Bebbia juncea (Benth.) Greene, was not abundant
in Vasek’s (1983) study and was the least abundant of the five shrub colonizers of the
1968 corridor in our study. Bebbia juncea also was uncommon in a study of abandoned
roads in Lake Mead National Recreation Area (Bolling and Walker 2000) and in a study
of abandoned military camps in the eastern Mojave Desert (Prose et al. 1987). This
species does appear capable of colonizing disturbed areas at low densities, however. Also
similar to Vasek’s (1983) classification, Larrea tridentata, categorized as a late-
successional, long-lived perennial, was uncommon even after 38 years on the 1968
corridor.
Our findings on both the 1968 and the 1998 corridors concur with the long time scales
reported in the literature for Mojave Desert plant succession (Lovich and Bainbridge
1999). For example, Webb and Wilshire (1980) found that perennial species composition
on dirt roads abandoned 51 years previously still sharply differed from adjacent control
areas at the Wahmonie ghost town site in the northern Mojave Desert. However, these
early successional shrub communities are not necessarily “bad,” depending on ecological
management objectives. In fact, in our study, plant species richness in early successional
shrub communities on the 1968 corridor was similar to the control, and exotic richness
was actually lower (Fig. 2). Plant assemblages similar to those on this corridor also
characterize natural washes in this region (Wells 1961). Based on minimal colonization
on the untreated 1998 corridor, however, these shrub communities take more than eight
years to develop under the climatic and site conditions characterizing our study. It is
possible that the direct planting of Larrea tridentata seedlings on the treated corridor
bypassed the development of an early successional shrub stage. The planting allowed the
late-succesional L. tridentata to circumvent high-mortality germination and early
seedling phases that make natural regeneration an infrequent event (Barbour 1968).
ACKNOWLEDGEMENTS
We thank Stacey Provencal, Mike Boyles, and Mark Sappington with the National Park
Service for facilitating our research permit for this study; the Southern Nevada Water
Authority for enabling sampling on their right-of-way; David Connally (Southern
Nevada Water Authority) for providing the disturbance history of the 1968 corridor; Utah
State Analytical Laboratories for analyzing soil samples; Sharon Altman (University of
Nevada Las Vegas) for creating Figure 2; and Jill Craig, Jef Jaeger, Denise Knapp, and
three anonymous reviewers for reviewing the manuscript. Funding was provided by the
National Park Service through a cooperative agreement with the University of Nevada
Las Vegas.
54 Crossosoma 33(2R), Fall-Winter 2007
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