Spatial Arrangement and Potential Detrimental Effects of Nassella tenuissima on an Urban Remnant Shortgrass Prairie in West Texas
Juan G. García-Cancel1,2,* and Robert D. Cox2
1Department of Environmental Science, University of Puerto Rico, Río Piedras, Puerto Rico, U.S.A. 00925-2537 00939 (ORCID 0009-0007-8033-8931) 2Department of Natural Resources Management, Davis Agricultural College, Texas Tech University, Lubbock, Texas, U.S.A. 79409 *Corresponding author.
Urban Naturalist, No. 80 (2025)
Abstract
We investigated the spatial distribution of Nassella tenuissima, an invasive grass, in an urban native remnant shortgrass prairie, as well as its effects on a native grass species, Bouteloua gracilis, in a glasshouse setting. We first mapped populations in a native prairie remnant to determine spatial distributions of adult N. tenuissima clumps, and detected a high level of aggregation in the field. We then set up an additive competition experiment to determine at which density N. tenuissima influences growth and nitrogen allocation of native grass seedlings. Higher N. tenuissima density had a detrimental effect on growth for B. gracilis seedlings, while B. gracilis had higher biomass and nitrogen accumulation with no N. tenuissima present. Our results suggest that the effects of invasive grasses can start with very low numbers in early life stages in the field. Management and control of these should be done sooner rather than later when density increases with more propagule rain. In urban settings, N. tenuissima may rapidly invade native prairie remnants, and could significantly alter biodiversity and ecosystem attributes.
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Urban Naturalist
Spatial Arrangement and Potential Detrimental Effects of
Nassella tenuissima on an Urban Remnant Shortgrass Prairie
in West Texas
Juan G. García-Cancel1,2* and Robert D. Cox2
Abstract: We investigated the spatial distribution of Nassella tenuissima, an invasive grass, in an
urban native remnant shortgrass prairie, as well as its effects on a native grass species, Bouteloua
gracilis, in a glasshouse setting. We first mapped populations in a native prairie remnant to determine
spatial distributions of adult N. tenuissima clumps, and detected a high level of aggregation in the
field. We then set up an additive competition experiment to determine at which density N. tenuissima
influences growth and nitrogen allocation of native grass seedlings. Higher N. tenuissima density had
a detrimental effect on growth for B. gracilis seedlings, while B. gracilis had higher biomass and
nitrogen accumulation with no N. tenuissima present. Our results suggest that the effects of invasive
grasses can start with very low numbers in early life stages in the field. Management and control of
these should be done sooner rather than later when density increases with more propagule rain. In
urban settings, N. tenuissima may rapidly invade native prairie remnants, and could significantly alter
biodiversity and ecosystem attributes.
Introduction
Terrestrial ecosystems around the planet are experiencing shifts in species assemblages
due to human activity, including both introductions of non-native species (DiTomaso et
al. 2017, Kreyling et al. 2011, Lockwood et al. 2007) and anthropogenic climate change
(Vitousek et al. 1997). Species that are transported by humans and are able to survive and
reproduce in their new systems could potentially become invasive if their environmental
impacts are on a large scale and affect human economic or cultural activities (Lockwood et
al. 2008). Plants in particular can be inconspicuous when initially introduced and only once
they have increased does their effect on the ecosystem become noticeable (Crooks 2005),
and perhaps nearly irreversible (Evans et al., 2017).
Invasive species can affect native species through multiple pathways; including through
soil nutrient dynamics, competition, shading effects, or by direct competition for soil resources
due to their extensive root system (García-Cancel and Thaxton 2018, Ibarra-Flores
et al. 1999, Parkinson et al. 2013, Rojas-Sandoval and Meléndez-Ackerman 2012). Some
invasive plants can alter the dynamics in ecosystems by exerting indirect competition pressures
with native plants by deterring herbivory on their tissues, forcing herbivores to eat
more palatable plants (Mapaura et al. 2020).
The spatial arrangement of plants in a landscape can also accentuate the effects they
might have on other local plants by monopolizing resources (Archer et al. 2017), or by
providing valuable aid at key life stages (García-Cancel and Thaxton 2018, Padilla and
Pugnaire 2006). Therefore, spatial arrangement of plant populations can show past his-
1Department of Environmental Science, University of Puerto Rico, Río Piedras, Puerto Rico, U. S. A.
00925-2537 00939 (ORCID 0009-0007-8033-8931) 2Department of Natural Resources Management,
Davis Agricultural College, Texas Tech University, Lubbock, Texas, U. S. A.79409 *Corresponding
author: j.garcia.cancel@gmail.com
Associate Editor: Sonja Knapp, Helmholtz Centre for Environment al Research–UFZ
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torical trends in propagule arrival and spread (Seabloom et al. 2006). Even geographically
restricted native plants could be released from their constraints and become invasive in
introduced regions if the biological barriers such as mountains, oceans or deserts are
bypassed (Seabloom et al. 2006, Vilà and D’Antonio 1998). The same can happen when
disturbance regimes have been altered and have weakened the native species pool, making
them susceptible to invasion (D’Antonio and Vitousek 1992, Suding et al. 2004), including
in grasslands (Seabloom et al. 2013).
Dispersal of non-native plants from urban centers is a common occurrence (Beaury
et al. 2021, Veldman and Putz 2010) that has accelerated the rate of biodiversity loss
and breakdown of ecosystem services globally (Suding et al. 2004). Some have argued
that urban green spaces can act as ecosystem service providers in lieu of native robust
ecosystems by providing services e.g. nectar providers (Yessoufou 2023); or sources of
adaptability to changed local soil conditions (Honfi et al. 2023). Others have found that
urban green places can act as reservoirs and sources of ornamental, potentially invasive
plant species capable of spreading into nearby native areas (Knapp et al. 2012, Reichard
and White 2001, Shah et al. 2025).
Nassella tenuissima (Trin.) Barkworth (Mexican feathergrass) is an invasive grass in
the Southern High Plains of the continental United States, but native to the arid Trans
Pecos region of Texas, northern Mexico and southeastern New Mexico, with some populations
in southern South America (Humphries and Florentine 2021, Jacobs et al. 1998). It
has been introduced in several locations as an ornamental plant (Fig. 1), and it has been
observed to remain in the seedbank up to 7 years or longer after removal of adult plants
(Moran et al. 2018). In environments with more water availability, such as in urban land-
Figure 1. Native locations of N. tenuissima in TX (blue counties) and known locations (green circles)
of N. tenuissima in the state of Texas as of April 21, 2021. (QGIS geographic information system).
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scaping, this grass can spread rapidly to nearby ecosystems due to its high seed output
(Humphries and Florentine 2021, Moran et al. 2018, Russell and Rector 2016).
Bouteloua gracilis (Kunth) Lag. ex Griffiths (Blue grama), (syn. Chondrosum gracile)
is a C4 warm-season perennial caespitose grass located mainly in the North American shortgrass
prairies (Anderson 2003, Wynia 2007). The species has a wide range in the North
American Central Plains, from south Canadian prairies (Wilson and Pärtel 2003) to northern
Mexico (Anderson 2003). The broader genus has a Pan-American distribution, with 57
described species (Siquieros-Delgado 2007).
B. gracilis is a fast-reproducing grass, able to reproduce asexually by stolons and
sexually by outcrossing wind-pollination and wind dispersed seeds (Anderson 2003). Its
foliage has a high nutrient content which is favored by livestock (Melgoza-Castillo et al.
2014), and it is readily used in restoration efforts due to easy germination rates (Bakker et
al. 2003). B. gracilis has the ability to suppress some invasive grass species with its high
germination rates occupying available space (Wilson and Pärtel 2003). On that regard it
has also been documented to have a strong ability to extract soil moisture by the profuse
root system of adult plants (Dormaar et al. 1994), though in some studies B. gracilis did
not rapidly spread in highly disturbed sites with the presence of other competitive perennials
(Samuel and Hart 1994).
The early effects of N. tenuissima invasion in a remnant shortgrass prairie embedded in
an urban matrix have not been studied. To study the spatial patterns that such an invasive
might have, we carried out a field study to determine the presence and spatial patterns of
N. tenuissima plants in the remnant rangeland in an urban matrix. We hypothesized that N.
tenuissima clumps will be clustered instead of being randomly distributed and that their
densities will be higher than randomly expected. We also conducted a greenhouse competition
experiment with N. tenuissima and the native shortgrass prairie grass Bouteloua
gracilis. We hypothesized that increasing density of N. tenuissima would impact biomass
production and nitrogen tissue allocation in B. gracilis individuals.
Methods
Experimental design: Field study
We studied the spatial distribution of N. tenuissima in a 160-acre remnant shortgrass
prairie (33˚ 36’9.36” N, -101˚ 54’ 2.52” W) managed by the Department of Natural Resources
Management, Texas Tech University, in Lubbock, TX (Fig. 1). The presence of N. tenuissima
had been observed as early as 2014. We established 25 parallel transects, measuring
around 700 m long and 3 m wide, 30 m apart. A positive sighing of a Mexican feathergrass
clump would be recorded by a geotagged photograph along these transects during the days
of May 15–16, 2019 and June 24–25, 2019. We then used Point Density Analysis as part of
the Density Analysis toolbox from ArcGIS (Esri©) to produce Average Nearest Neighbor
between grass clumps to determine if there was a clustering of the plants and a High/Low
Clustering (Getis-Ord General G) to determine the likelihood that the cluster pattern could
be the result of random chance (Getis and Ord 1992, Mitchell 20 05).
Experimental design: Glasshouse
We carried out addition competition experiments in the TTU glasshouses in Lubbock, TX,
USA (33° 35’ 3.066” N, 101° 53’ 13.1199” W) with seedlings of N. tenuissima and B. gracilis.
We used seeds collected from adult N. tenuissima from the remnant shortgrass prairie administered
by the Department of Natural Resources Management, Texas Tech University, in the
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summer of 2021. Seeds were rinsed with a 10 % chlorine bleach solution to limit the occurrence
of phytopathogenic fungi and bacteria and seeded for the experiment in Spring 2022. For the
native component of this experiment, we chose B. gracilis due to its widespread distribution and
importance for shortgrass prairie ecosystems in North America (Melgoza-Castillo et al. 2014),
in addition to its easy availability from commercial vendors. To that end, B. gracilis seeds for
this study were purchased from Bamert Seed Company, Muleshoe, TX.
We designed an experimental additive design, with the target species being B. gracilis and
the subsequent addition of N. tenuissima in increasing numbers. Seeds from both species were
germinated in sterile greenhouse soil media (BM6 All-Purpose Optimal Porosity Mix, Incubated
pH = 5.4–6.2 Berger ©) in Spring 2022 in controlled greenhouse conditions at 25° C. Once germinated,
seedlings were randomly assigned into five replicates of six treatments ranging from
0–5 N. tenuissima per pot. Each pot was a 2-gallon (7.5 L) nursery pot (19D x 24.63W x 20.63H
cm) and had a single B. gracilis individual and 0–5 individuals of N. tenuissima.
All pots were watered to saturation (on average 511.6 mL per 5 seconds) once a week and allowed
to dry between waterings. Gibson et al. (1999) warned about the use of additive designs in
greenhouse conditions, as researchers focus on harvest end data and not other factors that might
be at play in the plant dynamics; such as the interactions of shoot and roots and their relation to
plant species performance. We mitigated this by collecting data on available soil moisture using
gravimetric soil moisture measurements, and biomass growth output at the time of harvest, with
distinction for above and below ground organs and nutrient allocation in selected experiments.
Biomass samples were weighed after harvest before being oven dried, and weighed a second
time to estimate dry weights of the plant matter. All biomass measurements were analyzed using
one-way ANOVA using the program Infostat© (DiRienzo et al. 2008). Fixed factors were species
identity and invasive plant density as categorical data, while the dependent variables were
wet and dry biomass production and above and below ground maximum length of the native
seedlings. Biomass was weighed with a Mettler Toledo AL104 Balance and a standard measuring
tape was used for measuring maximum length above ground as well below ground.
After harvest, plant matter was oven dried at 55.5 °C (~140 °F) for a period of 3 weeks
and ground to 2 mm mesh size using a Wiley Mill for larger samples and smaller (< 0.5
mm) mesh size for smaller samples. For the tissue allocation of nitrogen under the treatments,
we selected plants from four treatments, these being Treatment #0, #1, #3, and
#5 to determine if increasing density of invasive grass also affected nitrogen deposition
in above and below ground tissues. If ground samples were too large, they were further
processed in the laboratory by freezing with liquid nitrogen and grinding with mortar and
pestle before being rolled in tin foil and analyzed in an elemental analyzer (Costech-4010;
Costech Analytical, Valencia, CA, USA). Spectral graphs from the combusted samples
were produced, as well as data from the available percent nitrogen found in the analyzed
samples. These data were analyzed by one-way ANOVA using the program Infostat©
(Di Rienzo et al. 201 1). The fixed factor was invasive plant density , while the dependent
variables were biomass production of target native seedlings and nitrogen percentages in
tissue samples. Ad hoc differences were analyzed using Tukey test.
Results
Field study
We found that N. tenuissima occupies large tracts of the 160-acre remnant prairie (Fig.
2), with a nearest neighbor ratio of 0.16 clump/m2 (z-score = -51.6811, p-value = 0.00). Expected
mean distance was 211.48 m, but observed mean distance was 34.49 m. The High/
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Figure 2. Point Density Map of known N. tenuissima clumps in the TTU Rangeland in June 2019.
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Low Clustering (Geris-Ord General G) analysis showed a High-cluster spatial pattern (Observed
General G = 0.02, z-score = 10.40, p-value < 0.01; Fig 2).
Greenhouse
Increased N. tenuissima plant density reduced biomass production for B. gracilis roots,
but not for shoot biomass (Table 1). Soil water content was not statistically different between
treatments (x̄ = 1.33, SE = 0.46, F = 1.28, df = 5, p = 0.31), and the presence of this
invasive grass did not result in any difference in organ length. However, root biomass was
significantly different between treatments (Table 1). Treatment #0 had significantly more
root biomass (Fig. 3) compared to any other treatment. The presence of N. tenuissima did
not affect B. gracilis shoot biomass (Table 1). Nitrogen content was statistically different
between treatments for below ground samples, with highest nitrogen content in Treatment
#0, but not for aboveground samples (Table 2).
Discussion
Spatial arrangement of the N. tenuissima clumps was highly clustered, implying that
dispersal is limited, but the patches they colonize have a pronounced thatch formation.
Mexican feathergrass thatch is problematic, being highly fibrous and resistant to degradation
(Moretto et al. 2001), which could impose a limitation to native seed germination
or emergence due the physical barrier imposed by the new thatch layer. Such barriers
have been noted in other study systems (Evans et al. 2017). Future studies could well
explore if the same barriers are present in the TTU Rangeland in Lubbock, TX.
N. tenuissima has been observed to easily expand once escaped from cultivation
(Moran et al. 2018) and create dense patches (Fig. 2), which we now confirm has some
effects in native plant growth and its nutritional potential of native plants for livestock
fodder. It has been widely planted and exported in the ornamental industry due to its
large range of drought tolerance and aesthetic value (Beaury et al. 2021). One of its
impacts on native communities in California, New Zealand, and Australia include unpalatability
for native herbivores, overgrazing more palatable grasses, and diminishing
productivity of rangelands where present (Humphries and Florentine 2021, Moran et al.
2018, Russell and Rector 2016). Even in regions with precipitation as low as 300 mm
Table 1. ANOVA Biomass Analysis for the effects of N. tenuissima density on B. gracilis growth in
the TTU Greenhouse. Values in bold are statistically significant. p-value = 0.05.
N Organ System Variable F df p-value
30 Shoot Length 1.12 5 0.3741
Wet net biomass 0.97 5 0.4577
Dry net biomass 1.18 5 0.3458
30 Root Length 0.79 5 0.5679
Wet net biomass 2.9 5 0.0347
Dry net biomass 2.97 5 0.0315
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Figure 3. Post hoc Tukey Test for B. gracilis dry root biomass under increasing N. tenuissima density
treatments for the summer 2022 growing season. p-value = 0.005.
Table 2. ANOVA analyses for the estimated organ nitrogen content in B. gracilis (nitrogen (g) per
biomass) under increasing N. tenuissima density treatments for summer 2022 growing season. Values
in bold are statistically significant. p-value = 0.05. Degrees of freedom = 3.
Organ System F p-value Treatment Means N Standard Error Comparison
0 0.05 5 0.01 A
Shoot 1.69 0.2085 1 0.04 5 0.01 A
3 0.04 5 0.01 A
5 0.02 5 0.01 A
0 0.20 5 0.02 A
Root 9.65 0.0007 1 0.13 5 0.02 A B
3 0.05 5 0.02 B
5 0.04 5 0.02 B
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can grow N. tenuissima plants, making the central prairies of North America a suitable
region for expansion.
In our experiment the presence of N. tenuissima did not cause remarkable differences
in B. gracilis organ length, but it did reduce root biomass (Table 1). Treatment #0 had
significantly more biomass for roots compared to the other treatments. This might have
influenced the values for gravimetric water content, as the dense and profuse root systems
of B. gracilis (Dormaar et al. 1994, Wynia 2007) might absorb water in comparable
similar rates to the N. tenuissima roots. Nitrogen content in tissues for summer 2022 was
higher for roots (Table 2) compared with shoots. However, some of the B. gracilis plants
in Treatments #1 and #3 had already reached maturity and produced seeds, and may have
begun withdrawing nutrients from the aboveground biomass.
Moretto et al. (2001) found that N. tenuissima and other unpalatable grasses in Argentina
immobilize N in their litter. Nitrogen immobilization would impact nutrient dynamics
in ecosystems where this grass has been introduced. For other Nassella species introduced
in South Africa, the effects of these grasses have been noted to change herbivore behavior
by forcing them to select more palatable vegetation, allowing the Nassella plants to
expand with little competition (Mapaura et al. 2020). Not just wildlife is impacted, but
also livestock as they cannot digest these grasses and when forced it can be harmful to the
digestive tract of the herbivore (Campbell 1998, Mapaura et al. 2020).
Another serious issue that can arise from these grasses is the high deposition of leaflitter
in yards and public spaces that could potentially create large fuel loads for fires (Bell
et al. 2009, Fusco et al. 2019, Fusco et al. 2021). Other studies have mentioned the pivotal
role fire has in grassland ecosystems (D’Antonio and Vitousek 1992, Neary et al. 1999,
Williams and Baruch 2000), yet the alteration in disturbance regimes or secession of these
altogether by land conversion can put human populations at risk (D’Antonio and Vitousek
1992, Fusco et al. 2021). Fire risk has risen in the past few decades (Setterfield et al. 2013,
Fusco et al. 2019, Fusco et al. 2021). Range expansion of non-native grasses complicates
this scenario (Di Tomasso et al. 2017) as some are present in urban settings (Fusco et al.
2021, García-Cancel 2023).
The introduction of Nassella tenuissima into new habitats via the ornamental market
has allowed it to spread into nearby areas due primarily to high seed production
(Humphries and Florentine 2021, Moran et al. 2018, Russell and Rector 2016). However,
native areas invaded by this species might experience the detrimental effects of
increased thatch, potential changes to fire cycles, and changes in nutrient cycling and
native species biomass production. Urban landscapers should minimize the use of this
grass due to the potential for harm to ecosystem services and interactions of native
communities. More imminent to human safety is the potential of this grass to be a fire
hazard, as it produces copious amount of leaf litter where present. This could provide an
economic sink as mitigation of fire hazards and changing climatic patterns favor drying
conditions in the study area.
Acknowledgments
The authors would like to acknowledge Dr. Vikram Baliga for administrative support in greenhouse
facilities in the Texas Tech University, Lubbock, TX. We also would like to thank Dr. Nicholas
G. Smith and Dr. Evan Perkowksi for the laboratory space and guidance with the Costech-4010 Elemental
Analyzer. No specific grant from any funding agency, commercial or not-for-profit sectors
was received for this research.
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