The influence of subterranean termites on the hydrological characteristics of a Chihuahuan desert ecosystem

Summary Rainfall simulation at an average intensity of 124 mm·h^-1 was used to compare infiltration and run off on arid areas where subterranean termites had been eliminated four years prior to the initiation of the study (termite free) with adjacent areas populated by subterranean termites (termites present). Infiltration rates on termite free plots with less than 5% perennial plant cover were significantly lower 51.3±6.8 mm·h^-1 than rates on comparable termites present plots 88.4±5.6 mm·h^-1. On plots centered on Larrea tridentata shrubs, there were no differences in infiltration rates with or without termites. Plots with shrub cover had the highest infiltration rates 101±6 mm·h^-1. Highest run-off volumes were recorded from termite free <5% grass cover plots and the lowest from plots with shrubs. There were no differences in suspended sediment concentrations from termites present and termite free plots. Average bed load concentration was more than three times greater from termite free, <5% cover plots than from termites present, <5% cover plots.The reduction in infiltration, high run-off volumes and high bedloads from termite free areas without shrub cover is related to increased soil bulk density resulting from the collapse of subterranean galleries of the termites that provide avenues of bulk flow into the soil. Subterranean termites affect the hydrology of Chihuahuan desert systems by enhancing water infiltration and retention of top soil. The presence of a shrub canopy and litter layer cancels any effect of subterranean termites on hydrological parameters. Since approximately 2/3 of the area is not under shrub canopies, subterranean termites are considered to be essential for the maintenance of the soil water characteristics that support the present vegetation.     

Contributions of subterranean termites to the “economy” of Chihuahuan desert ecosystems

Summary We examined the role of subterranean termites in decomposition of cattle dung, various herbaceous plant species and wood in a Chihuahuan desert ecosystem. From July–September, termites removed dung at a rate of 0.63 g day^-1 accounting for a percent mass loss of 19.5–100%. During the autumn subterranean termites consumed more than 50% of the leaves of the shrub Larrea tridentata, the grass Erioneuron pulchellum and annual plant Lepidium lasiocarpum and Baileya multiradiata but used very little of two other annuals Eriastrum diffusum and Eriogonum trichopes. Yucca inflorescence stalks on plots with termites lost 23% of their original mass in 30 months while those on termite free soils lost 11%. Elimination of termites resulted in reduction of fluff grass, Erioneuron pulchellum biomass, thereby affecting the structure of the ecosystem.     

The contribution of abiotic processes to buried litter decomposition in the northern Chihuahuan desert

Summary Creosobebush (Larrea tridentata) fine litter was treated with either the general biocide HgCl₂ and CuSO₄ or water (controls) and buried 5 cm beneath the soil surface in the northern Chihuahuan Desert. The treated litter showed significantly less mass loss than controls during the three month summer-autumn field study; controls lost about 20% of the original mass while treated litter lost less than 2%. In addition, the total nitrogen content of the control litter increased from an initial concentration of about 14.08 g kg^-1 to 17.62 g kg^-1 dry weight by the end of the study, while treated litter nitrogen content decreased to 13.30 g kg^-1. Results suggest abiotic processes other than leaching have little effect on the decomposition of buried litter in this environment.     

Invading Monotypic Stands of Phalaris arundinacea: A Test of Fire, Herbicide, and Woody and Herbaceous Native Plant Groups

Phalaris arundinacea L. is an aggressive species that can dominate wetlands by producing monotypic stands that suppress native vegetation. In this study invasion windows were created for native species in monotypic stands of P. arundinacea with either fire or herbicide. Three native species groups, herbaceous plants, herbaceous seeds, and woody shrubs, were planted into plots burned or treated with herbicide in the early spring. Fire did not create an effective invasion window for native species; there was no difference in P. arundinacea root and shoot biomass or cover between burned and control plots (p≥ 0.998). Herbicide treatment created an invasion window for native species by reducing P. arundinacea root and shoot biomass for two growing seasons, but that invasion window was fast closing by the end of the second growing season because P. arundinacea shoot biomass had nearly reached the shoot biomass levels in the control plots (p= 0.053). Transplant mortality, frost, and animal herbivory prevented the herbaceous species and woody seedlings from becoming fully established in the plots treated with herbicide during the first year of the experiment. Transplanted monocots had a greater survival than dicots. By the second growing season the herbaceous group had the greatest mean areal cover (5%), compared to the woody seedlings (3%) and seed group (0%). Long‐term monitoring of the plots will determine whether the herbaceous transplants will compete effectively with P. arundinacea and whether the woody species will survive, shade the P. arundinacea, and accelerate forest succession.     

Winter survival of North American grassland birds is driven by weather and grassland condition in the Chihuahuan Desert

Populations of grassland birds that overwinter in the Chihuahuan Desert are declining more rapidly than other grassland birds, and survival during the non‐breeding season may have a strong influence on population trends of these species. Habitat loss and deterioration due to desertification may be contributing to these declines, and the winter ecology of grassland birds under these changing environmental conditions remains relatively unexplored. To fill this information gap, we estimated the survival of two grassland‐obligate sparrows, Baird's Sparrows (Ammodramus bairdii) and Grasshopper Sparrows (A. savannarum), on their wintering grounds in the Chihuahuan Desert, and investigated the role of habitat structure and weather on survival rates. We deployed radio‐transmitters on Baird's (N = 49) and Grasshopper (N = 126) sparrows near Janos, Chihuahua, and tracked birds from November to March during the winters of 2012–2013 and 2013–2014. Causes of mortality included avian predators, mammals, and possibly weather. We estimated an overall weekly winter survival probability of ? = 92.73% (95% CI[s] = 88.63–95.44%) for Baird's Sparrows in 2012–2013. We estimated a weekly winter survival probability of ? = 93.48% (95% CI[s] = 90.29–96.67%) and ? = 98.78% (95% CI[s] = 97.88–99.68%) for Grasshopper Sparrow in 2012–2013 and 2013–2014, respectively. Weekly winter survival was lower with colder daily minimum temperatures for both species and in areas with taller shrubs for Grasshopper Sparrows, with the shrubs potentially increasing predation risk by providing perches for Loggerhead Shrikes (Lanius ludovicianus). Our results highlight the need to maintain healthy grass structure in wintering areas to provide birds with food, protection from predators, and adequate cover from inclement weather. Our results also demonstrate that the presence of shrubs can lower winter survival, and suggest that shrub encroachment into the winter habitat of these sparrows may be an important driver of their population declines. Shrub removal could increase survival of wintering sparrows in the Chihuahuan Desert by reducing availability of perches for avian predators, thus reducing predation risk.     

Assessment of woody species encroachment in the grasslands of Nechisar National Park,Ethiopia

Ecological survey was executed to assess woody species encroachment into the grassland plain of Nechisar National Park (NNP). Forty‐one woody species were recorded. Dichrostachys cinerea Wight & Arn., Acacia mellifera (Vahl) Benth., Acacia nilotica (L) Willd., Acacia senegal (L.) Willd., Acacia seyal Del. and Acacia tortilis (Forssk.) Hayne were among the major encroaching woody species. The majority of the woody species were found to be highly aggregated in their pattern of distribution, while only few species showed some degree of randomness. The mean woody species density was ca. 1995 woody plants ha^?1. Mean cover of woody, grass, unpalatable forbs and total herbaceous species were 31%, 58%, 68% and 121%, respectively. The woody species density and cover, unpalatable forbs and bare land cover were significantly higher in the highly grazed and fire‐suppressed part of the grassland plain. Pearson correlation coefficient matrix indicated that woody species cover and density were negatively correlated with total herbaceous and grass cover. The high woody, unpalatable forbs and bare land cover indicated the progressively increasing perennial grass species diversity deterioration in the grass plain of the Park. Decline in the grassland condition, unless reversed, will jeopardize the biological diversity as well as the aesthetic value of the NNP.     

Response of plant species and life form diversity to variable fire histories and biomass in the jarrah forest of south‐west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light. We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south‐west Australia. Fire frequency was defined as the total number of fires over a 30‐year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency.     

Andropogon gayanus (Gamba Grass) Invasion Increases Fire-mediated Nitrogen Losses in the Tropical Savannas of Northern Australia

Invasive alien grasses can substantially alter fuel loads and fire regimes which could have significant consequences for fire-mediated nutrient losses. The effects of the alien grass Andropogon gayanus Kunth. (Gamba grass) on fire-mediated nutrient losses was evaluated in Australia’s tropical savannas. Losses of macronutrients during fire were determined by comparing the nutrient pools contained in the fine fuel before fire and in the ash after fire. Pre-fire grass nutrient pools were significantly higher in A. gayanus plots than in native grass plots for all nutrients measured (N, P, K, S, Ca, and Mg). Nutrient losses were substantially higher in A. gayanus plots, with 113% higher losses for N, 80% for P, 56% for K, 63 for S, 355% for Ca, and 345% for Mg. However, only losses of N and Mg varied significantly between grass types. A simplified savanna ecosystem nutrient budget estimated that A. gayanus fires led to the net N loss of 20 kg ha⁻¹ y⁻¹. This is a conservative estimate because total fuel loads were relatively low (7.85 t ha⁻¹) for A. gayanus invaded plots leading to a relatively moderate intensity fire (6,408 kW m⁻¹). Higher A. gayanus fuel loads and fire intensities could potentially lead to losses of up to 61.5 kg N ha⁻¹ from the grass fuel. Over the long term, this is likely to lead to depletion of soil nutrients, particularly N, in the already low-fertility tropical savanna soils.     

Effects of restriction of wild herbivore movement on woody and herbaceous vegetation in the Okavango Delta Botswana

We conducted herbaceous and woody vegetation surveys across Botswana's southern Okavango Buffalo Fence, which separates wildlife management from tribal grazing areas, to determine whether the restriction of herbivore movement by fencing has influenced vegetation composition, diversity and structure. We sampled herbaceous and woody vegetation at twenty paired sites every 2 km along the fence. For the herbaceous layer, ten 0.25 m² quadrats were laid every 10 m perpendicular to the fence; while for the woody vegetation, variable quadrat plots were used. Paired t‐tests were run. Results show little difference in forb or grass composition between the two sides. However, the cover and diversity of many woody species were greatly reduced across most height classes on the wildlife management side. Overall woody cover on the wildlife side of the fence was nearly half that of the tribal grazing side (t = 2.83, P = 0.011, df = 19), while overall wood diversity was also significantly less on the wildlife side (t = 3.29, P = 0.004, df = 19). We conclude that the concentration of wildlife due to the fence, while improving habitat for some herbivore species, is having a detrimental effect on plant diversity in general.     

Evaluating nurse plants for restoring native woody species to degraded subtropical woodlands

Harsh habitats dominated by invasive species are difficult to restore. Invasive grasses in arid environments slow succession toward more desired composition, yet grass removal exacerbates high light and temperature, making the use of “nurse plants” an appealing strategy. In this study of degraded subtropical woodlands dominated by alien grasses in Hawai'i, we evaluated whether individuals of two native (Dodonaea viscosa, Leptocophylla tameiameia) and one non‐native (Morella faya) woody species (1) act as natural nodes of recruitment for native woody species and (2) can be used to enhance survivorship of outplanted native woody species. To address these questions, we quantified the presence and persistence of seedlings naturally recruiting beneath adult nurse shrubs and compared survival and growth of experimentally outplanted seedlings of seven native woody species under the nurse species compared to intact and cleared alien‐grass plots. We found that the two native nurse shrubs recruit their own offspring, but do not act as establishment nodes for other species. Morella faya recruited even fewer seedlings than native shrubs. Thus, outplanting will be necessary to increase abundance and diversity of native woody species. Outplant survival was the highest under shrubs compared to away from them with few differences between nurse species. The worst habitat for native seedling survival and growth was within the unmanaged invasive grass matrix. Although the two native nurse species did not differentially affect outplant survival, D. viscosa is the most widespread and easily propagated and is thus more likely to be useful as an initial nurse species. The outplanted species showed variable responses to nurse habitats that we attribute to resource requirements resulting from their typical successional stage and nitrogen fixation capability.     

Does shrub invasion indirectly limit grass establishment via seedling herbivory? A test at grassland‐shrubland ecotones

Question: Does shrub invasion at ecotones indirectly limit grass establishment by increasing mammalian seedling herbivory? Location: Chihuahuan Desert, New Mexico, USA. Methods: We tested the hypothesis that herbivore‐related mortality of seedlings of the dominant perennial grass Bouteloua eriopoda would be highest in shrub‐dominated portions of grassland‐shrubland ecotones. We tested the hypothesis in two Chihuahuan Desert sites featuring similar shrub encroachment patterns but different shrub species, grass cover, and different abundances of small mammals. Within each site we transplanted B. eriopoda seedlings to grass‐dominated, middle, and shrub‐dominated positions of replicate ecotones during the time of year (mid‐summer) when they would naturally appear and monitored seedling fates. We estimated population size/activity of putative small mammal herbivores. Results: Seedlings were killed by mammals in greater numbers in shrubland than in grassland or middle ecotone positions at the site with large herbivore numbers. At the site with low herbivore numbers, most seedlings were killed in middle ecotone positions. The abundance patterns of herbivores did not parallel patterns of seedling herbivory across the ecotones or between sites. Conclusions: Seedling herbivory is an important process and is related to vegetation composition, but the mechanisms underlying the relationship are not clear. We speculate that variation in small mammal foraging behavior may contribute to seedling herbivory patterns. Restoration strategies in the Chihuahuan Desert need to account for the abundance and/or behavior of native herbivores.     

Effects of kangaroo rat exclusion on vegetation structure and plant species diversity in the Chihuahuan Desert

Long-term (1977–90) experimental exclusion of three species of kangaroo rats from study plots in the Chihuahuan Desert resulted in significant increases in abundance of a tall annual grass (Aristida adscensionis) and a perennial bunch grass (Eragrostis lehmanniana). This change in the vegetative cover affected use of these plots by several other rodent species and by foraging birds. The mechanism producing this change probably involves a combination of decreased soil disturbance and reduced predation on large-sized seeds when kangaroo rats are absent. Species diversity of summer annual dicots was greater on plots where kangaroo rats were present, as predicted by keystone predator models. However, it is not clear whether this was caused directly by activities of the kangaroo rats or indirectly as a consequence of the increase in grass cover. No experimental effect on species diversity of winter annual dicots was detected. Our study site was located in a natural transition between desert scrub and grassland, where abiotic conditions and the effects of organisms may be particularly influential in determining the structure and composition of vegetation. Under these conditions kangaroo rats have a dramatic effect on plant cover and species composition.     

Mechanisms for the keystone status of kangaroo rats: graminivory rather than granivory?

Graminivory by kangaroo rats (Dipodomys spp.) was investigated as a potential mechanism for the keystone role of these rodents in the dynamics of desert grasslands. Experiments confirmed that Ord's kangaroo rats (Dipodomys ordii) cut and consumed a large proportion of the tillers of three Chihuahuan Desert tussock-forming grass species. Field observations indicated that the characteristically cut grass tillers were absent from all-rodent and medium-sized kangaroo rat exclosures, but were frequent in large-sized kangaroo rat and rabbit exclosures, indicating that the medium-sized kangaroo rats (D. ordii, D. merriami) were responsible for grass cutting. Tiller waste as a percentage of peak standing crop ranged from 7% in grassland habitats to 0.7% in Flourensia cernua shrubland. Of the 13 species of perennial, tussock-forming grasses measured, only one, Muhlenbergia porteri, had no tillers cut by kangaroo rats. This study demonstrates that the keystone role of kangaroo rats in Chihuahuan Desert grassland ecosystems is probably the result of their graminivory. Received: 28 October 1996 / Accepted: 26 February 1997     

Changes in heathland vegetation under goat grazing: effects of breed and stocking rate

Questions: How are heathland vegetation dynamics affected by different goat grazing management? Location: Cantabrian heathlands in Illano, Asturias, northern Spain. Methods: During 4 years, vegetation dynamics (structural composition, canopy height and floristic diversity) were studied under three goat grazing treatments with three replicates: high stocking rate (11.7 goats ha^?1) with a local Celtiberic breed, and high (15 goats ha^?1) and low (6.7 goats ha^?1) stocking rates with a commercial Cashmere breed. Results: The relative cover of woody plants, particularly heather species, decreased more while herbaceous cover increased more under local Celtiberic than under Cashmere breed grazing. Within Cashmere treatments, the cover and height of live shrubs decreased more and the herbaceous cover increased more under high than under low stocking rate. Redundancy analysis showed a significant effect of treatment × year interaction on floristic composition. Greater species richness was recorded under local goat grazing, but Shannon diversity index fell in the fourth year on these plots because of dominance by two grass species. Conclusions: Local Celtiberic goat grazing at such a high stocking rate (11.7 goats ha^?1) hinders the development of sustainable systems on these heathlands, both in environmental and productive terms, owing to the limitations in soil fertility. Nevertheless, Celtiberic goats could be useful for controlling excessive shrub encroachment and reducing fire hazard. Cashmere goat grazing at high stocking rate promoted the highest Shannon diversity by generating a better balance between woody and herbaceous plants, while shrub dominance was not altered under the low stocking rate.     

Plant diversity patterns in the Aljibe Mountains (S. Spain): a comprehensive account

The Aljibe Mountains are located in the southern tip of the Iberian Peninsula and have a remarkable biogeographical interest. The complete plant species list (trees, climbers, shrubs, perennial and annual herbs, ferns, lichens, bryophytes and macroscopic algae) was recorded in four 0.1 ha plots from each of the most representative community types (Quercus suber woodland, Q. canariensis forest, open heathland and Q. coccifera shrubland). Up to 119 plant species were found in total in the Q. suber woodland plot. The diversity of woody plants was analysed from 44 samples of cover (100 m line), and the herbaceous layer was explored in 200 quadrats (of 0.5 × 0.5 m). Three biodiversity components (species richness, endemism, and taxonomic singularity) were evaluated in both shrub and herbaceous layers. Open heathlands showed the highest richness of endemic species, both woody and herbaceous. The highest number of woody species was found in the evergreen Q. suber woodland, and of herbaceous species in the semi-deciduous Q. canariensis woodland. Taxonomic singularity was higher in Q. canariensis woodlands and Q. coccifera shrublands for woody species, but there were no significant differences in the herbaceous layer. Local species diversity of heathlands in this region resembles that of South African heathlands (fynbos), despite the obvious geographic and floristic distance, and contrasts with the low diversity of biogeographically closer, European temperate heathlands. The Aljibe Mountains show high diversity values for different life forms (from trees to mosses) and spatial scales (from community to region), and are rich in endemic species. Thus, this area should be recognised as a relevant unit within the Mediterranean plant diversity hot spots.     

Tree and grass rooting patterns in an African humid savanna

Abstract. Spatial and temporal soil partitioning between roots of the two savanna plant components, i.e. trees and grasses, were investigated in a West African humid savanna. Vertical root phytomass distribution was described for grass roots, large (> 2 mm) and fine (< 2 mm) tree roots, in open sites and beneath tree canopies. These profiles were established monthly over one year of vegetation growth. Natural ¹³C abundance measurement was used to determine the woody/herbaceous phytomass ratio in root samples. Tree and grass root distributions widely overlapped and both were mostly located in the top 20 cm of the soil. Grass root phytomass decreased with depth whereas woody root phytomass peaked at about 10 cm depth. No time partitioning was detected. These structural results do not support the hypothesis of soil resource partitioning between trees and grasses and are thus consistent with functional results previously reported.     

Termite- and Mulch-Mediated Rehabilitation of Vegetation on Crusted Soil in West Africa

The rehabilitation of vegetation on structurally crusted soils by triggering termite activity through mulch was studied on three soil types in northern Burkina Faso, West Africa. A split-plot design was used in a fenced environment for the experiment. Insecticide (Dieldrin) was used at a rate of 500 g a.i. (active ingredient)/ha to create nontermite and termite plots. Three mulch types consisting of straw (Pennisetum pedicellatum), woody material (Pterocarpus lucens), and a composite mulch (straw and woody material) applied at a rate of 3, 6, and 4 tons/ha, respectively, were used to trigger termite activity. The grasses and woody species on the plots were surveyed. Nontermite plots responded weakly to mulch treatments, but even in the first year vegetation established on termite + mulch plots. Termite activity resulted in the increase of plant cover, plant species number, phytomass production, and rainfall use efficiency. Infiltrated water use efficiency and plant diversity were not statistically different among treatments during the first 2 years but were in the third. Woody species established only on termite plots. The three types of mulch plots showed greater vegetation development than bare plots, which remained bare throughout the experiment. Analysis of the termite and mulch interaction indicated that mulch plots without termites did not perform better than bare plots, especially in the case of woody plant regeneration. Vegetation rehabilitation was best with composite and straw mulches with termites, followed by woody mulch with termites; it was worst on bare plots.     

Changing dominance of key plant species across a Mediterranean climate region: implications for fuel types and future fire regimes

Herbaceous and woody plants represent different fuel types in flammable ecosystems, due to contrasting patterns of growth and flammability in response to productivity (moisture availability). However, other factors, such as soil type, fire regimes and competitive interactions may also influence the relative composition of herbaceous and woody plants within a community. The Mediterranean climate region of south eastern Australia is transitional between two contrasting fuel systems; herbaceous dominated in the dry north, versus woody plant dominated shrublands in the relatively moist south. Across the rainfall gradient of the region, there are confounded changes in dominant soil types and fire frequency. We used model-subset selection using Akaike’s Information Criterion to examine potential driving mechanisms of community compositional change from herbaceous (e.g. Triodia scariosa, Austrostipa sp.) to woody plants (e.g. Beyeria opaca, Leptospermum coriaceum, Acacia ligulata) by measuring relative cover across combinations of rainfall, time since the last fire (TSF) and soil type. We examined the relative influence of environmental versus competitive interactions on determining the cover of perennial hummock grass, T. scariosa, and co-occurring woody shrubs. Rainfall and soil types, rather than competition, were the over-arching determinants of the relative cover of grasses and shrubs. Given the sensitivity to rainfall, our results indicate there is strong potential for the nature of fuel, flammability and fire regimes to be altered in the future via climate change in this region.     

Cattle and Weedy Shrubs as Restoration Tools of Tropical Montane Rainforest

Over the last 150 years, a large proportion of forests in Latin America have been converted to pastures. When these pastures are abandoned, grasses may slow re‐establishment of woody species and limit forest regeneration. In this study, we explored the use of cattle in facilitating the establishment of woody vegetation in Colombian montane pastures, dominated by the African grasses Pennisetum clandestinum (Kikuyo) and Melinis minutiflora (Yaraguá). First, we described woody and herbaceous vegetation in grazed and non‐grazed pastures. Second, we tested the effect of grazing and seed addition on the establishment and growth of woody species. We also determined if the effect of grazing was different in P. clandestinum and M. minutiflora pastures. We found that low stocking density of cattle greatly increased density, number of branches per individual (a measure of “shrubiness”), and basal area of woody species, but also reduced woody plant species richness and diversity. In the grazed area, the shrubs Baccharis latifolia (Chilca) and Salvia sp. (Salvia) were the most abundant. The combined effect of grazing and shading from the shrubs reduced herbaceous vegetation by 52 to 92%. In the grazing/seed addition experiment, grazing increased establishment of woody seedlings, particularly of the shrub Verbesina arborea (camargo), but the largest effect was seed addition. Where grasses are an important barrier to regeneration, grazing can facilitate the establishment of shrubs that create a microhabitat more suitable for the establishment of montane forest tree species.