\u3cem\u3eBromus tectorum\u3c/em\u3e Litter Alters Photosynthetic Characteristics of Biological Soil Crusts from a Semiarid Shrubland

Invasion by the exotic annual grass Bromus tectorum has increased the cover and connectivity of fine litter in the sagebrush steppes of western North America. This litter tends to cover biological soil crusts, which could affect their metabolism and growth. To investigate this possible phenomenon, biological soil crusts dominated by either the moss Bryum argenteum or the lichen Diploschistes muscorum were covered with B.tectorum litter (litter treatment) or left uncovered (control treatment) and exposed to natural field conditions. After periods of five and ten months, we removed the litter and compared the photosynthetic performance of biological soil crusts from the two treatments. Litter induced photosynthetic changes in our samples. In bothB. argenteum and D. muscorum, biological soil crusts that had been covered with litter for ten months had lower rates of gross photosynthesis and lower chlorophyll content than control samples. Similarly in both biological soil crust types, litter reduced the rate of dark respiration. For D. muscorum, the reduction in dark respiration fully compensated for the decrease in gross photosynthesis, resulting in similar values of net photosynthesis in the two treatments. In contrast, for B. argenteum, net photosynthesis was four-times greater in the control than the litter treatment. Also under litter cover, D. muscorum showed three common adaptations to shade conditions: a decrease in the light compensation point, in the light intensity needed to achieve 95% of maximal net photosynthesis, and in the chlorophyll a/b ratio. None of these changes was apparent in B. argenteum. Overall, our results indicate that photosynthetic responses to the presence of litter varied among species of the crust biota and that the litter can reduce the photosynthetic capacity of biological soil crusts. These results help to explain field observations of decreases in biological soil crust cover and changes in biological soil crust composition with increases in litter cover, and suggest that the landscape-wide invasion by B. tectorum may have substantial effects on biological soil crust performance and therefore their capacity to function in semiarid shrublands

Bromus tectorum litter alters photosynthetic characteristics of biological soil crusts from a semiarid shrubland

a b s t r a c t Invasion by the exotic annual grass Bromus tectorum has increased the cover and connectivity of fine litter in the sagebrush steppes of western North America. This litter tends to cover biological soil crusts, which could affect their metabolism and growth. To investigate this possible phenomenon, biological soil crusts dominated by either the moss Bryum argenteum or the lichen Diploschistes muscorum were covered with B. tectorum litter (litter treatment) or left uncovered (control treatment) and exposed to natural field conditions. After periods of five and ten months, we removed the litter and compared the photosynthetic performance of biological soil crusts from the two treatments. Litter induced photosynthetic changes in our samples. In both B. argenteum and D. muscorum, biological soil crusts that had been covered with litter for ten months had lower rates of gross photosynthesis and lower chlorophyll content than control samples. Similarly in both biological soil crust types, litter reduced the rate of dark respiration. For D. muscorum, the reduction in dark respiration fully compensated for the decrease in gross photosynthesis, resulting in similar values of net photosynthesis in the two treatments. In contrast, for B. argenteum, net photosynthesis was four-times greater in the control than the litter treatment. Also under litter cover, D. muscorum showed three common adaptations to shade conditions: a decrease in the light compensation point, in the light intensity needed to achieve 95% of maximal net photosynthesis, and in the chlorophyll a/b ratio. None of these changes was apparent in B. argenteum. Overall, our results indicate that photosynthetic responses to the presence of litter varied among species of the crust biota and that the litter can reduce the photosynthetic capacity of biological soil crusts. These results help to explain field observations of decreases in biological soil crust cover and changes in biological soil crust composition with increases in litter cover, and suggest that the landscape-wide invasion by B. tectorum may have substantial effects on biological soil crust performance and therefore their capacity to function in semiarid shrublands

Effects of a Companion Plant on the Formation of Mycorrhizal Propagules in \u3cem\u3eArtemisia tridentata\u3c/em\u3e Seedlings

Inoculation of seedlings with arbuscular mycorrhizal fungi (AMF) can increase their establishment after outplanting. The success of this practice depends partly on the extent of root colonization and abundance of AMF propagules in the outplanted seedlings. We conducted a greenhouse experiment to investigate the effects of a companion plant, the native grass Poa secunda J Presl (Sandberg bluegrass), on the formation of spores and vesicles, AMF colonization, and AMF taxa present in the roots of the shrub Artemisia tridentata Nutt (big sagebrush). These effects were tested at two phosphorus (P) fertilization levels, 5 μM and 250 μM. Neither coplanting nor differences in P had an effect on spore density in the potting mix. In contrast, coplanting increased vesicular colonization of A. tridentata from 5% to 18%, but only at low P. Differences in P also affected vesicular colonization of P. secunda, which was 10% and 30% at high and low P, respectively. Arbuscular colonization of A. tridentata was not affected by the treatments and ranged between 12% and 20%. In P. secunda, arbuscular colonization was lower but increased from high to low P. Coplanted seedlings exposed to low P also had the highest levels of total AMF colonization, 70% for A. tridentata and 63% for P. secunda. On the basis of partial sequences of the 28S ribosomal RNA gene, coplanting did not affect the AMF taxa, which were within the Glomeraceae. In some taxa within this family, root fragments containing vesicles are the main propagules. Particularly in this situation, increases in vesicle density caused by coplanting and low P are likely to facilitate mycorrhization of A. tridentata after outplanting, resulting in higher levels of colonization than those naturally occurring in the soil. Such outcomes are critical for assessing the extent to which A. tridentata establishment is limited by insufficient AMF colonization

Herbivory Amplifies Adverse Effects of Drought on Seedling Recruitment in a Keystone Species of Western North American Rangelands

Biotic interactions can affect a plant’s ability to withstand drought. Such an effect may impact the restoration of the imperiled western North American sagebrush steppe, where seedlings are exposed to summer drought. This study investigated the impact of herbivory on seedlings’ drought tolerance for a keystone species in this steppe, the shrub Artemisia tridentata. Herbivory effects were investigated in two field experiments where seedlings were without tree protectors or within plastic or metal-mesh tree protectors. Treatment effects were statistically evaluated on herbivory, survival, leaf water potential, and inflorescence development. Herbivory occurrence was 80% higher in seedlings without protectors. This damage occurred in early spring and was likely caused by ground squirrels. Most plants recovered, but herbivory was associated with higher mortality during the summer when seedlings experienced water potentials between −2.5 and −7 MPa. However, there were no differences in water potential between treatments, suggesting that the browsed plants were less tolerant of the low water potentials experienced. Twenty months after outplanting, the survival of plants without protectors was 40 to 60% lower than those with protectors. The percentage of live plants developing inflorescences was approximately threefold higher in plants with protectors. Overall, spring herbivory amplified susceptibility to drought and delayed reproductive development

Arbuscular Mycorrhizae Alter Photosynthetic Responses to Drought in Seedlings of <i>Artemisia tridentata</i>

The establishment of Artemisia tridentata, a keystone species of the sagebrush steppe, is often limited by summer drought. Symbioses with arbuscular mycorrhizal fungi (AMF) can help plants to cope with drought. We investigated this possible effect on A. tridentata seedlings inoculated with native AMF and exposed to drought in greenhouse and field settings. In greenhouse experiments, AMF colonization increased intrinsic water use efficiency under water stress and delayed the decrease in photosynthesis caused by drought, or this decrease occurred at a lower soil water content. In the field, we evaluated the effect of AMF inoculation on colonization, leaf water potential, survival, and inflorescence development. Inoculation increased AMF colonization, and the seedlings experienced water stress, as evidenced by water potentials between −2 and −4 MPa and reduced stomatal conductance. However, survival remained high, and no differences in water potentials or survival occurred between treatments. Only the percentage of plants with inflorescence was higher in inoculated than non-inoculated seedlings. Overall, the greenhouse results support that AMF colonization enhances drought tolerance in A. tridentata seedlings. Yet, the significance of these results in increasing survival in nature remains to be tested under more severe drought than the plants experienced in our field experiment