The symmetry of competitive interactions in mixed Norway spruce,silver fir and European beech forests

Questions

We aim for a better understanding of the different modes of intra‐ and inter‐specific competition in two‐ and three‐species mixed‐forests. How can the effect of different modes of competitive interactions be detected and integrated into individual tree growth models? Are species interactions in spruce–fir–beech forests more associated with size‐symmetric or size‐asymmetric competition? Do competitive interactions between two of these species change from two‐ to three‐species mixtures?

Location

Temperate mixed‐species forests in Central Europe (Switzerland).

Methods

We used data from the Swiss National Forest Inventory to fit basal area increment models at the individual tree level, including the effect of ecological site conditions and indices of size‐symmetric and size‐asymmetric competition. Interaction terms between species‐specific competition indices were used to disentangle significant differences in species interactions from two‐ to three‐species mixtures.

Results

The growth of spruce and fir was positively affected by increasing proportions of the other species in spruce–fir mixtures, but negative effects were detected with increasing presence of beech. We found that competitive interactions for spruce and fir were more related to size‐symmetric competition, indicating that species interactions might be more associated with competition for below‐ground resources. Under constant amounts of stand basal area, the growth of beech clearly benefited from the increasing admixture of spruce and fir. For this species, patterns of size‐symmetric and size‐asymmetric competitive interactions were similar, indicating that beech is a strong self‐competitor for both above‐ground and below‐ground resources. Only for silver fir and beech, we found significant changes in species interactions from two‐ to three‐species mixtures, but these were not as prominent as the effects due to differences between intra‐ and inter‐specific competition.

Conclusions

Species interactions in spruce–fir–beech, or other mixed forests, can be characterized depending on the mode of competition, allowing interpretations of whether they occur mainly above or below ground level. Our outcomes illustrate that species‐specific competition indices can be integrated in individual tree growth functions to express the different modes of competition between species, and highlight the importance of considering the symmetry of competition alongside competitive interactions in models aimed at depicting growth in mixed‐species forests.

     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Size‐balanced community reorganization in response to nutrients and warming

It is widely accepted that global warming will adversely affect ecological communities. As ecosystems are simultaneously exposed to other anthropogenic influences, it is important to address the effects of climate change in the context of many stressors. Nutrient enrichment might offset some of the energy demands that warming can exert on organisms by stimulating growth at the base of the food web. It is important to know whether indirect effects of warming will be as ecologically significant as direct physiological effects. Declining body size is increasingly viewed as a universal response to warming, with the potential to alter trophic interactions. To address these issues, we used an outdoor array of marine mesocosms to examine the impacts of warming, nutrient enrichment and altered top‐predator body size on a community comprised of the predator (shore crab Carcinus maenas), various grazing detritivores (amphipods) and algal resources. Warming increased mortality rates of crabs, but had no effect on their moulting rates. Nutrient enrichment and warming had near diametrically opposed effects on the assemblage, confirming that the ecological effects of these two stressors can cancel each other out. This suggests that nutrient‐enriched systems might act as an energy refuge to populations of species under metabolic constraints due to warming. While there was a strong difference in assemblages between mesocosms containing crabs compared to mesocosms without crabs, decreasing crab size had no detectable effect on the amphipod or algal assemblages. This suggests that in allometrically balanced communities, the expected long‐term effect of warming (declining body size) is not of similar ecological consequence to the direct physiological effects of warming, at least not over the six week duration of the experiment described here. More research is needed to determine the long‐term effects of declining body size on the bioenergetic balance of natural communities.     

Foraging behavior of migrant warblers in mixed‐species flocks in Venezuelan shade coffee: interspecific differences,tree species selection,and effects of drought

Shade coffee has been identified as an important habitat for Nearctic‐Neotropical migrants during the non‐breeding season, including species of conservation concern such as Cerulean Warblers (Setophaga cerulea). To better understand habitat features important for migrants in shade coffee, we studied the foraging behavior of migrants in mixed‐species flocks at six shade‐coffee farms in the Cordillera de Merida, Venezuela, in 2008–2009 and the El Niño drought year of 2009–2010. We examined interspecific differences in foraging behavior and tree species selection of three foliage‐gleaning migrants, Blackburnian (Setophaga fusca), Cerulean, and Tennessee (Oreothlypis peregrina) warblers, and aerial‐foraging American Redstarts (Setophaga ruticilla). For morphologically similar Blackburnian and Cerulean warblers, we also examined factors influencing foraging rates (attack and movement rates), capture of large prey, and maneuver/substrate type. We found that aerial‐foraging American Redstarts foraged lower, used more aerial maneuvers, showed no tree species selection, and were less likely to forage in flocks than foliage‐gleaners. Although foraging rates were similar for Blackburnian and Cerulean warblers, the three foliage‐gleaners differed in foraging height and use of maneuvers. Cerulean Warblers foraged lower than the other two species, whereas Blackburnian Warblers used the greatest proportion of woody gleans. All three foliage‐gleaners selected Inga spp. (a commonly planted shade tree in shade‐coffee farms) for foraging, and Blackburnian and Cerulean warblers captured a greater proportion of large prey in Inga spp. than in other tree species. During the drought year, Blackburnian and Cerulean warblers captured half as many large prey and used a greater proportion of woody‐gleans. We found that interactions among behavioral, floristic, and environmental drivers influenced the foraging behavior of migrants wintering in shade coffee. Our results support those of previous studies suggesting that migrants partition resources behaviorally during the non‐breeding season, that foliage‐gleaners may benefit from the presence of shade trees, especially Inga spp., in agroforestry systems, and that drought may influence the foraging behavior of foliage‐gleaning migrants, presumably due to reduced prey availability.     

西藏东部主要建群树种径向生长对极端干旱的响应差异

不同坡向、不同海拔树木生长对极端干旱事件的响应可能不同,然而这方面的认识不足。为此,选取西藏东部珠角拉山阴、阳坡的建群树种川西云杉 (Picea likiangensis var. rubescens) 和大果圆柏 (Juniperus tibetica),在不同海拔高度建立了树轮宽度年表,分析了径向生长的气候响应,以及对极端干旱事件的抵抗力和恢复力。结果表明:阳坡大果圆柏和阴坡川西云杉的树木生长对气候的响应存在相似性,均与前一年3-6月、11-12月气温显著正相关,与当年4-5月气温显著负相关,与当年4-5月降水和帕尔默干旱指数 (PDSI, Palmer Drought Severity Index) 显著正相关 (P <0.05)。阳坡大果圆柏的抵抗力显著低于阴坡川西云杉。随着海拔升高,阴坡川西云杉树木个体的抵抗力和恢复力均显著提高,而阳坡大果圆柏树木个体抵抗力、恢复力在不同海拔无显著区别。结合混合效应模型表明树木抵抗力主要受当年4-5月平均最高气温限制,树木恢复力主要受干旱事件后四年4-5月平均最高气温限制 (P <0.01),说明生长季高温引起的极端干旱是树木径向生长下降的主要原因。     

Drought tolerance and growth in populations of a wide‐ranging tree species indicate climate change risks for the boreal north

Choosing drought‐tolerant planting stock in reforestation programs may help adapt forests to climate change. To inform such reforestation strategies, we test lodgepole pine (Pinus contorta Doug. ex Loud. var latifolia Englm.) population response to drought and infer potential benefits of a northward transfer of seeds from drier, southern environments. The objective is addressed by combining dendroecological growth analysis with long‐term genetic field trials. Over 500 trees originating from 23 populations across western North America were destructively sampled in three experimental sites in southern British Columbia, representing a climate warming scenario. Growth after 32 years from provenances transferred southward or northward over long distances was significantly lower than growth of local populations. All populations were affected by a severe natural drought event in 2002. The provenances from the most southern locations showed the highest drought tolerance but low productivity. Local provenances were productive and drought tolerant. Provenances from the boreal north showed lower productivity and less drought tolerance on southern test sites than all other sources, implying that maladaptation to drought may prevent boreal populations from taking full advantage of more favorable growing conditions under projected climate change.     

Seed germination responses to seasonal temperature and drought stress are species‐specific but not related to seed size in a desert steppe: Implications for effect of climate change on community structure

Investigating how seed germination of multiple species in an ecosystem responds to environmental conditions is crucial for understanding the mechanisms for community structure and biodiversity maintenance. However, knowledge of seed germination response of species to environmental conditions is still scarce at the community level. We hypothesized that responses of seed germination to environmental conditions differ among species at the community level, and that germination response is not correlated with seed size. To test this hypothesis, we determined the response of seed germination of 20 common species in the Siziwang Desert Steppe, China, to seasonal temperature regimes (representing April, May, June, and July) and drought stress (0, ?0.003, ?0.027, ?0.155, and ?0.87 MPa). Seed germination percentage increased with increasing temperature regime, but Allium ramosum, Allium tenuissimum, Artemisia annua, Artemisia mongolica, Artemisia scoparia, Artemisia sieversiana, Bassia dasyphylla, Kochia prastrata, and Neopallasia pectinata germinated to >60% in the lowest temperature regime (April). Germination decreased with increasing water stress, but Allium ramosum, Artemisia annua, Artemisia scoparia, Bassia dasyphylla, Heteropappus altaicus, Kochia prastrata, Neopallasia pectinata, and Potentilla tanacetifolia germinated to near 60% at ?0.87 MPa. Among these eight species, germination of six was tolerant to both temperature and water stress. Mean germination percentage in the four temperature regimes and the five water potentials was not significantly correlated with seed mass or seed area, which were highly correlated. Our results suggest that the species‐specific germination responses to environmental conditions are important in structuring the desert steppe community and have implications for predicting community structure under climate change. Thus, the predicted warmer and dryer climate will favor germination of drought‐tolerant species, resulting in altered proportions of germinants of different species and subsequently change in community composition of the desert steppe.     

Implications of the mesophyll conductance to CO2 for photosynthesis and water‐use efficiency during long‐term water stress and recovery in two contrasting Eucalyptus species

Water stress (WS) slows growth and photosynthesis (A_n), but most knowledge comes from short‐time studies that do not account for longer term acclimation processes that are especially relevant in tree species. Using two Eucalyptus species that contrast in drought tolerance, we induced moderate and severe water deficits by withholding water until stomatal conductance (g_sw) decreased to two pre‐defined values for 24 d, WS was maintained at the target g_sw for 29 d and then plants were re‐watered. Additionally, we developed new equations to simulate the effect on mesophyll conductance (g_m) of accounting for the resistance to refixation of CO₂. The diffusive limitations to CO₂, dominated by the stomata, were the most important constraints to A_n. Full recovery of A_n was reached after re‐watering, characterized by quick recovery of g_m and even higher biochemical capacity, in contrast to the slower recovery of g_sw. The acclimation to long‐term WS led to decreased mesophyll and biochemical limitations, in contrast to studies in which stress was imposed more rapidly. Finally, we provide evidence that higher g_m under WS contributes to higher intrinsic water‐use efficiency (iWUE) and reduces the leaf oxidative stress, highlighting the importance of g_m as a target for breeding/genetic engineering.