Measured and modelled leaf and stand‐scale productivity across a soil moisture gradient and a severe drought

Environmental controls on carbon dynamics operate at a range of interacting scales from the leaf to landscape. The key questions of this study addressed the influence of water and nitrogen (N) availability on Pinus palustris (Mill.) physiology and primary productivity across leaf and canopy scales, linking the soil‐plant‐atmosphere (SPA) model to leaf and stand‐scale flux and leaf trait/canopy data. We present previously unreported ecophysiological parameters (e.g. V_cmax and J_max) for P. palustris and the first modelled estimates of its annual gross primary productivity (GPP) across xeric and mesic sites and under extreme drought. Annual mesic site P. palustris GPP was ～23% greater than at the xeric site. However, at the leaf level, xeric trees had higher net photosynthetic rates, and water and light use efficiency. At the canopy scale, GPP was limited by light interception (canopy level), but co‐limited by nitrogen and water at the leaf level. Contrary to expectations, the impacts of an intense growing season drought were greater at the mesic site. Modelling indicated a 10% greater decrease in mesic GPP compared with the xeric site. Xeric P. palustris trees exhibited drought‐tolerant behaviour that contrasted with mesic trees' drought‐avoidance behaviour.     

Effect of climatic variability on δ13C and tree-ring growth in piñon pine (Pinus edulis)

Understanding the response of long-lived species to natural climatic variability at multiple scales is a prerequisite for forecasting ecosystem responses to global climate change. This study investigated the response of piñon pine (Pinus edulis) to natural climatic variability using information on physiology and growth as recorded in leaves and tree rings. δ¹³C of annual leaf cohorts (δ¹³C_leaf) and tree rings (δ¹³C_ring) were measured at an ecotonal/xeric site and a mid-range/mesic site. Ring width indices (RWI) were used to estimate annual growth of individual trees. Relationships between seasonal and annual climate parameters and δ¹³C and growth were investigated. δ¹³C–climate relationships were stronger for δ¹³C_leaf than for δ¹³C_ring especially at the xeric site. The mean monthly maximum summer temperatures over May through September (summer T _max) had the strongest influence on δ¹³C_leaf. There was a strong negative relationship between RWI with summer T _max and a strong positive relationship between RWI with October to October precipitation (water–year PPN) at both sites. This suggests that piñon pine populations could be vulnerable to decreased growth and, perhaps mortality, in response to warmer, drier conditions predicted by models of global climate change.     

Above- and Belowground Net Primary Production in a Temperate Mixed Deciduous Forest

Our current ability to detect and predict changes in forest ecosystem productivity is constrained by several limitations. These include a poor understanding of belowground productivity, the short duration of most analyses, and a need for greater examination of species- or community-specific variability in productivity studies. We quantified aboveground net primary productivity (ANPP) over 3 years (1999–2001), and both belowground NPP (BNPP) and total NPP over 2 years (2000–2001) in both mesic and xeric site community types of the mixed mesophytic forest of southeastern Kentucky to examine landscape variability in productivity and its relation with soil resource [water and nitrogen (N)] availability. Across sites, ANPP was significantly correlated with N availability (R² = 0.58, P = 0.028) while BNPP was best predicted by soil moisture content (R² = 0.72, P = 0.008). Because of these offsetting patterns, total NPP was unrelated to either soil resource. Interannual variability in growing season precipitation during the study resulted in a 50% decline in mesic site litter production, possibly due to a lag effect following a moderate drought year in 1999. As a result, ANPP in mesic sites declined 27% in 2000 compared to 1999, while xeric sites had no aboveground production differences related to precipitation variability. If global climate change produces more frequent occurrences of drought, then the response of mesic sites to prolonged moisture deficiency and the consequences of shifting carbon (C) allocation on C storage will become important questions.     

Phenology constrains opportunistic growth response in <Emphasis Type="Italic">Bromus tectorum</Emphasis> L.

Seasonal resource availability may act as a constraint on plant phenology and thereby influence the range of growth responses observed among populations of annual species, especially those occupying a wide range of environments. We compared a mesic and a xeric population of the non-native, annual grass, Bromus tectorum, to examine phenology in response to interspecific competition and water availability. Using a target-neighborhood approach, we assessed how phenological patterns of the two populations affected morphological and growth responses to enhanced resource availability represented by late-season soil moisture. The xeric population exhibited a highly constrained phenology and was unable to extend the growing season despite available soil resources. Because of the low phenotypic variation, allocation to reproduction was similar across resource conditions. In contrast, the mesic population flowered later and showed a more opportunistic phenology in response to late-season water availability. The mesic population was not able to maintain consistent reproductive allocation at low resource levels. The responses of the two populations to late-season water availability were not affected by the density of neighboring plants. We suggest that post-introduction selection pressure on B. tectorum in the xeric habitat has resulted in a more fixed phenology which limits opportunistic response to unpredictable, particularly late-season resource availability. Opportunistic and fixed responses represent contrasting strategies for optimizing fitness in temporally varying environments and, while both play important roles for ensuring reproductive success, these results suggest that local adaptation to temporal resource variation may reflect a balance between flexible and inflexible phenology.     

Phenotypic plasticity and population differentiation in seeds and seedlings of the grass Anthoxanthum odoratum

Summary Seeds of Anthoxanthum odoratum were transplanted reciprocally between a xeric and a mesic field population that were genetically differentiated in adult traits. In one experiment seeds were reciprocally buried in bags in the soil, in a second experiment seeds were reciprocally sown in small plots. For most traits, site effects were much larger than seed-source effects. Germination, emergence, mortality of buried seed and recruitment were significantly higher at the xeric site than at the mesic site, irrespective of population of origin. Seed dormancy, was significantly higher for seed originating from the mesic than from the xeric population. Seedling recruits originating from the xeric population tended to be larger at both sites. Fecundity of seedling recruits depended on the environment; fecundities of plants growing in the xeric site had more than double the fecundity of plants growing in the mesic site. Phenotypic plasticity rather than population differences determined variation in performance in the seed and seedling stages.     

Growth reactions of Pinus sylvestris L. and Quercus pubescens Willd. to drought years at a xeric site in Valais, Switzerland

In Valais, an inner-Alpine dry valley in Switzerland, low-elevation Scots pine (Pinus sylvestris L.) forests are changing. While pine shows high mortality rates, deciduous species, in particular pubescent oak (Quercus pubescens Willd.), are becoming more abundant. We hypothesise that increasing drought and the species-specific drought tolerance are key factors in these processes. In this study, the growth reaction to drought years of pine and oak growing at a xeric site in Valais was analysed using dendrochronological and wood anatomical methods. Congruent with theoretical expectations, the tree-ring widths of both species, the mean lumen area of earlywood vessels in oak and the number of tracheids in a radial row in pine decreased in response to dry conditions. However, both species also showed reactions deviating from those known from mesic sites: In oak, the mean lumen area of latewood vessels increased in drought years. In pine, in the driest year of the period (1976), the mean radial diameter increased in latewood and decreased only slightly in earlywood. These results emphasises that the process of wood formation and cell functionality at xeric sites is not completely understood yet. Both species seem to have difficulties to adapt the size of their water-conducting cells to strongly reduced water availability in drought years. Additionally, the cell number is strongly reduced. Thus it remains unclear if both species can maintain sufficient water transport under increasingly dry conditions.     

Plasticity in gas‐exchange physiology of mature Scots pine and European larch drive short‐ and long‐term adjustments to changes in water availability

Adjustment mechanisms of trees to changes in soil‐water availability over long periods are poorly understood, but crucial to improve estimates of forest development in a changing climate. We compared mature trees of Scots pine (Pinus sylvestris) and European larch (Larix decidua) growing along water‐permeable channels (irrigated) and under natural conditions (control) at three sites in inner‐Alpine dry valleys. At two sites, the irrigation had been stopped in the 1980s. We combined measurements of basal area increment (BAI), tree height and gas‐exchange physiology (Δ¹³C) for the period 1970–2009. At one site, the Δ¹³C of irrigated pine trees was higher than that of the control in all years, while at the other sites, it differed in pine and larch only in years with dry climatic conditions. During the first decade after the sudden change in water availability, the BAI and Δ¹³C of originally irrigated pine and larch trees decreased instantly, but subsequently reached higher levels than those of the control by 2009 (15 years afterwards). We found a high plasticity in the gas‐exchange physiology of pine and larch and site‐specific responses to changes in water availability. Our study highlights the ability of trees to adjust to new conditions, thus showing high resilience.     

Variation in community structure of gall‐inducing insects associated with a tropical plant supports the hypothesis of competition in stressful habitats

Environmental factors act as drivers of species coexistence or competition. Mesic environments favor the action of parasites and predators on gall communities, while the factors that determine the structure of gall communities in xeric environments remain unknown. We evaluated the structure of gall communities along an environmental gradient defined by intrinsic plant characteristics, soil fertility, and aridity, and investigated the role of competition as a structuring force of gall communities in xeric environments. We created null models to compare observed and simulated patterns of co‐occurrence of galls and used the C‐score index to assess community aggregation or segregation. We used the NES C‐score (standardized C‐score) to compare patterns of co‐occurrence with parameters of environmental quality. Xeric environments had poorer and more arid soils and more sclerophyllous plants than mesic environments, which was reflected in the distribution patterns of gall communities. Values of the C‐score index revealed a segregated distribution of gall morphospecies in xeric environments, but a random distribution in mesic environments. The low availability of resources for oviposition and the high density of gallers in xeric environments reinforce interspecific competition as an important structuring force for gall communities in these environments.     

Sensitivity of French temperate coniferous forests to climate variability and extreme events (Abies alba,Picea abies and Pinus sylvestris)

Questions: (1) How do extreme climatic events and climate variability influence radial growth of conifers (silver fir, Norway spruce, Scots pine)? (2) How do elevation and soil water capacity (SWC) modulate sensitivity to climate? Location: The sampled conifer stands are in France, in western lowland and mountain forests, at elevations from 400 to 1700 m, and an SWC from 50 to 190 mm. Methods: We established stand chronologies for total ring width, earlywood and latewood width for the 33 studied stands (985 trees in total). Responses to climate were analysed using pointer years and bootstrapped response functions. Principal component analysis was applied to pointer years and response function coefficients in order to elucidate the ecological structure of the studied stands. Results: Extreme winter frosts are responsible for greater growth reductions in silver fir than in Norway spruce, especially at the upper elevation, while Scots pine was the least sensitive species. Exceptional spring droughts caused a notable growth decrease, especially when local conditions were dry (altitude<1000 m and SWC<100 mm for silver fir, western lowlands for Scots pine). Earlywood of silver fir depended on previous September and November and current‐year February temperature, after which current June and July water supply influenced latewood. Earlywood of Norway spruce was influenced by previous September temperature, after which current spring and summer droughts influenced both ring components. In Scots pine, earlywood and latewood depended on the current summer water balance. Local conditions mainly modulated latewood formation. Conclusions: If the climate becomes drier, low‐elevation dry stands or trees growing in western lowlands may face problems, as their growth is highly dependent on soil moisture availability.     

Species‐specific differences in water uptake depth of mature temperate trees vary with water availability in the soil

• Temperate tree species differ in their physiological sensitivity to declining soil moisture and drought. Although species‐specific responses to drought have often been suggested to be the result of different water uptake depths, empirical evidence for such a mechanism is scarce.
• Here we test if differences in water uptake depths can explain previously observed species‐specific physiological responses of temperate trees to drought and if the water uptake depth of different species varies in response to declining soil moisture. For this purpose, we employed stable oxygen and hydrogen isotopes of soil and xylem water that we collected over the course of three growing seasons in a mature temperate forest in Switzerland.
• Our data show that all investigated species utilise water from shallow soil layers during times of sufficient soil water supply. However, Fraxinus excelsior, Fagus sylvatica and Acer pseudoplatanus were able to shift their water uptake to deeper soil layers when soil water availability decreased in the topsoil. In contrast, Picea abies, was not able to shift its water uptake to deeper soil layers.
• We conclude from our data that more drought‐resistant tree species are able to shift their water uptake to deeper soil layers when water availability in the topsoil is becoming scarce. In addition, we were able to show that water uptake depth of temperate tree species is a trait with high plasticity that needs to be characterised across a range of environmental conditions.

     

Carbon, nitrogen, and hydrogen isotope ratios in creekside trees in western Kansas

Three species of creekside trees were monitored weekly during the 2007 and 2008 growing seasons. The 2007 growing season was wet early, but became drier as the season progressed. In contrast, the 2008 growing season was dry early, but became wetter as the season progressed. Creekside trees were measured to determine effects of changing water regimes on leaf-level processes. Lonicera tatarica plants were compared to Morus alba and Celtis occidentalis trees. Leaves were monitored for changes in stomatal conductance, transpiration, δ¹³C, δ¹⁵N, δD, leaf temperature, and heat losses via latent, sensible, and radiative pathways. δD of creek water was more similar to ground water than to rain water, but the creek was partially influenced by summer rains. δD of bulk leaf material was significantly higher in individuals of C. occidentalis compared to the other species, consistent with source water coming from shallower soil layers. Despite decreasing water levels, none of these tree species showed signs of water stress. There were no significant differences between species in stomatal conductance or transpiration. Leaf δ¹³C was significantly lower in individuals of L. tatarica compared to the other species. Differences in δ¹³C were attributed to a lower carboxylation capacity, consistent with lower leaf nitrogen content in L. tatarica plants. Leaf δ¹⁵N was significantly lower in individuals of L. tatarica compared to the other species, consistent with uptake of a different N source. Two of the three sites appeared to be affected by inorganic N from fertilizer run-off. Evidence is presented that these species acquired water and nitrogen from different sources, resulting from differences in root uptake patterns.     

Radial-growth and wood-anatomical changes in overaged Quercus pyrenaica coppice stands: functional responses in a new Mediterranean landscape

Recent land-use changes in intensively managed forests such as Mediterranean coppice stands might profoundly alter their structure and function. We assessed how the abandonment of traditional management practices in coppice stands, which consisted of short cutting-cycles (10–15 years), has caused overaging (stems are usually much older than when they were coppiced) and altered their wood anatomy and hydraulic architecture. We studied the recent changes of wood anatomy, radial growth, and hydraulic architecture in two stands of Quercus pyrenaica, a transitional Mediterranean oak with ring-porous wood forming coppice stands in W–NW Spain. We selected a xeric and a mesic site because of their contrasting climates and disturbance histories. The xeric site experienced an intense defoliation after the severe 1993–1994 summer drought. The mesic site was thinned in late 1994. We studied the temporal variability in width, vessel number and diameter, and predicted the hydraulic conductivities (K _h) of earlywood and latewood. In the mesic site, we estimated the vulnerability to xylem cavitation of earlywood vessels. Overaging caused a steep decline in latewood production at a cambial age of 14 years., which was close to the customary cutting cycle of Q. pyrenaica. The diameter distribution of vessels was bimodal, and latewood vessels only accounted for 4% of the K _h. Overaging, acting as a predisposing factor in the decline episode, was observed at the xeric site, where most trees did not produce latewood in 1993–1995. At the mesic site, thinned trees formed wider tree-rings, more latewood and multiseriate tree-rings than overaged trees. The growth enhancement remained 8 years after thinning. Most of the hydraulic conductivity in earlywood was lost in a narrow range of potentials, between −2.5 and −3.5 MPa. We have shown how hydraulic conductivity and radial growth are closely related in Q. pyrenaica and how aging modulates this relationship.     

On the influence of provenance to soil quality enhanced stress reaction of young beech trees to summer drought

Climate projections propose that drought stress will become challenging for establishing trees. The magnitude of stress is dependent on tree species, provenance, and most likely also highly influenced by soil quality. European Beech (Fagus sylvatica) is of major ecological and economical importance in Central European forests. The species has an especially wide physiological and ecological amplitude enabling growth under various soil conditions within its distribution area in Central Europe. We studied the effects of extreme drought on beech saplings (second year) of four climatically distinct provenances growing on different soils (sandy loam and loamy sand) in a full factorial pot experiment. Foliar δ¹³C, δ¹⁵N, C, and N as well as above‐ and belowground growth parameters served as measures for stress level and plant growth. Low‐quality soil enhanced the effect of drought compared with qualitatively better soil for the above‐ and belowground growth parameters, but foliar δ¹³C values revealed that plant stress was still remarkable in loamy soil. For beeches of one provenance, negative sandy soil effects were clearly smaller than for the others, whereas for another provenance drought effects in sandy soil were sometimes fatal. Foliar δ¹⁵N was correlated with plant size during the experiment. Plasticity of beech provenances in their reaction to drought versus control conditions varied clearly. Although a general trend of declining growth under control or drought conditions in sandy soil was found compared to loamy soil, the magnitude of the effect of soil quality was highly provenance specific. Provenances seemed to show adaptations not only to drought but also to soil quality. Accordingly, scientists should integrate information about climatic pre‐adaptation and soil quality within the home range of populations for species distribution modeling and foresters should evaluate soil quality and climatic parameters when choosing donor populations for reforestation projects.     

Adjustments in hydraulic architecture of Pinus palustris maintain similar stomatal conductance in xeric and mesic habitats

We investigated relationships between whole-tree hydraulic architecture and stomatal conductance in Pinus palustris Mill. (longleaf pine) across habitats that differed in soil properties and habitat structure. Trees occupying a xeric habitat (characterized by sandy, well-drained soils, higher nitrogen availability and lower overstory tree density) were shorter in stature and had lower sapwood-to-leaf area ratio (A(S):A(L)) than trees in a mesic habitat. The soil-leaf water potential gradient (psiS - psiL) and leaf-specific hydraulic conductance (kL) were similar between sites, as was tissue-specific hydraulic conductivity (Ks) of roots. Leaf and canopy stomatal conductance (gs and Gs, respectively) were also similar between sites, and they tended to be somewhat higher at the xeric site during morning hours when vapour pressure deficit (D) was low. A hydraulic model incorporating tree height, A(S):A(L) and psiS-psiL accurately described the observed variation in individual tree G(Sref) (G(S) at D = 1 kPa) across sites and indicated that tree height was an important determinant of G(Sref) across sites. This, combined with a 42% higher root-to-leaf area ratio (A(R):A(L)) at the xeric site, suggests that xeric site trees are hydraulically well equipped to realize equal--and sometimes higher potential for conductance compared with trees on mesic sites. However, a slightly more sensitive stomatal closure response to increasing D observed in xeric site trees suggests that this potential for higher conductance may only be reached when D is low and when the capacity of the hydraulic system to supply water to foliage is not greatly challenged.     

Spatial Heterogeneity of Denitrification in Semi-Arid Floodplains

Riparian ecosystems are recognized as sinks for inorganic nitrogen (N). Denitrification, a heterotrophic microbial process, often accounts for a significant fraction of the N removed. Characteristics of both riparian soils and hydrologic vectors may constrain the locations where denitrification can occur within riparian ecosystems by influencing the distribution of substrates, water, and suitable redox conditions. We employed spatially explicit methods to quantify heterogeneity of soil characteristics and potential rate of denitrification in semi-arid riparian ecosystems. These results allow us to evaluate the relative contributions of hydrologic vectors (water courses that convey materials) and soil resources (materials required by biota) to spatial heterogeneity of denitrification. During dry and monsoon seasons we contrasted a mesic site, characterized by shallow groundwater and annual inundation by floods, with a xeric site that is inundated less often and has a deeper water table. Potential denitrification was detected throughout the mesic floodplain and the average rate of denitrification was greater at the mesic site than at the xeric site, indicating the influence of water availability on denitrification. At the xeric reach, sharp declines in pools of soil resources and rate of denitrification occurred away from the stream, demonstrating the importance of the stream in determining spatial patterns. Using geographically weighted regression analysis, we determined that soil organic matter and soil nitrate were significant predictors of denitrification at the xeric site, but that factors influencing denitrification varied spatially. Spatial heterogeneity of carbon (C) and N substrates in soils likely influenced spatial patterns of denitrification, but distribution of C and N substrates was ultimately organized by hydrologic vectors. Droughts will increase the abundance of reaches with hydrogeomorphic templates similar to the xeric reach studied here. Consequences of such a transition may include a reduced rate of denitrification and patchy distribution of denitrification in floodplain soils, which will decrease the contribution of riparian ecosystems to N removal. TKH designed and completed the study and wrote the paper; EAW contributed methods and edited the paper; NBG designed the study and edited the paper.     

Thermal habitat segregation among morphotypes of whitefish (Coregonus lavaretus: Salmonidae) and invasive vendace (C. albula): a mechanism for co‐existence?

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Variation in vegetation and seed bank in a Chilean semi‐arid community affected by ENSO 1997

Abstract. We followed plant cover and soil seed density of shrubs and herbs in two markedly contrasting years with regard to annual rainfall in two opposite‐facing slopes (mesic vs xeric) and an intervening, relatively wet, ravine of a typical creek in semi‐arid Chile. During the ENSO year 1997 cover of ephemerals increased in all three sites; 43% vs 8% on the xeric slope 75% vs 26% on the mesic slope and 57% vs 32% in the ravine. The number of species was almost three times higher on the xeric slope (27 vs 10 species), increased by 47% on the mesic slope (28 vs 19) and by 14% in the ravine (24 vs 21). Cover of shrubs plus perennial grasses increased from 52% in 1996 to 59% in 1997 on the xeric slope, but no significant cover changes were found on the mesic slope or the ravine. On the xeric slope peak density of seeds was 4500/m² in 1996, while during the ENSO year it was 24000/m². On the mesic slope equivalent values were 3000 and 17000 seeds/m² while in the ravine figures were 8500 and 27000 seeds/m². The strong responses of ephemerals and seed bank to the ENSO‐driven wet year of 1997 demonstrate the importance of this phenomenon in replenishing, recovering and probably maintaining ephemeral vegetation in this region.     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Relevance of using whole-ring stable isotopes of black spruce trees in the perspective of climate reconstruction

Studies in dendroisotope chemistry suggested that latewood cellulose contains better climatic records than whole-ring cellulose. However, this approach has never been tested on northeastern Canadian spruce trees. This study compares dendroisotopic series of cellulose from late and whole ring, and analyses their statistical relationships with hydro-climatic variables with the aim of selecting the best suited protocol for future hydro-climatic reconstruction in the downstream sector of Churchill River basin of Labrador, Canada. To this end, δ¹³C and δ¹⁸O series from latewood (LW) and whole ring (WR) α-cellulose of black spruce trees (Picea mariana [Mill.] B.S.P.) were produced for the 1940–2010 period. The results show strong correlations between LW and WR isotopic series suggesting that there are no important variation in the isotopic ratios during the growing year and that black spruce trees use photosynthates of the current growing season to form their earlywood. Moreover, LW and WR δ¹³C and δ¹⁸O show similar relationships with both maximum temperature (T_max) and Churchill River discharge. Correlations are higher when combining δ¹³C and δ¹⁸O for LW and WR. Overall, those correlations support the indirect relationship between tree-ring isotopic series and river discharge, as they are integrators of several climatic variables and derived parameters (T_max, relative humidity, evapotranspiration, etc.). The LW and WR isotopic series give similar statistical relationships with hydro-climatic variables, and the WR treatment is faster (separation easier compared to LW). Thus, for black spruce the use of combined isotopic series in WR can be favored over LW for hydro-climatic reconstruction in the study region.     

Revisiting water loss in insects: a large scale view

Desiccation resistance in insects has long been thought to covary with environmental water availability, and to involve changes in both cuticular and respiratory transpiration. Here, we adopt a large-scale approach to address both issues. Water loss rate and precipitation are positively related at global scales. A significant proportion (68%) of the interspecific variation in water loss rate is explained at the genus level or above. The relationship between metabolic rate and water loss rate differs substantially between mesic and xeric species. While these variables covary as a consequence of their independent covariation with body mass in mesic species, this is not the case in xeric species. In the latter, there is a strong relationship between the residuals of the water loss rate-body mass and metabolic rate-body mass relationships, and water loss rate is much reduced. Moreover, because metabolic rate does not differ significantly between xeric and mesic species of a similar size, respiratory transpiration constitutes a greater proportion of total water loss in xeric than in mesic species of a similar size. This implies that respiratory transpiration and the extent to which it can be modified must be of considerable importance in xeric insect species, although finer scale studies suggest otherwise.