Plant and small mammal richness correlate positively in a biodiversity hotspot

Hotspots of biodiversity are important areas in facilitating an understanding of species richness and its maintenance. Herbivores can increase plant richness by reducing dominant plant species thus providing space for subdominant species. As small mammals are abundant in the Succulent Karoo and therefore might affect plant richness by means of herbivory, we tested if this mechanism might exist in the Succulent Karoo in southern Africa, a biodiversity hotspot due to its extraordinary plant richness. At ten ecologically different study sites we measured plant and small mammal richness and diversity and determined 11 abiotic factors including soil composition, altitude and rainfall. We found positive correlations between plant richness and the number of small mammal species. A general linear model revealed that the number of small mammal species was more important than abiotic factors in explaining variation in plant richness. To test whether small mammals might directly influence plant richness, we studied the influence of the bush-Karoo rat Otomys unisulcatus, a central place forager, on the plant community. The immediate surroundings of occupied O. unisulcatus nests showed significantly higher plant richness than control areas. We conclude that small mammals can have a positive effect on plant richness in the Succulent Karoo. While experimental data are needed to support these correlative results, the results of our study indicate that areas of high small mammal richness should be included in conservation programs of the Succulent Karoo.     

Do increased summer precipitation and N deposition alter fine root dynamics in a Mojave Desert ecosystem?

Climate change is expected to impact the amount and distribution of precipitation in the arid southwestern United States. In addition, nitrogen (N) deposition is increasing in these regions due to increased urbanization. Responses of belowground plant activity to increases in soil water content and N have shown inconsistent patterns between biomes. In arid lands, plant productivity is limited by water and N availability so it is expected that changes in these factors will affect fine root dynamics. The objectives of this study were to quantify the effects of increased summer precipitation and N deposition on fine root dynamics in a Mojave Desert ecosystem during a 2‐year field experiment using minirhizotron measurements. Root length density, production, and mortality were measured in field plots in the Mojave Desert receiving three 25 mm summer rain events and/or 40 kg N ha^?1 yr^?1. Increased summer precipitation and N additions did not have an overall significant effect on any of the measured root parameters. However, differences in winter precipitation resulting from interannual variability in rainfall appeared to affect root parameters with root production and turnover increasing following a wet winter most likely due to stimulation of annual grasses. In addition, roots were distributed more deeply in the soil following the wet winter. Root length density was initially higher under canopies compared to canopy interspaces, but converged toward the end of the study. In addition, roots tended to be distributed more deeply into the soil in canopy interspace areas. Results from this study indicated that increased summer precipitation and N deposition in response to climate change and urbanization are not likely to affect fine root dynamics in these Mojave Desert ecosystems, despite studies showing aboveground plant physiological responses to these environmental perturbations. However, changes in the amount and possibly distribution of winter precipitation may affect fine root dynamics.     

Rainfall and fluctuating plant populations in relation to distributions and numbers of desert rodents in southern Nevada

Summary Fluctuations in rainfall and size of desert rodent and plant populations for each of five consecutive years were documented on 68 sites in Mojave, Great Basin, and transition desert communities of central-southern Nevada. Post-reproduction rodent densities in the summer usually followed the seasonal patterns of winter annual success the previous spring, in turn directly related to the success of germination the preceding autumn.In the Dipodomys merriami environment, at the lower elevations with relatively low rainfall and high temperatures of the Mojave Desert, D. merriami and D. deserti numbers (indicating success or failure of reproduction) closely followed the seasonal and site patterns of winter annual success. In the Dipodomys microps environment, at the middle and higher elevations with the relatively high rainfall and lower temperatures of the transition and Great Basin deserts, D. microps numbers followed winter annual success patterns in years of high or low rainfall in the months preceding reproduction, but there was reproduction in years of intermediate rainfall whether or not there were winter annuals in the environment. Reproductive patterns of D. merriami in the D. microps environment followed those of D. microps. Perognathus longimembris, the ubiquitous heteromyid of the region, also exhibited both reproductive patterns, depending on whether it was in the D. merriami or the D. microps environment. The Mojave Desert P. formosus, edaphically restricted in the region, followed the patterns of the D. merriami environment and its numbers were highly correlated with winter annual success.Herbaceous perennials, capable of large germinations of plants behaving as biennials or winter annuals, are shown to have large effects on D. merriami numbers. Size of populations of Eurasian annual species of Bromus and Salsola were not correlatable with rodent reproduction patterns.From all patterns in all environments, it is inferred that environmental conditions affecting water balance in the rodents play the key role in initiation of reproduction. Environmental moisture levels during the winter season are usually under the control of rainfall of late autumn-early winter, coincidentally the period of greatest predictability of heavy rains. The herbaceous plants, present in the winter months most years, are available sources of water (and vitamins) at the time of initiation of reproductive activity. For D. microps, identified with the mesic environments of the region, the herbaceous plants may or may not be necessary sources of water for reproduction, depending on the seasonal rainfall regime. For D. merriami, herbaceous plants appear to serve as a water reservoir required for reproduction in its usual environments. The same relationships evidently obtain for the other heteromyids, and the cricetids as a group.Studies conducted under U.S. Atomic Energy Commission Contracts AT(04-1) Gen-12 and AT(11-1) 2307, and at the Nevada Test Site as part of CETO Project 61.5.4     

Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.)

Climatic and biogeographical associations of southern African dung beetles (Scarabaeinae, Coprinae) were analysed from a collection amassed between 1971 and 1986. Endemism to Africa south of 15°S was much greater in southwesterly climates (winter rainfall, bimodal spring/autumn rainfall, arid late summer rainfall) than to the north‐east in the moist, mid‐summer rainfall region. Major biogeographical groups centred to the south‐west comprised predominantly southern African endemics, Western/Eastern Cape coast, Karoo, Karoo/Namib, northern Namibia and the south‐western Kalahari. Biogeographical groups centred on the south‐eastern highlands and the subtropical east coast (mid‐summer rainfall) also comprised predominantly southern African endemics. All other major groups centred to the north‐east in the mid‐summer rainfall region comprised predominantly species with widespread tropical biogeographical affiliations, pan‐southern Africa but centred in the east, pan‐mid‐summer rainfall region, eastern mid‐summer rainfall region, tropical east Zimbabwe/central Moçambique, subtropical/tropical game reserves (non‐ruminant dung specialists). There were cross‐climatic differences in taxonomic composition of the fauna. Within the winter rainfall region, percentage species composition of Scarabaeinae was greater whereas that of the coprine tribe, Onthophagini, was lower than within the other three climatic regions. Percentage species richness of most other tribes of Coprinae differed little between most climates but that of Scarabaeinae declined from west to east (Canthonini, Scarabaeini), east to west (Sisyphini) or to either side of the late summer rainfall region (Gymnopleurini).     

Reference intervals and physiologic alterations in hematologic and biochemical values of free-ranging desert tortoises in the Mojave Desert

Desert tortoise (Gopherus agassizii) populations have experienced precipitous declines resulting from the cumulative impact of habitat loss, and human and disease-related mortality. Evaluation of hematologic and biochemical responses of desert tortoises to physiologic and environmental factors can facilitate the assessment of stress and disease in tortoises and contribute to management decisions and population recovery. The goal of this study was to obtain and analyze clinical laboratory data from free-ranging desert tortoises at three sites in the Mojave Desert (California, USA) between October 1990 and October 1995, to establish reference intervals, and to develop guidelines for the interpretation of laboratory data under a variety of environmental and physiologic conditions. Body weight, carapace length, and venous blood samples for a complete blood count and clinical chemistry profile were obtained from 98 clinically healthy adult desert tortoises of both sexes at the Desert Tortoise Research Natural area (western Mojave), Goffs (eastern Mojave) and Ivanpah Valley (northeastern Mojave). Samples were obtained four times per year, in winter (February/March), spring (May/June), summer (July/August), and fall (October). Years of near-, above- and below-average rainfall were represented in the 5 yr period. Minimum, maximum and median values, and central 95 percentiles were used as reference intervals and measures of central tendency for tortoises at each site and/or season. Data were analyzed using repeated measures analysis of variance for significant (P < 0.01) variation on the basis of sex, site, season, and interactions between these variables. Significant sex differences were observed for packed cell volume, hemoglobin concentration, aspartate transaminase activity, and cholesterol, triglyceride, calcium, and phosphorus concentrations. Marked seasonal variation was observed in most parameters in conjunction with reproductive cycle, hibernation, or seasonal rainfall. Year-to-year differences and long-term alterations primarily reflected winter rainfall amounts. Site differences were minimal, and largely reflected geographic differences in precipitation patterns, such that results from these studies can be applied to other tortoise populations in environments with known rainfall and forage availability patterns.     

Seasonal Acclimation of Stem Photosynthesis in Woody Legume Species from the Mojave and Sonoran Deserts of California

Photosynthesis (Pn) was measured in stems of two desert legumes, Caesalpinia virgata at a low elevation site (118 m) in the Sonoran Desert and Senna armata at a higher elevation (950 m) in the Mojave Desert. The lower elevation site experienced higher spring and summer temperatures than the higher elevation site, but the air vapor pressure, irradiance, and rainfall patterns were similar. Mid-morning maximum stem Pn was highest in May for C. virgata (7.8 [mu]mol m-2 s-1) and in July for S. armata (5.8 [mu]mol m-2 s-1). The seasonal variation in maximum stem Pn was not associated with changes in bulk tissue water potential or chlorenchyma tissue nitrogen concentration. The main environmental regulators of seasonal stem Pn were temperature and leaf to air vapor pressure gradient. Light-response curves indicated no major differences in apparent quantum yield or light compensation point between the spring and summer, but light-saturated stem Pn at ambient temperature decreased for C. virgata between these seasons. The optimal temperature for stem Pn remained the same for both species between the spring and the summer. However, stem Pn of both species increased at all temperatures between the spring and summer. Potential stem Pn under optimal conditions and CO2-saturated stem Pn increased for both species between spring and summer. The increase in stem Pn potential allowed these species to maintain stem Pn during the summer even though stem Pn responses to temperature and vapor pressure did not acclimate to seasonal climatic conditions.     

Alien plant assemblages near roads in arid and semi-arid South Africa

Herbaceous and woody alien plants visible from a moving vehicle were recorded along 1 km roadside transects at 5 km intervals over a distance of 5869 km in the semi-arid and arid Fynbos, Succulent Karoo, Nama Karoo and Arid Savanna (Kalahari) biomes in South Africa. Each 1 km transect was classified by biome and vegetation type, mean annual rainfall, rainfall seasonality, soil surface type and landuse adjoining the roadside. Although travelling speed affected the range and frequency of plant species observed, the method was repeatable at a speed of 100 km h^?1. Alien plants occurred in 98% of 119 Fynbos, 81% of 204 Succulent Karoo, 72% of 661 Nama Karoo, 47% of 171 Arid Savanna and 100% of seventeen Grassland transects. Alien species richness per site was correlated with mean annual rainfall, but in all regions, sites adjacent to cultivation had significantly more alien species than sites adjoining rangeland. The alien plant assemblage of the arid winter-rainfall Succulent Karoo included species from mesic winter-rainfall lowland Fynbos and the arid Nama Karoo receives unseasonal rainfall. The frequencies of Prosopis spp., Atriplex spp. and Opuntia ficus-indica were not significantly greater near cultivation, and these perennial plants, all of which are dispersed by indigenous and domestic animals, can invade natural rangeland in arid and semi-arid southern Africa.     

Plant species richness, endemism, and genetic resources in Namibia

Namibia is a floristically diverse, arid to mesic country, with several highly distinct taxa. Including naturalized plants, there are about 4334 vascular plant species and infraspecific taxa within the country's borders, a substantial increase from the existing major reference work. Dominant families are the Poaceae (422species), Fabaceae (377), Asteraceae (385) and Mesembryanthemaceae (177). Freshwater algae and most other groups of lower plants remain poorly known. Concentrations of plant species richness are found in the Succulent Karoo biome, Kaokoveld, Otavi highland/Karstveld area, Okavango Basin, and Khomas highlands. Recent studies have led to a new estimate of 687 endemic plant species, defined as those contained wholly within Namibia's borders, amounting to about 17% of the Namibian flora. At least a further 275 species are Namib Desert endemics shared between the Kaokoveld and southern Angola (75spp.) and between the Succulent Karoo and northwestern South Africa (200spp.). Research on plant genetic resources is focused on species of potential or actual agricultural importance, such as pearl millet, Pennisetum glaucum, and cucurbits. Many wild plants have considerable genetic diversity and development potential. Primary threats to plant diversity fall in the category of poor land management and inappropriate development.     

Remarkable medium-term dynamics of leaf succulent Mesembryanthemaceae shrubs in the winter-rainfall desert of northwestern Namaqualand,South Africa

Populations of shrubs in a winter-rainfall (ca. 70 mm yr_-1 desert community (Succulent Karoo), dominated by leaf succulents, showed remarkable dynamics over a 17-yr period. After a severe drought in 1979, which caused high plant mortality, perennial species number in a permanent 10 m×10 m plot doubled between 1980 and 1996, when the maximum of 41 species was recorded. Numbers of individuals of evergreen, leaf succulent shrubs also doubled over the same period, but showed fluctuations in response to dry years during the monitoring period. Detailed observations on the four dominant leaf succulent shrubs (all members of the Mesembryanthemaceae) between 1983 and 1996, showed species-specific patterns in population turnover. Mortality of all observed individuals ranged between 60% and 85%, and the proportion of the population that was recruited over this period ranged from 62% to 89%. In only one species did individuals persist throughout the monitoring period. Mean ages of individuals, excluding seedlings with lifespans of <1 yr, and individuals observed at the beginning of the monitoring period, ranged from 4.6 yr to 5.6 yr. Patterns of mortality and recruitment showed substantial differences among species and were not all attributable to rainfall patterns. Overall, the turnover of the shrub populations over the 15-yr monitoring period was remarkably high for a system of desert perennials. These unusual population patterns may explain the unique structure of leaf succulent-dominated communities in the Succulent Karoo.     

Carbon isotope ratio measurements of succulent plants in southern Africa

Summary Succulent plants representing 16 families and a variety of growth forms originating from winter, summer, and year-round rainfall regimes in southern Africa were analyzed for carbon isotope ratios. Most families had species with ¹³C values indicative of CAM, particularly those from winter and year-round rainfall regimes. Plants with ¹³C values intermediate between CAM and C₃, indicating flexible photosynthetic pathways, were generally leafy perennials subject to seasonal tissue dehydration. Reproductive tissue tended to have less negative ¹³C values than vegetative tissue on the same plant, indicating drought-season origin of the former.CIW-DPB Publication No. 609     

Biogeography of prostrate-leaved geophytes in semi-arid South Africa: hypotheses on functionality

Nowhere is the species diversity of geophytes greater than in the five mediterranean-climate ecosystems of the world. Of these, the Cape mediterranean zone of South Africa is the most speciose. While the relative diversity and importance of geophytes of all of the other four mediterranean regions of the world drops off sharply as one moves into adjacent winter-rainfall desert regions, geophytes in the semi-arid to arid Succulent Karoo (including Namaqualand) remain a very important component of the flora, both in terms of abundance and diversity (comprising 13 to 29% of the regional floras in this region). Apart from species richness, there are also a number of interesting geophyte growth forms in this region. One unusual growth form is geophytes with flattened leaves that lie prostrate on the soil surface. At least eight families (Amaryllidaceae, Colchicaceae, Eriospermaceae, Geraniaceae, Hyacinthaceae, Iridaceae, Orchidaceae and Oxalidaceae) exhibit this growth form. While this growth form is relatively common in many geophyte lineages in the Succulent Karoo biome and the Cape mediterranean zone (Fynbos biome), and occurs infrequently through the summer-rainfall temperate regions of Africa, it is virtually absent in other regions worldwide. A null hypothesis is that the prostrate leaved trait is a neutral characteristic, however biogeographical data do not support this. A neutral trait would be unlikely to show such a clear pattern of distribution. Several alternative hypotheses on the adaptive significance of this growth form are discussed. These include: avoidance of herbivory, reduction in competition from neighbors, creation of a CO₂ enriched environment below the leaves, reduction of water loss around the roots, reduction of water loss through transpiration, precipitation of dew on the leaves and maintenance of optimal leaf temperatures for growth.     

Importance of Seasonality for the Response of a Mesic Temperate Grassland to Increased Precipitation Variability and Warming

Timing of precipitation events within the growing season and the non-uniformity of warming might be decisive for alterations in productivity and community composition, with consequences for ecosystem functioning. The responses of aboveground production, community composition, functional group and species evenness to altered intra-annual precipitation variability and their interactions with winter or summer warming were examined in European, mesic temperate grassland. Increased precipitation variability with an induced spring drought resulted in a 17% reduction in ANPP, and late drought reduced ANPP by 18% compared to regular rainfall patterns throughout the entire growing season. Winter warming increased ANPP by 12%, whereas summer warming showed no significant effect on biomass but decreased species richness. The effects of increased precipitation variability and warming on ANPP were independent of each other. Forbs benefited from high precipitation variability with spring drought events, likely due to reduced competitive pressure by decreasing, water stressed grasses. Increased precipitation variability coinciding with higher summer temperatures led to reduced species evenness and likely promoted the establishment of specialists and drought-tolerant species. Seasonality of climatic factors, here early versus late drought events in the high precipitation variability treatments, was important in driving shifts in community composition but not for decreases in ANPP. Non-uniform warming, here winter versus summer, affected the direction of response of both community composition and ANPP. Variability of resources is affecting ecosystem processes and species interactions. Recognition of seasonality and non-uniformity of climatic factors will improve predictions of plant performance and biotic interactions in response to climate change.     

Dynamics of cover, UV-protective pigments, and quantum yield in biological soil crust communities of an undisturbed Mojave Desert shrubland

Biological soil crusts are an integral part of dryland ecosystems. We monitored the cover of lichens and mosses, cyanobacterial biomass, concentrations of UV-protective pigments in both free-living and lichenized cyanobacteria, and quantum yield in the soil lichen species Collema in an undisturbed Mojave Desert shrubland. During our sampling time, the site received historically high and low levels of precipitation, whereas temperatures were close to normal. Lichen cover, dominated by Collema tenax and C. coccophorum, and moss cover, dominated by Syntrichia caninervis, responded to both increases and decreases in precipitation. This finding for Collema spp. at a hot Mojave Desert site is in contrast to a similar study conducted at a cool desert site on the Colorado Plateau in SE Utah, USA, where Collema spp. cover dropped in response to elevated temperatures, but did not respond to changes in rainfall. The concentrations of UV-protective pigments in free-living cyanobacteria at the Mojave Desert site were also strongly and positively related to rainfall received between sampling times (R² values ranged from 0.78 to 0.99). However, pigment levels in the lichenized cyanobacteria showed little correlation with rainfall. Quantum yield in Collema spp. was closely correlated with rainfall. Climate models in this region predict a 3.5–4.0 °C rise in temperature and a 15–20% decline in winter precipitation by 2099. Based on our data, this rise in temperature is unlikely to have a strong effect on the dominant species of the soil crusts. However, the predicted drop in precipitation will likely lead to a decrease in soil lichen and moss cover, and high stress or mortality in soil cyanobacteria as levels of UV-protective pigments decline. In addition, surface-disturbing activities (e.g., recreation, military activities, fire) are rapidly increasing in the Mojave Desert, and these disturbances quickly remove soil lichens and mosses. These stresses combined are likely to lead to shifts in species composition and the local extirpation of some lichen or moss species. As these organisms are critical components of nutrient cycling, soil fertility, and soil stability, such changes are likely to reverberate throughout these ecosystems.     

新疆莫索湾地区短命植物的初步研究

莫索湾地区位于准噶尔盆地西南部主要由固定、半固定以及半流动的沙丘、沙地组成,是古尔班通古特沙漠的组成部分。是典型的温带荒漠气候,代表性植被是小半乔木荒漠。在这种严酷的大陆性荒漠气候条件下,由下列原因使短命植物得以发育;1.年降水量虽少,但冬有积雪,春有一定降雨,为短命植物的发育提供了有利的水分条件;2．基质为疏松沙层,毛细管作用微弱,在少雨干旱的环境中,具有一定的保水特性。春季至夏初间,湿沙层的存在为具浅根系的短命植物整个生长发育期间提供了十分有利的水分供应;3．虽然冬严寒,夏炎热,但春季和暖。其热量状况可满足短命植物进行高效光合生产的需要,尤其是较大的日温差,对光合产物的积累非常有利。莫索湾地区共有短命植物42种,分属于38属,16科。其生活型组成有短营养期多年生和短营养期一年生植物两大类群,根据繁殖特点和地下器官的形态特征等生态生物学特点进一步细分为7类。本区短命植物是两种梭梭群落的重要成员,盖度很大,在春季产量结构中占有重要地位,起着明显的层片作用。另外,在莫索湾地区短命植物常以植物群聚形式出现在裸露基质上,构成群落发展的先锋阶段。     

Hierarchical processes define spatial pattern of avian assemblages restricted and endemic to the arid Karoo, South Africa

Abstract Aim To identify and quantify biotic and abiotic factors associated with the regional gradients in the distribution and abundance of bird communities restricted and endemic to the Succulent and Nama Karoo biomes of South Africa. Location The arid Nama and Succulent Karoo biomes in South Africa. Methods The quarter degree grid cell (QDGC) was used to extract environmental data, while the bird data previously atlased, was linked to the same geo‐referenced system, using a geographical information system (GIS). Bird species were grouped into different life‐history assemblages. A quantitative, systematic analysis of the different bird communities spanning the Karoo was undertaken to examine contributions of broad‐ and local‐scale physical environmental and biotic factors to regional variations in the species composition, using multivariate statistical and spatial analytical tools. These included two indirect gradient methods; principal components analysis (PCA) and detrended correspondence analysis (DCA), and two direct gradient methods; canonical correspondence analysis (CCA) and redundancy analysis (RDA). Results Principal components analysis results showed that the selected environmental variables accounted for about 85% of the variation in the region. The first two principal gradients defined regional temperature seasonality and variability especially in winter and summer. The third principal gradient mainly defined summer rainfall areas in association with the coefficient of variation of rain and regional primary production, while the fourth gradient defined winter rainfall areas, growth days and elements of landscape structure. CCA/RDA analysis produced shortened hierarchically ranked explanatory variables for each bird assemblage. Stepwise gradient analysis results showed summer rain, rainfall concentration, topographic heterogeneity and annual evapotranspiration, as the most important climate variables explaining species occurrence. Landscape, in terms of percentage transformation, morphology, coefficient of variation of primary productivity and distance between suitable habitat patches, were also important, but to a lesser degree. Total variation explained (TVE) by the supplied variables was between 23 and 37% of variation. Less than 20% of TVE was the intrinsic spatial component of environmental influence, indicating that any unmeasured factors were independent of spatial structuring. For all the eight bird assemblages, climate contributed most to TVE (24–57%). Landscape characteristics (human‐induced transformation, vegetation in terms of size if grassy clumps and the average distances between them) contributed theleast to TVE for all the different assemblages (0–6%), especially the granivorous assemblage where it was not significant at all (0%). Seasonal extremes and variability were more important in explaining species gradients than were annual climatic conditions, with the exception of annual potential evapotranspiration. Main conclusions This study was able to synthesize species environment relations at the broad scale and demonstrated the association of arid zone endemic species occurrence with climate extremes and seasonality. Given the predicted climate change scenarios for South Africa, this regional gradient study provides a quantitative ecological basis for finer scale modelling and analysis, developing regional strategies for conserving biodiversity as well as predicting and planning for the effects of global climate change. However, most importantly, it clearly showed that bird species restricted and endemic to the arid Karoo biome may be more sensitive to climate rather than vegetation structure as previously thought.     

Phenological Sensitivity of Early and Late Flowering Species Under Seasonal Warming and Altered Precipitation in a Seminatural Temperate Grassland Ecosystem

Shifts in flowering phenology of plants are indicators of climate change. The great majority of existing phenological studies refer solely to gradual warming. However, knowledge on how flowering phenology responds to changes in seasonal variation of warming and precipitation regimes is missing. We report the onset of 22 early (flowering before/within May) and 23 late flowering (flowering after May) species in response to manipulated seasonal warming (equal to + 1.2°C; last 100-year summer/winter warming), additional winter rainfall, and modified precipitation variability (including a 1000-year extreme drought event followed by heavy rainfall) over the growing season in two consecutive years for a species-rich temperate grassland ecosystem. The average onset of flowering (over 2 years) was significantly advanced 3.1 days by winter warming and 1.5 days by summer warming compared to control. Early flowering species responded to seasonal warming in both years, while late-flowering species responded in only 1 year to summer warming. The average onset of early flowering species was significantly advanced, 4.9 days by winter warming and 2.3 days by summer warming. Species-specific analysis showed that even within the early flowering community there were divergences. A positive correlation between plant height and shift in flowering onset was detected under winter warming (R² = 0.20, p = 0.005). The average onsets of early and late flowering community were affected by neither winter rain nor growing season precipitation variability. Seasonal differences in warming, and particularly winter warming, might alter community dynamics among early and late flowering species which can cause shifts in the seasonal performances of temperate ecosystems.     

Rhizome dynamics and resource storage in Phragmites australis

Seasonal changes in rhizome concentrations of total nonstructural carbohydrates (TNC), water soluble carbohydrates (WSC), and mineral nutrients (N, P and K) were monitored in two Phragmites australis stands in southern Sweden. Rhizome biomass, rhizome length per unit ground area, and specific weight (weight/ length ratio) of the rhizomes were monitored in one of the stands.Rhizome biomass decreased during spring, increased during summer and decreased during winter. However, changes in spring and summer were small (< 500 g DW m-2) compared to the mean rhizome biomass (approximately 3000 g DW m^–2). Winter losses were larger, approximately 1000 g DW m-2, and to a substantial extent involved structural biomass, indicating rhizome mortality. Seasonal changes in rhizome length per unit ground area revealed a rhizome mortality of about 30% during the winter period, and also indicated that an intensive period of formation of new rhizomes occurred in June.Rhizome concentrations of TNC and WSC decreased during the spring, when carbohydrates were translocated to support shoot growth. However, rhizome standing stock of TNC remained large (> 1000 g m^–2). Concentrations and standing stocks of mineral nutrients decreased during spring/ early summer and increased during summer/ fall. Only N, however, showed a pattern consistent with a spring depletion caused by translocation to shoots. This pattern indicates sufficient root uptake of P and K to support spring growth, and supports other evidence that N is generally the limiting mineral nutrient for Phragmites.The biomass data, as well as increased rhizome specific weight and TNC concentrations, clearly suggests that reloading of rhizomes with energy reserves starts in June, not towards the end of the growing season as has been suggested previously. This resource allocation strategy of Phragmites has consequences for vegetation management.Our data indicate that carbohydrate reserves are much larger than needed to support spring growth. We propose that large stores are needed to ensure establishment of spring shoots when deep water or stochastic environmental events, such as high rhizome mortality in winter or loss of spring shoots due to late season frost, increase the demand for reserves.     

Temperature influences on root growth for Encelia farinosa (Asteraceae), Pleuraphis rigida (Poaceae), and Agave deserti (Agavaceae) under current and doubled CO2 concentrations

To help evaluate root distribution patterns, elongation rates of individual roots were measured as a function of soil temperature for Encelia farinosa (a C₃ species), Pleuraphis rigida (C₄), and Agave deserti (CAM), sympatric codominants in the northwestern Sonoran Desert. Measurements were made at current and doubled CO₂ concentrations under winter and summer conditions of air temperature (day/night temperatures of 17 C/10 C and 33 C/22 C, respectively). The three species had different optimal temperatures for root elongation (T_opt) under winter conditions (25 C for E. farinosa, 35 C for P. rigida, and 30 C for A. deserti); T_opt increased by 2-3 C under summer conditions for all three species. The limiting temperatures for elongation also acclimated from winter to summer conditions. The rate of root elongation at T_opt was higher under summer than winter conditions for E. farinosa (9 vs. 6 mm d⁻¹) and P. rigida (20 vs. 14 mm d⁻¹), reflecting conditions for maximum photosynthesis; no difference occurred for A. deserti (9 vs. 10 mm d⁻¹). Decreased elongation rates at extreme temperatures were associated with less cell division and reduced cell extension. The doubled CO₂ concentration increased average daily root elongation rates for A. deserti under both winter (7%) and summer (12%) conditions, reflecting increased cell extension, but had no effect for the other two species. Simulations of root elongation as a function of soil temperatures showed that maximum elongation would occur at different depths (16-20 cm for E. farinosa, 4-8 cm for P. rigida, and 0-4 cm for A. deserti) and during different seasons (winter to spring for E. farinosa, spring to summer for P. rigida, and all year for A. deserti), contributing to their niche separation. Shading of the soil surface moderated daily variations in soil temperature, reducing seasonal root elongation for winter and spring and increasing elongation for summer. Shading also altered root distribution patterns, e.g., optimal rooting depth for A. deserti and especially P. rigida increased for a hot summer day.     

Effects of climatic variation on field metabolism and water relations of desert tortoises

We used the doubly labeled water method to measure the field metabolic rates (FMRs, in kJ kg^?1?day^?1) and water flux rates (WIRs, in ml H₂O?kg^?1?day^?1) of adult desert tortoises (Gopherus agassizii) in three parts of the Mojave Desert in California over a 3.5-year period, in order to develop insights into the physiological responses of this threatened species to climate variation among sites and years. FMR, WIR, and the water economy index (WEI, in ml H₂O?kJ^?1, an indicator of drinking of free water) differed extensively among seasons, among study sites, between sexes, and among years. In high-rainfall years, males had higher FMRs than females. Average daily rates of energy and water use by desert tortoises were extraordinarily variable: 28-fold differences in FMR and 237-fold differences in WIR were measured. Some of this variation was due to seasonal conditions, with rates being low during cold winter months and higher in the warm seasons. However, much of the variation was due to responses to year-to-year variation in rainfall. Annual spring peaks in FMR and WIR were higher in wet years than in drought years. Site differences in seasonal patterns were apparently due to geographic differences in rainfall patterns (more summer rain at eastern Mojave sites). In spring 1992, during an El Niño (ENSO) event, the WEI was greater than the maximal value obtainable from consuming succulent vegetation, indicating copious drinking of rainwater at that time. The physiological and behavioral flexibility of desert tortoises, evident in individuals living at all three study sites, appears central to their ability to survive droughts and benefit from periods of resource abundance. The strong effects of the El Niño (ENSO) weather pattern on tortoise physiology, reproduction, and survival elucidated in this and other studies suggest that local manifestations of global climate events could have a long-term influence on the tortoise populations in the Mojave Desert.