Seedling establishment in a masting desert shrub parallels the pattern for forest trees

The masting phenomenon along with its accompanying suite of seedling adaptive traits has been well studied in forest trees but has rarely been examined in desert shrubs. Blackbrush (Coleogyne ramosissima) is a regionally dominant North American desert shrub whose seeds are produced in mast events and scatter-hoarded by rodents. We followed the fate of seedlings in intact stands vs. small-scale disturbances at four contrasting sites for nine growing seasons following emergence after a mast year. The primary cause of first-year mortality was post-emergence cache excavation and seedling predation, with contrasting impacts at sites with different heteromyid rodent seed predators. Long-term establishment patterns were strongly affected by rodent activity in the weeks following emergence. Survivorship curves generally showed decreased mortality risk with age but differed among sites even after the first year. There were no detectable effects of inter-annual precipitation variability or site climatic differences on survival. Intraspecific competition from conspecific adults had strong impacts on survival and growth, both of which were higher on small-scale disturbances, but similar in openings and under shrub crowns in intact stands. This suggests that adult plants preempted soil resources in the interspaces. Aside from effects on seedling predation, there was little evidence for facilitation or interference beneath adult plant crowns. Plants in intact stands were still small and clearly juvenile after nine years, showing that blackbrush forms cohorts of suppressed plants similar to the seedling banks of closed forests. Seedling banks function in the absence of a persistent seed bank in replacement after adult plant death (gap formation), which is temporally uncoupled from masting and associated recruitment events. This study demonstrates that the seedling establishment syndrome associated with masting has evolved in desert shrublands as well as in forests.     

Mycorrhizal symbiosis increases growth,reproduction and recruitment of Abutilon theophrasti Medic. in the field

We examined in the field the effect of the vesicular-arbuscular (VA) mycorhizal symbiosis on the reproductive success of Abutilon theophrasti Medic., an early successional annual member of the Malvaceae. Mycorrhizal infection greatly enhanced vegetative growth, and flower, fruit and seed production, resulting in significantly greater recruitment the following year. In addition, the seeds produced by mycorrhizal plants were significantly larger and contained significantly more phosphorus than seeds from non-mycorrhizal plants, an effect which may improve offspring vigor. Infection by mycorrhizal fungi may thus contribute to the overall fitness of a host plant and strongly influence long-term plant population dynamics.     

Nitrogen and phosphorus dynamics in hot desert streams of Southwestern U.S.A.

Nitrogen to phosphorus ratios and concentrations of nitrate and soluble reactive phosphate are presented for an array of Southwestern streams as evidence that nitrogen is the limiting nutrient where such limitation occurs. Nitrate uptake in sections of intermittent streams was attributable to autotrophic activity. Uptake of soluble reactive phosphate was unrelated to any indicator of autotrophic activity, thus concentrations of this nutrient in desert and semi-desert stream waters may be controlled by other factors.     

Geigeria alata in the Namib desert: seed heteromorphism in an extremely arid environment

Abstract. Seed heteromorphism in annual Geigeria alata in the Namib Desert was investigated in relation to environmental conditions. Geigeria alata shows heteromorphism in seed dispersal distance depending on the position of the seeds on the parent plant. Seeds produced in capitula on branches (aerial) travel long distances, while seeds produced at the base of the plant remain in the vicinity of the parent plant. The ratio of aerial to basal seed production was investigated under different environmental conditions following an aridity gradient. Although there were significant differences in seed and capitula production between seasons and between different positions on the plant, no overall trend was found with increasing aridity.     

Nitrogen enrichment of host plants has mostly beneficial effects on the life-history traits of nettle-feeding butterflies

Butterflies rank among the most threatened animal groups throughout Europe. However, current population trends differ among species. The nettle-feeding butterflies Aglais io and Aglais urticae cope successfully with the anthropogenic land-use change. Both species are assumed to be pre-adapted to higher nitrogen contents in their host plant, stinging nettle (Urtica dioica). However, it is currently unknown, whether this pre-adaptation enables both Aglais species to cope successfully or even to benefit from the excessive nitrogen availabilities in nettles growing in modern farmlands. For this reason, this study focused on the response of both Aglais species to unfertilized nettles compared to nettles receiving 150 or 300 kg N ha⁻¹ yr⁻¹ (i.e., common fertilizer quantities of modern-day agriculture). Fertilized nettles were characterized by higher nitrogen concentrations and lower C:N ratios compared to the control group. In both Aglais species, the individuals feeding on fertilized nettles had higher survival rates, shorter larval periods and heavier pupae and, in A. urticae also longer forewings. All these trait shifts are beneficial for the individuals, lowering their risk to die before reproduction and increasing their reproductive potential. These responses agree with the well-accepted nitrogen-limitation hypothesis predicting a positive relationship between the nitrogen content of the diet and the performance of herbivorous insects. Furthermore, our findings suggest that the increasing abundance of both Aglais species may result not only from the increasing spread of nettles into the farmland but also from changes in their quality due to the eutrophication of the landscape during recent decades.     

Size-related flowering and fecundity in the tropical canopy tree species, Shorea acuminata (Dipterocarpaceae) during two consecutive general flowerings

We monitored the reproductive status of all trees with diameters at breast height (dbh) >30 cm in a 40-ha plot at Pasoh, west Malaysia, and investigated the individual fecundity of 15 Shorea acuminata Dyer (Dipterocarpaceae) trees using seed-trapping methods during two consecutive general flowering periods in 2001 (GF2001) and 2002 (GF2002). The proportion of flowering trees was higher, and not dependent on size, in GF2002 (84.2%), than in GF2001 (54.5%), when flowering mainly occurred in trees with a dbh ≤70 cm. Fecundity parameters of individual trees per event varied widely (221,000–35,200,000 flowers, 0–139,000 mature seeds, and 1.04–177 kg total dry matter mass of fruit (TDM) per tree). Monotonic increases with increasing tree size were observed for flower production and TDM amongst trees up to 90 cm in dbh, but not for mature seed production or for any of these parameters amongst larger trees. The pattern of reproductive investment during the two consecutive reproductive events clearly differed between medium-sized and large trees; the former concentrated their reproductive investment in one of the reproductive events whereas the latter allocated their investment more evenly to both reproductive events. Our results suggest size-related differences in the resource allocation pattern for reproduction. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

氮磷添加对荒漠草原植物-凋落物-土壤生态化学计量特征的影响

为明确植物、凋落物和土壤养分含量及化学计量比对土壤中添加多种限制性养分的响应,阐明“植物-凋落物-土壤”连续体化学计量动态及各组分之间的协同作用,以宁夏荒漠草原为研究对象,于2018年开始进行氮(N)、磷(P)养分添加控制试验。试验处理包括对照(CK)、N添加、P添加、NP共同添加4个处理。结果表明：(1)NP共同添加显著增加了荒漠草原植物N和P含量、以及凋落物和土壤P含量,显著降低了荒漠草原植物C∶N和C∶P、以及土壤和凋落物C∶P和N∶P。P添加显著增加了荒漠草原植物、凋落物和土壤P含量,显著降低了植物、凋落物、土壤C∶P和N∶P。N添加分别增加了植物、凋落物N含量和N∶P,但对植物N含量影响未达到显著水平。(2)C、N、P含量和N∶P大小均表现为植物>凋落物>土壤,C∶N和C∶P均表现为凋落物>植物。(3)N添加提高了荒漠草原植物对P再吸收效率,降低了荒漠草原植物对N利用效率;P添加提高荒漠草原植物对N再吸收效率,降低荒漠草原对P的利用效率;NP共同添加提高了荒漠草原植物对N和P再吸收效率,降低了荒漠草原植物对N和P利用效率。(4)植物-凋落物-土壤的N、P含量...     

Dynamics of cohorts of cladodes and related effects on reproduction in the shrub Retama sphaerocarpa in semi-arid south-eastern Spain

Changes in the size and age structure of the canopy of the leguminous shrub Retama sphaerocarpa in semi-arid south-eastern Spain were investigated by monitoring growth and survivorship of cladodes (photosynthetic stems) on marked shoots over a period of 26 months. Three basic morphological types of cladodes – long shoots, short shoots, and secondary short shoots – were distinguished.The canopy of the shrubs contained three annual cohorts of cladodes in spring and summer. The number and size of shoots produced each year was highly variable and was apparently related to the amount of rainfall during the preceding cool season. The spring cohort of 1994 produced only 37% of cladodes compared with 1993. Two cohorts of shoots were actually initiated in spring and late summer of each year, but the second cohort produced only 2–12% of the number of cladodes compared with the spring cohort. The proportions of the different cohorts in the canopy changed rapidly from April to August, but only slowly during the remainder of the year when only two annual cohorts remained after extensive litterfall in late summer. This late summer litterfall caused a substantial reduction in green canopy area (40–50%) which was achieved mainly be shedding of one year old cladodes. The life expectancy of cladodes decreased with increasing order of their morphological type from 850 ± 25 days in long shoots to 563 ± 4 days and 546 ± 9 days in short shoots and second order short shoots, respectively.Flowering and fruiting took place from May to July, almost exclusively on one year old cladodes, and coincided with the maximum development of the canopy. Flowering intensity was high in 1994, when individual shoots supported a mean number of approximately 150 flowers. Shoots produced an average of 12.6 ± 0.6 and 5.3 ± 1.0 fruits per shoot in 1993 and 1994, respectively. Most of the annual fruit crop (80–90%) was shed during litterfall in late summer. A proportion of 10–20% was retained in the canopy for up to 12 months, however, with some fruits persisting for more than 22 months.     

Small mammal seed consumption in the Karoo,South Africa: further evidence for divergence in desert biotic processes

Summary Annual seed consumption by a small mammal community in the semi-arid Karoo, South Africa, was calculated using population, biomass and dietary estimates and published estimates of field metabolic rate. The community comprised a macroscelid elephant shrew Macroscelides proboscideus, and two rodents Gerbillurus paeba and Desmodillus auricularis. Only G. paeba consumed seed during the study, and then in low amounts (annual mean = 0.4% of diet, insects = 53%, herbage = 47%), with M. proboscideus and D. auriocularis consuming mainly insects (88%) and herbage (90%), respectively. Energy requirements for the G. paeba population averaged 568 kJ ha^–1 day^–1. Total seed consumption was estimated to be 140 g ha^–1, or 155 000 seeds ha^–1 which represents 0.5% of the annual seed production for this site. These levels of granivory by small mammals are significantly lower than those recorded for North American deserts, and raises the question as to why granivory varies between these systems. These findings also refute the hypothesis that as a result of the harshness and similarity of physical conditions in deserts, the properties of desert communities are convergent.     

Effects of within-plant Variation in Leaf Quality on the Ecology of G. viridula

ABSTRACT In order to find out the relationship between the leaf quality of R. obtusifolius and the ecology of G. viridula , a series of experiments was carried out. The effects of leaf age on rates of growth of instars of G. viridula showed that the leaf age was closely related to the development of the larvae of G. viridula . Also, the results showed quite clearly that the N concentration of R. obtusifolius leaves have significant effects on the growth rates of G. viridula larvae. It is suggested that C: N ratio of food plant plays an important role in the population dynamics of phytophagous insects. A small increase in leaf N concentration in a nutrient poor habitat will substantially increase the G. viridula population.     

阔叶红松林不同演替阶段灌木器官的N、P分配特征

氮（N）、磷（P）是植物生长发育重要的限制元素,N、P以及N、P间的关系是化学计量学研究的重要内容。植物不同器官对N、P的需求存在差异,因此N、P的分配与权衡对植物的生长发育起到重要的调节作用。为了探究阔叶红松（Pinus koraiensis）林不同演替阶段主要灌木器官的N、P化学计量特征及其分配格局,了解灌木的养分限制因子及其分配策略,在黑龙江凉水国家级自然保护区的阔叶红松林四个演替阶段——白桦（Betula platyphylla）次生林、阔叶混交林、针阔混交林和阔叶红松林内,选取主要灌木种,分析了叶片、枝条、茎干和根的N、P化学计量特征以及异速生长关系。结果发现：灌木不同器官之间的N、P含量差异显著,其中叶的N、P含量均为最高,茎干的N、P含量均为最低。所有器官中,叶的N ： P最高,为7.59;根的N ： P最低,为5.47。灌木叶、茎干、根之间的N ： P存在显著差异,而茎干、根与枝条之间的N ： P没有显著差异。阔叶红松林不同演替阶段对灌木同一器官N、P化学计量的影响不同,灌木叶片的N含量在不同演替中具有显著差异,其他器官的差异不显著;不同演替阶段灌木器官的P含量均无显著差异。在不同演替阶段中各器官的N ： P均小于14,说明N元素是阔叶红松林各演替阶段灌木生长的限制因子,而P对灌木的生长没有起到限制作用。研究表明在不同演替阶段中,N、P的异速生长关系普遍存在于灌木器官中,呈现显著的正相关关系;整体上各器官之间的斜率没有显著差异,说明灌木采取保守分配策略,以此来适应环境和满足自身生长发育的需要。     

Nitrogen supply effects on leaf dynamics and nutrient input into the soil of plant species in a sub-arctic tundra ecosystem

Global warming will lead to increased nitrogen supply in tundra ecosystems. How increased N supply affected leaf production, leaf turnover and dead leaf N input into the soil of Empetrum nigrum and Andromeda polifolia (evergreens), Eriophorum vaginatum (graminoid) and Betula nana (deciduous) in a sub-arctic tundra in northern Sweden between 2003 and 2007 was experimentally investigated. There was considerable interspecific variation in the response of leaf production to N addition, varying from negative, no response to a positive response. Nitrogen addition effects on leaf turnover also showed considerable variation among species, varying from no effect to increased leaf turnover (up to 27% in Eriophorum). Nitrogen addition resulted in a four to fivefold increase in N content in the dead leaves of both evergreens and a 65% increase in Eriophorum. Surprisingly, there was no increase in Betula. The response of dead leaf P contents to N addition was rather species specific. There was no response in Empetrum, whereas there were significant increases in Andromeda (+214%) and Eriophorum (+32%), and a decrease of 47% in Betula. As an overall result of the changes in leaf production, leaf turnover and dead leaf N and P contents, nitrogen addition increased in all species except Betula the amount of N and, for Andromeda and Eriophorum the amount of P transferred to the soil due to leaf litter inputs. However, the way in which this was achieved differed substantially among species due to interspecific differences in the response of the component processes (leaf production, leaf turnover, dead leaf nutrient content).     

Changes in nitrogen resorption traits of six temperate grassland species along a multi-level N addition gradient

Nitrogen (N) resorption from senescing leaves is an important mechanism of N conservation for terrestrial plant species, but changes in N-resorption traits over wide-range and multi-level N addition gradients have not been well characterized. Here, a 3-year N addition experiment was conducted to determine the effects of N addition on N resorption of six temperate grassland species belonging to three different life-forms: Stipa krylovii Roshev. (grass), Cleistogenes squarrosa (T.) Keng (grass), Artemisia frigida Willd. (semishrub), Melissitus ruthenica C.W.Wang (semishrub and N-fixer), Potentilla acaulis L. (forb) and Allium bidentatum Fisch.ex Prokh. (forb). Generally, N concentrations in green leaves increased asymptotically for all species. N concentrations in senescent leaves for most species (5/6) also increased asymptotically, except that the N concentration in senescent leaves of A. bidentatum was independent of N addition. N-resorption efficiency decreased with increasing N addition level only for S. krylovii and A. frigida, while no clear responses were found for other species. These results suggest that long-term N fertilization increased N uptake and decreased N-resorption proficiency, but the effects on N-resorption efficiency were species-specific for different temperate grassland species in northern China. These inter-specific differences in N resorption may influence the positive feedback between species dominance and N availability and thus soil N cycling in the grassland ecosystem in this region.     

The role of vegetation–microclimate feedback in promoting shrub encroachment in the northern Chihuahuan desert

Many arid and semi‐arid landscapes around the world are affected by a shift from grassland to shrubland vegetation, presumably induced by climate warming, increasing atmospheric CO₂ concentrations, and/or changing land use. This major change in vegetation cover is likely sustained by positive feedbacks with the physical environment. Recent research has focused on a feedback with microclimate, whereby cold intolerant shrubs increase the minimum nocturnal temperatures in their surroundings. Despite the rich literature on the impact of land cover change on local climate conditions, changes in microclimate resulting from shrub expansion into desert grasslands have remained poorly investigated. It is unclear to what extent such a feedback can affect the maximum extent of shrub expansion and the configuration of a stable encroachment front. Here, we focus on the case of the northern Chihuahuan desert, where creosotebush (Larrea tridentata) has been replacing grasslands over the past 100–150 years. We use a process‐based coupled atmosphere‐vegetation model to investigate the role of this feedback in sustaining shrub encroachment in the region. Simulations indicate that the feedback allows juvenile shrubs to establish in the grassland during average years and, once established, reduce their vulnerability to freeze‐induced mortality by creating a warmer microclimate. Such a feedback is crucial in extreme cold winters as it may reduce shrub mortality. We identify the existence of a critical zone in the surroundings of the encroachment front, in which vegetation dynamics are bistable: in this zone, vegetation can be stable both as grassland and as shrubland. The existence of these alternative stable states explains why in most cases the shift from grass to shrub cover is found to be abrupt and often difficult to revert.     

Responses of leaf C:N:P stoichiometry to water supply in the desert shrub Zygophyllum xanthoxylum

• Based on the elemental composition of major biochemical molecules associated with different biological functions, the ‘growth rate hypothesis’ proposed that organisms with a higher growth rate would be coupled to lower C:N, especially lower C:P and N:P ratios. However, the applicability of the growth rate hypothesis for plants is unclear, especially for shrubs growing under different water supply.
• We performed an experiment with eight soil moisture levels (soil water content: 4%, 6%, 8%, 13%, 18%, 23%, 26% and 28%) to evaluate the effects of water availability on leaf C:N:P stoichiometry in the shrub Zygophyllum xanthoxylum.
• We found that leaves grew slowly and favored accumulation of P over C and N under both high and low water supply. Thus, leaf C:P and N:P ratios were unimodally related to soil water content, in parallel with individual leaf area and mass. As a result, there were significant positive correlations between leaf C:P and N:P with leaf growth (u).
• Our result that slower‐growing leaves had lower C:P and N:P ratios does not support the growth rate hypothesis, which predicted a negative association of N:P ratio with growth rate, but it is consistent with recent theoretical derivations of growth–stoichiometry relations in plants, where N:P ratio is predicted to increase with increasing growth for very low growth rates, suggesting leaf growth limitation by C and N rather than P for drought and water saturation.

     

Nitrogen addition increases intraspecific competition in the invasive wetland plant Alternanthera philoxeroides,but not in its native congener Alternanthera sessilis

Nitrogen is often released in pulses with different frequencies, and N supply pulses may affect growth, reproduction, and biomass allocation of plants. However, few studies have examined how N supply pulses affect intraspecific competition of clonal plants and whether such an effect depends on the N supply amount. We grew one (no competition) or 12 ramets (with intraspecific competition) of both an invasive clonal plant Alternanthera philoxeroides and its native congener Alternanthera sessilis in five different N treatments: control (no N addition), low/high amount with low/high frequencies (pulses). Nitrogen addition significantly increased the growth of both species, while intraspecific competition decreased it. Nitrogen addition significantly increased intraspecific competitive intensity of A. philoxeroides as measured by the log response ratio of growth traits, but did not affect that of A. sessilis. Despite the N supply amount, N pulses had little effect on the growth and thus intraspecific competition of the two species. Therefore, increasing N deposition may change population structure and dynamics and the invasion succession of A. philoxeroides, but changes in N pulses may not.     

Differential responses of the floating-leaved aquatic plant Nymphoides peltata to gradual versus rapid increases in water levels

We compared the growth responses of the floating-leaved species Nymphoides peltata to gradual and rapid rising water levels under two nutrient concentrations (1 g and 12 g of slow released fertilizer (N-P-K: 16-8-12) per container filled with 8 kg washed sand), and predicted the population expansion after these floods. The results showed that the capacity for petiole elongation was dependent on leaf age, and only leaves that were no more than five days old had the capability to reach the water surface when the water level increased rapidly from 50 cm to 300 cm. Plants subjected to a gradual rising water level tracked the increase in water depth whose petioles elongated at 3.96 ± 1.70 cm per day and 4.80 ± 0.16 cm per day under low and high nutrient concentrations respectively throughout the experiment period. When water levels were rapidly raised, leaf petioles elongated rapidly at 25.48 ± 1.51 cm per day and 26.64 ± 2.24 cm per day under low and high nutrient concentrations respectively during the first ten days. Plants under a constant water level maintained highest mean leaf recruitment (mean 3.0 ± 0.33 leaves and 24.4 ± 5.87 leaves every ten days under low and high nutrient concentrations, respectively). Therefore, more young leaves existed in the canopy ensuring that when the water level increases, young leaves can rapidly emerge after submergence. Gradual water level rise did not significantly affect biomass and ramet production (4.75 ± 1.41 g and 5.50 ± 1.22 ramets in low nutrient; 48.49 ± 21.45 g and 35.67 ± 11.78 ramets in high nutrient), but rapid water level rise negatively affected ramet production in both nutrient concentrations (3.00 ± 1.26 ramets and 11.25 ± 4.19 ramets in low and high nutrients, respectively). The results indicated that continual leaf recruitment and rapid petiole elongation were both important ways in which N. peltata adapted to increasing water levels. Extreme flooding may be a disturbance factor that affects plant growth and the population expansion of N. peltata, while small gradual water level rise should not harm this species.     

华北太行山前平原农田生态系统中氮、磷、钾循环与平衡研究

以栾城县为例,分析了华北太行山前平原农田生态系统N、P、K三要素养分循环与平衡特征及其变化趋势,评价了农田养分平衡状况,提出了合理施肥对策。结果表明,1985～2000年栾城县农田养分平衡中,N素经历了由轻微赤字向盈余的转变过程,由1985年的1．4％赤字转变到2000年盈余48．6％;K由极度亏缺(82％赤字),逐渐向平衡发展,到2000年已有4．6％的轻微盈余,但仍有大量田块是赤字平衡;P则始终有大量盈余。目前农田养分收支存在一定程度的N、P投入过量,K投入不足的问题,以土壤肥力状况和作物种植结构而论,栾城县应采取“稳氮、控磷、增钾”的施肥对策。避免过多的盈余化肥氮进入环境,同时应继续推广秸秆还田技术,提高养分循环再利用效率。