AMPHIPLOIDY AND INTROGRESSION IN STIPA

Johnson , B. Lennart . (U. California, Los Angeles.) Amphiploidy and introgression in Stipa. Amer. Jour. Bot. 49(3): 253–262. Illus. 1962.—Frequent specimens heretofore referred to Stipa californica (2n = 36) were found to be similar to an undescribed polyploid (2n = 68) species of Stipa on the ratio of lemma to palea length. On other individual diagnostic characters, S. californica and the polyploid were not separable. However, on the discriminant function (Z) comprising 5 such attributes, their mean difference was highly significant. The polyploid is given the new species name, Stipa nevadensis . Appropriate tests using the function Z excluded S. californica as a possible parent of the polyploid and pointed to S. lettermani (2n = 32) and the combined species S. occidentalis (2n = 36) and S. elmeri (2n = 36) as the putative parents. The last 2 species were barely separable on multigenic differences measured by the discriminant function, and the mean Z for S. californica fell near the mid point between them and S. columbiana (2n = 36). This fact together with evidence from the frequency distribution of Z suggested a hybrid origin for S. californica and introgression with its putative parents.     

内蒙古地带性针茅植物对CO2和气候变化的适应性研究进展

收集了1992—2013年关于模拟CO2浓度升高及气候变化(温度升高、降水变化)对内蒙古地带性草原群落的5个建群种针茅植物(贝加尔针茅、本氏针茅、大针茅、克氏针茅、短花针茅)影响的实验研究结果表明,模拟CO2浓度升高、增温和增雨将提高针茅植物的光合作用和株高生长,但CO2处理时间延长会导致光合适应;温度和降雨变化将改变针茅植物的物候进程,但物种之间反应有差异;CO2浓度升高有助于针茅植物生物量增加,增温和干旱则相反,CO2浓度升高对干旱的影响具有补偿作用;干旱和涝渍胁迫将提高针茅植物植株C/N,CO2浓度升高将加剧水分胁迫下针茅植物植株C/N的增加效应,导致牧草品质下降。由于当前在适应性指标、针茅植物对气候变化协同作用的适应机理及其敏感性研究等方面存在的不足,导致目前无法全面比较各针茅植物对CO2和温度、降水变化的响应差异及其敏感性,因而无法预测未来在全球变化背景下,这几种针茅植物的动态变化及其在地理分布上的迁移替代规律。为科学应对气候变化,未来应加强内蒙古地带性针茅植物的适应性指标、针茅植物对多因子协同作用的适应机理及敏感性研究。     

Stipa valdemonensis (Poaceae), a new species from Sicily

Abstract

A new species of Stipa, endemic to Sicily, named Stipa valdemonensis is described here. The new taxon is related to Stipa crassiculmis. Owing to the small number of individuals observed, in few restricted localities only, it is assigned the IUCN threat status “vulnerable”.     

NATURAL HYBRIDS BETWEEN ORYZOPSIS AND STIPA. III. ORYZOPSIS HYMENOIDES × STIPA PINETORUM

Johnson, B. Lennart. (U. California, Los Angeles.) Natural hybrids between Oryzopsis and Stipa. III. Oryzopsis hymenoides × Stipa pinetorum. Amer. Jour. Bot. 50(3): 228–234. Illus. 1963.—On the basis of morphological characters, the spontaneous hybrid Oryzopsis hymenoides × Stipa pinetorum is included in 0. bloomeri which consists of a number of sterile, natural hybrids between O. hymenoides (2n = 48) and various American species of Stipa (2n = 32-82). While intermediate between its parents in many attributes, the pinetorum hybrid is different from the other hybrids included in O. bloomeri with respect to lemma-hair length and other characters which are diagnostic for S. pinetorum (2n = 32). The hybrid has 2n = 40 chromosomes, but some plants of the hybrid and some of O. hymenoides had small supernumerary, somatic chromosomes. The parents formed only bivalents at meiosis. The hybrid showed some affinity among 14 chromosomes per sporocyte. This affinity is nearly as great as that shown by other Stiporyzopsis hybrids with 56 or 58 chromosomes, and is consistent with the earlier suggestion that bivalent formation in polyploid species of Stipa may be gene-controlled.     

Comparative Physiology and Growth of Two Perennial Tussock Grass Species in a Semi-Arid Environment

The physiology and growth ofLygeum spartumandStipa tenacissima,two perennial tussock grasses which dominate wide areas of semi-aridsouth-eastern Spain, were compared at times of high and lowavailability of water, in autumn and summer respectively, tostudy the adaptation of this growth form to arid environments.The two species differed in morphological and physiologicaltraits.Stipatussocks were larger and had opportunistic growth,andStipaleaves had a smaller specific area and lower diffusiveconductance to water vapour. The two grasses were similar inmaximum photosynthetic rate, leaf nitrogen concentration, andin the response to high light conditions but had different tissuewater relations.Stipawas better suited to cope with droughtand erratic rainfall, because of its more effective controlof water loss and its growth patterns.Lygeumseems to be adaptedto less dry conditions and more saline soils. The tussock growthform provides an adaptive advantage in these infertile environmentsby reducing radiation absorbance. leaf extension; Lygeum spartum; perennial grassland; semi-arid environments; Stipa tenacissima; tussock grass; water relations; Adaptation; growth; photosynthesis     

Differences in the compensatory growth of two co-occurring grass species in relation to water availability

We compared the potential for compensatory growth of two grass species from the Mongolian steppe that differ in their ability to persist under grazing: the rhizomatous Leymus chinensis and the caespitose Stipa krylovii, and investigated how this ability might be affected by drought. Plants were grown in a greenhouse under wet and dry conditions and subjected to a clipping treatment (biweekly removal of 75–90% of the aerial mass). Leymus exhibited a much stronger compensatory growth after clipping than Stipa. Leymus showed a significant increase in its relative growth rate (RGR) after clipping, while for Stipa RGR was negatively affected. Clipped Leymus plants maintained leaf productivity levels that were similar to undamaged individuals, while leaf-productivity in clipped Stipa dropped to less than half of that of the controls. In Leymus, there was less compensatory growth under dry than under wet conditions, while in Stipa the compensation was increased under drought. This difference probably reflects the fact that Stipa is more drought-tolerant than Leymus. The greater compensatory growth of Leymus compared to Stipa mainly resulted from a greater stimulation of its net assimilation rate (NAR), and its greater capacity to store and reallocate carbohydrates by clipping. The greater increase in NAR was probably the result of a stronger reduction in self-shading, because Leymus shoots were much denser than those of Stipa, which resulted in a higher increase in light penetration to remaining leaves after clipping. The results of this study suggest that the greater ability of Leymus to persist under grazing is the result of its larger capacity for compensatory growth.     

Germination ecology of Central Asian Stipa spp: differences among species, seed provenances, and the importance of field studies

Stipa-species are wide-spread in Central Asia, but sexual reproduction in the dry steppes is rare. To facilitate conservation and restoration of these important rangelands, we studied germination characteristics of three common Mongolian Stipa-species under field- and lab conditions. Seeds of Stipa krylovii, Stipa gobica and Stipa glareosa were sown at the study site in Southern Mongolia over two consecutive years during which period tests were carried out to ascertain whether competition or herbivory are the main constraints of seedling establishment. In addition, we tested germination and seed viability in the laboratory under two different temperature regimes (20/10°C and 8/4°C), as well as the effect of cold-stratification. The lab experiments also included S. krylovii seeds originating from three climatically different provenances. None of the three Stipa-species seedlings emerged during the first 2 years of the field study. However, after an unusually intense rain event in the third year, 3% of S. krylovii, 0.6% of S. glareosa and 0.1% of S. gobica seeds germinated in the study plots. The factors ‘sowing-year’ and ‘vegetation’ significantly affected seedling emergence, whereas grazing had no effect at all. Under laboratory conditions a high percentage of viable seeds of S. gobica and S. glareosa germinated at both incubation temperatures, and cold-stratification had no effect on germination or viability. Germination of S. krylovii seeds required warmer temperatures and cold-stratification had a positive effect. Such evidence for dormancy was more pronounced in seeds from the moister, northern provenances. Germination of Stipa-species in the field is rare and only possible under exceptionally moist conditions. Conservation should thus concentrate on steppe conservation rather than on restoration. Where artificial reseeding is necessary, differences among species and also among different seed provenances should be taken into account.     

EXPERIMENTAL EVIDENCE FOR GENETIC VARIATION IN COMPATIBILITY BETWEEN THE FUNGUS ATKINSONELLA HYPOXYLON AND ITS THREE HOST GRASSES

Variation in compatibility has been documented within and among several natural plant populations infected by fungal pathogens. In this study, seeds and isolates of the fungus Atkinsonella hypoxylon (Ascomycetes, Clavicipitaceae) were collected from three populations of the grass Danthonia spicata, two populations of D. compressa, and four populations of Stipa leucotricha. Each fungal strain was reciprocally inoculated into seedlings grown in aseptic culture from its original host population, into seedlings from other conspecific host populations, and into seedlings from the other two host species. There were three distinct patterns of compatibility, as evidenced by the ability of the fungus to grow on the seedling and to colonize new tillers. Fungal strains from one host genus were incompatible with seedlings from the other host genus. Strains from the two Danthonia species were broadly compatible among Danthonia populations and had very high rates of infection, while strains from Stipa also were broadly compatible among Stipa populations but had relatively low rates of infection. Literature surveys indicate that, in contrast to pathogenic microorganisms, mutualistic microorganisms typically exhibit broad patterns of compatibility among hosts, which lack resistance to infection. The effect of A. hypoxylon on host fitness is most detrimental in Stipa, where the fewest seedlings became infected, and most beneficial in Danthonia, where most seedlings became infected.     

半干旱草地长期封育进程中针茅植物根系格局变化特征

以云雾山不同封育年限草地针茅植物根系和土壤为研究对象,对其根系特征、土壤特性及两者关系进行研究,以探讨分析封育对针茅根系格局的影响。结果表明:(1)针茅植物根系生物量、根长密度、根表面积和根体积在封育初期轻微下降,之后缓慢上升,并在封育30 a草地得到显著增加。(2)随封育年限增加,各根系指标在3种针茅物种间的组成格局具有类似变化规律,具体表现为:长芒草在放牧草地所占比例最高,之后逐渐降低,并在封育30 a草地消失;大针茅所占比例呈先升后降变化规律,并在封育22 a草地达到最大值;甘青针茅仅出现于封育30 a草地,且占据优势地位。(3)大针茅和甘青针茅0—0.6 mm径级根系比例高于大针茅,使其根系直径显著低于大针茅,比根长和比根面积显著高于大针茅;此外,长芒草根组织密度显著高于长芒草和甘青针茅。(4)长期封育在显著提高土壤水分、养分含量和土壤氮磷比的同时显著降低土壤碳氮比,但对微生物生物量碳、氮无明显影响。(5)针茅根系特征与土壤指标的关联性分析显示针茅根系受土壤氮资源的显著影响。     

Bioclimatic regions influence genetic structure of four Jordanian Stipa species

Strong environmental gradients can affect the genetic structure of plant populations, but little is known as to whether closely related species respond similarly or idiosyncratically to ecogeographic variation. We analysed the extent to which gradients in temperature and rainfall shape the genetic structure of four Stipa species in four bioclimatic regions in Jordan. Genetic diversity, differentiation and structure of Stipa species were investigated using amplified fragment length polymorphism (AFLP) molecular markers. For each of the four study species, we sampled 120 individuals from ten populations situated in distinct bioclimatic regions and assessed the degree of genetic diversity and genetic differentiation within and among populations. The widespread ruderals Stipa capensis and S. parviflora had higher genetic diversity than the geographically restricted semi‐desert species S. arabica and S. lagascae. In three of the four species, genetic diversity strongly decreased with precipitation, while genetic diversity increased with temperature in S. capensis. Most genetic diversity resided among populations in the semi‐desert species (Φ_ST = 0.572/0.595 in S. arabica/lagascae) but within populations in the ruderal species (Φ_ST = 0.355/0.387 S. capensis/parviflora). Principal coordinate analysis ( PCoA) and STRUCTURE analysis showed that Stipa populations of all species clustered ecogeographically. A genome scan revealed that divergent selection at particular AFLP loci contributed to genetic differentiation. Irrespective of their different life histories, Stipa species responded similarly to the bioclimatic gradient in Jordan. We conclude that, in addition to predominant random processes, steep climatic gradients might shape the genetic structure of plant populations.     

Temporal dynamics of herbivory and water availability interactively modulate the outcome of a grass–shrub interaction in a semi‐arid ecosystem

The study of plant–plant interactions along grazing and abiotic stress gradients is a major research topic in plant ecology, but the joint effects of both stressors on the outcome of plant–plant interactions remains poorly understood. We used two different factorial experiments conducted in a semi‐arid Mediterranean steppe to assess: 1) the role of the perennial grass Stipa tenacissima, a low‐palatability species, providing protection from rabbit herbivory to the shrub Retama sphaerocarpa (experiment 1), and 2) the effects of environmental amelioration provided by Stipa on the recovery of Retama after rabbit damage under two contrasted levels of water availability (experiment 2). In the experiment 1, water stress worked as an indirect modulator of herbivore protection by Stipa. This species protected Retama seedlings from rabbit herbivory during the wetter conditions of spring and winter, but this effect dissapeared when rabbit pressure on Retama increased during summer drought due to the decrease in alternative food resources. In the experiment 2, Stipa exerted a negative effect on the survival of Retama seedlings during the three years of the experiment, regardless of inter‐annual differences in rainfall or the watering level applied. This negative effect was mainly due to excessive shading. However, Stipa increased Retama recovery after initial rabbit impact, overriding in part this negative shade effect. Conversely, Stipa impact on the Fv/Fm of Retama seedlings depended on the intra‐annual water dynamics and its experimental manipulation, overall contradicting predictions from the stress–gradient hypothesis. The complex interactions found between herbivory, microclimatic amelioration from Stipa, and water availability as drivers of Retama performance illustrate the importance of considering the temporal dynamics of both biotic and abiotic stressors to fully understand the outcome of plant–plant interactions.     

温带草原7种针茅植物根系特征及其对环境因子变化的适应

为比较针茅(Stipa)植物适应策略,以大尺度梯度下(>1600 km)温带草原主要针茅植物为对象,系统研究了8个样点7种针茅根系生物量、根冠比、解剖结构和生理调节物质的差异及其对环境因子的适应。在由东北至西南的区域上,随降水量下降针茅植物根系的抗旱特征增强或适应策略趋于复杂,不同针茅植物根系对水分变化(或旱季和雨季)有着不同的适应策略。综合分析表明贝加尔针茅(S.baicalensis)、大针茅(S.grandis)及克氏针茅(S.krylovii)(多伦样点)的生长受干旱制约,对降水高度敏感,雨季降水促进其生物量快速积累。沙生针茅(S.glareosa)、短花针茅(S.breviflora)、戈壁针茅(S.gobica)、本氏针茅(S.bungeana)等通过增大根冠比和渗透调节物质累积等途径提高根系吸水和保水能力,抵御干旱胁迫。偏相关分析显示实验区域针茅植物根系性状与降水量和海拔高度存在显著的相关性。     

Stipa narynica sp. nov. (Poaceae) from the western Tian‐Shan Mountains

Stipa narynica M. Nobis sp. nov. from the western Tian‐Shan Mts (western Kyrgyzstan) is described and illustrated. The new species belongs to sect. Smirnovia Tzvel. Morphologically the species is similar to S. aktauensis, a species from which it is easily distinguished by its longer glumes (41–)44–55(–62) mm vs 35–45 mm, longer anthecium (11.5–) 12.0–14.2(–14.8) mm vs 9.5–11.2(–11.7) mm, longer hairs on the seta (5.5–)6.0–8.0(–8.5) mm vs (3.0–)3.8–5.5(–6.0) mm and by the top of the lemma which in S. narynica is either glabrous or with a poorly developed ring of short hairs, while in S. aktauensis, the top of lemma is always distinctly and densely pilose. The lemma and leaf blade structure of both species were examined by means of SEM. The main features differentiating S. narynica and S. aktauensis and distribution of the taxa are presented. An original key to the middle Asiatic species of the sect. Smirnovia, the most similar to Stipa narynica is provided.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

Nutrient resorption from senescing leaves in two Stipa species native to central Argentina

Abstract Nutrient resorption from senescing leaves enables plants to conserve and reuse nutrients. As such, it could be expected that plant species adapted to infertile soils have a higher nutrient resorption efficiency (percentage reduction of nutrients between green and senesced leaves) and/or higher nutrient resorption proficiency (absolute reduction of nutrients in senesced leaves) than those adapted to fertile soils. Our objective was to compare nitrogen (N) and phosphorous (P) resorption of two congener grasses that successfully occupy uplands of relatively low fertility (Stipa gynerioides) or lowlands of relatively high fertility (Stipa brachychaeta) in natural grasslands of central Argentina. The two Stipa species did not differ in N and P resorption efficiency, but S. gynerioides had a higher N and P resorption proficiency than S. brachychaeta. As a consequence, leaf‐level N and P use efficiency were higher in the species adapted to low fertility conditions than in the species adapted to high fertility conditions. The higher nutrient resorption proficiency of S. gynerioides was also associated with relatively low leaf‐litter decomposition and nutrient release rates found in a previous study.     

The role of drought,disturbance, and seed dispersal in dominance shifts in a temperate grassland

Aims

More intense and frequent droughts predicted for the future will heavily affect grasslands worldwide and, along with various other disturbances, possibly trigger major vegetation shifts. Therefore, documenting these changes and understanding the mechanisms behind them is essential. Our study aimed to investigate how dominant species in a semiarid grassland respond to a combination of small-scale experimental disturbances and naturally occurring droughts and uncover the underlying mechanisms.

Location

Central Hungary.

Methods

We conducted a small-scale disturbance experiment in a semiarid temperate grassland and followed recovery for 18 years, including severe drought events. In 1999, we established 16 sites, eight dominated by Festuca vaginata and another eight by Stipa borysthenica, the two dominant species of sand grasslands in the region. At each site, three permanent plots were marked that received either a cutting or digging treatment or remained as controls. We monitored the cover and density changes of Festuca and Stipa annually.

Results

In the early years following the disturbance, Festuca recovered at a similar rate under both disturbance treatments, while Stipa recovered faster in cut than in dug plots. When natural drought events caused major diebacks of both species, Stipa recovered very quickly and regained dominance in initially Stipa-dominated plots, and it also took over in initially Festuca-dominated control and cut plots. However, digging at Festuca-dominated sites delayed drought-induced Stipa colonisation and thus favoured Festuca recovery. We found that the poor performance of Stipa in dug plots was related to sharply reduced seedling establishment, which resulted from the low number of seeds captured by the bare soil surface after digging.

Conclusions

Our results demonstrate that recurring drought events may induce dominance shifts in temperate grasslands, but small-scale disturbances can modulate vegetation responses. Our findings emphasise the importance of post-disturbance regeneration patterns in drought-induced vegetation shifts and show that seed dispersal strategy may have a major effect on vegetation dynamics.     

Edaphische Charakteristik einigerStipa-Lokalitäten in Österreich

Zusammenfassung Wir analysierten in Österreich eine Reihe von Böden verschiedener Lokalitäten vonStipa-Arten und stellten bei ihnen den Feuchtigkeitsgehalt in %, den Humus- und Stickstoffgehalt in %, den Gehalt an austauschbarem Calcium und Magnesium in mg/100 g Trockengewicht, den Gehalt an Nitraten und den Gehalt der aufnehmbaren Phosphorsäure ebenfalls in mg/100 g Trockengewicht fest.Durch Bewertung der Ergebnisse und beim Vergleich der früher erzielten Resultate charakterisierten wir die edaphischen Standortseigenschaften der Arten der GattungStipa, beschrieben die Standortseigenschaften in verschiedenen Gebieten Österreichs, versuchten schließlich auf Grund der Dispersion und des Durchschnittes der analytischen Werte einige Bodeneigenschaften der Standorte festzustellen und durch den Vergleich mit früheren Ergebnissen die edaphischen Unterschiede zwischenStipa pulcherrima, Stipa capillata undStipa joannis auszudrücken. Wir überzeugten uns, daß die edaphischen Eigenschaften, die wir auf Grund eines umfangreichen, statistisch ausgewerteten Materials von Böden derStipa-Fundorte in der Tschechoslowakei feststellten, auch für die österreichischen Verhältnisse gelten.     

Stipa adamii sp. nov. (Poaceae) from the western Tian‐Shan,and some remarks on the taxa of the section Smirnovia occurring in Kazakhstan

Stipa adamii M. Nobis sp. nov. (Poaceae), endemic to Kazakhstan, is described and illustrated. The species belongs to the section Smirnovia Tzvel. and is distributed in the central Karatau Mountains (western Tian‐Shan). It is morphologically close to Stipa karataviensis Roshev. but is easily distinguishable by longer glumes, lemmas, awns and the callus. The awn is scabrous in the lower part and the callus is bearded in S. adamii whereas both organs are glabrous in S. karataviensis. Characters distinguishing S. karataviensis and S. adamii are presented in a table, together with photographs of the callus. The analysis of morphological characters of the taxa provides new information on their variability. A key to the species of Stipa sect. Smirnovia occurring in Kazakhstan, typification of the name S. manrakica and synonymization of the name S. saurica as a syn. nov. are included.