Ecology of the pteridophytes on the southern slopes of Mt. Kilimanjaro – I. Altitudinal distribution

140 taxa of 61 genera in 24 families of pteridophytes were recorded on the southern slopes of Mt. Kilimanjaro. These represent about one third of the entire pteridophyte flora of Tanzania. The families richest in species are the Aspleniaceae, the Adiantaceae, the Dryopteridaceae, the Thelypteridaceae and the Hymenophyllaceae. Due to its luxuriant montane rain forest, which receives a precipitation of up to over 3000 mm, Mt. Kilimanjaro is distinctly richer in pteridophyte species than other volcanoes in East Africa. However, compared with the mountains of the Eastern Arc, the number of pteridophytes on Mt. Kilimanjaro is smaller. This can be explained by the widely destroyed submontane (intermediate) forest rather than by the higher age of the Eastern Arc Mts.The altitudinal distribution of all ferns was investigated in 24 transects. On the southern slopes of Mt. Kilimanjaro they were found in an altitudinal range of 3640 m. Cyclosorus quadrangularis, Azolla nilotica, Azolla africana andMarsilea minuta are restricted to the foothills, while Polystichum wilsonii, Cystopteris nivalis and Asplenium adiantum-nigrum are species found in the highest altitudes.Based on unidimensionally constrained clustering and on the analysis of the lowermost and uppermost occurrence of species, floristic discontinuities within the transects were determined. From these data and from an evaluation of the distribution of ecological groups and life forms, 11 altitudinal zones could be distinguished: a colline zone (–900 m asl), a submontane zone (900–1600 m asl) with lower and upper subzones, a montane zone (1600-2800 m asl) divided into 4 subzones, a subalpine zone (2800–3900 m asl) with lower, middle and upper subzones, and finally a (lower) alpine zone above 3900 m. The highest species numbers were observed in the lower montane forest belt between 1600 and 2000 m altitude. The zonation of ferns found at Mt. Kilimanjaro corresponds well with the vegetational zonation described by other authors using bryophytes as indicators in different parts of the humid tropics.     

柠檬酸和EDTA对铜污染土壤环境中吊兰生长的影响

通过盆栽试验研究了在铜污染条件下,柠檬酸和EDTA作为活化剂对铜污染土壤中吊兰生长状况的影响.结果表明,柠檬酸和EDTA对吊兰富集量的影响与其对土壤中铜的活化能力呈显著性正相关.柠檬酸对土壤铜有较强的活化作用,能够有效提高吊兰对铜的吸收,且在浓度为5mmol/L时效果最为明显,而较高的铜富集量又抑制了吊兰的生长;EDTA对吊兰富集能力的影响相对较弱,对吊兰的生长也无显著影响.相比而言,柠檬酸对铜污染土壤中吊兰生长状况的影响比EDTA大.     

Community structure of ground beetles (Coleoptera: Carabidae) along an altitudinal gradient on Mt. Sobaeksan,Korea

We conducted this study to determine the diversity patterns, community structures, and seasonality of ground beetle assemblages along an altitudinal gradient (437–1420 m) on Mt. Sobaeksan, Korea. Ground beetles were collected by pitfall traps installed along an altitudinal gradient (437, 757, 1100, and 1420 m). A total of 32 species belonging to seven subfamilies were identified from 3259 collected ground beetles. The diversity pattern of the ground beetle assemblage according to altitude was neither monotonic decreased nor hump-shaped. However, subfamily assemblages and wing form diversity patterns differed according to altitude and may be correlated with altitude or some other environmental variables. The dominant species were Synuchus cycloderus (29.4%) and Eucarabus sternbergi sobaeksanensis (15.4%) and their seasonal activities according to altitude were similar. According to non-metric multidimensional scaling, ground beetles and altitudes could be divided into two distinct groups: a low altitude group (437–757 m) and a high altitude group (1100–1420 m). Some species were particularly abundant at high altitudes, such as Aulonocarabus koreanus kwonileeique and Poecilus nitidicollis, while others were highly abundant at low altitudes, such as Pterostichus audax, Pterostichus ishikawai, and Synuchus species.     

Comparative phenological and water potential patterns of three Calligonum species in the eastern great Erg of Tunisia

Calligonum azel, C. comosum and C. arich (Polygonaceae) are three dominant and economically important species widely distributed in active sand dunes in the southern desert of Tunisia, which differ in growth form and may co-occur under the same climatic constraints. The aims of this work were to compare their phenological and water potential patterns, as well as branch, flower and fruit production during three years at monthly intervals. Predawn (Ψ_pd) and midday (Ψ_md) water potentials followed similar trends with a few exceptions. For all species, Ψ_md reached values more negative than −2.5 MPa during the hot and dry summer, with extremes in July. Ψ_pd differed between the species, and C. comosum attained the most negative values (−1.71 MPa). Seasonal differences in Ψ_pd were less pronounced for Calligonum species established in dunes as compared to C. comosum from interdunes. Spatial and temporal variations in soil water content provoked changes in water potentials of these species. Diurnal amplitudes resulting from the difference between Ψ_md and Ψ_pd were more pronounced during the dry season compared to the wet months. The phenological pattern of the three species showed a similar sequence of phenophases with growth activity occurring mostly between spring and summer. We found a species-dependent response to summer drought, as C. comosum ended all phenophases in June, while C. azel and C. arich extended their vegetative growth into the summer period of low precipitation. Water availability for the plants was better on the dune slopes. Differences were statistically significant among species, concerning green branch production and numbers of initially formed flowers and mature fruits. Our data suggest that changes in the studied parameters may depend on the rooting depth of the species.     

A new species of Muscari subgenus Leopoldia from the Iberian Peninsula

The biological analysis of several populations confirms the existence in the Iberian Peninsula of two species of Muscari subgenus Leopoldia: M. comosum and M. matritensis (the latter described as new). Karyologically, they differ in at least a translocation rearrangement, in the presence of heterochromatin, which is lacking in M. matritensis, and in some chromosomal characters. At the genic level, the distance between M. comosum and M. matritensis is high (0.378). Morphologically, they show some differences in floral structure which are correlated with different reproductive systems, allogamy in M. comosum and autogamy in M. matritensis. The possible evolutionary relationships between these taxa is discussed.     

云南花椒园中昆虫群落特征的海拔间差异分析

分析了金沙江畔7个不同海拔区域花椒园中昆虫群落特征的动态变化及其与温、湿度变化的关系。结果表明:花椒园中昆虫群落的多样性、均匀度、丰富度、Ss/Si、Sn/Sp指标随海拔升高先增加后减少,而个体数、优势集中性指数则随海拔升高先减少后增加;低海拔(450—750 m)区域花椒园中由于总体温度高、湿度低,较高海拔(1300—1550 m)区域总体温度低、湿度高,均不利昆虫群落多样性、均匀度、丰富度、Ss/Si、Sn/Sp指标的提高和群落的稳定,而中等海拔(750—1300 m)区域的温度、湿度较为适中,昆虫群落的物种数及个体数量均相对较高,昆虫群落多样性、均匀度、丰富度、Ss/Si、Sn/Sp指标也较高,群落较稳定;相反,低海拔、较高海拔区域花椒园中昆虫群落优势集中性指数较高,而中等海拔区域却相对较低。综合分析表明,海拔高度及温湿度对花椒园昆虫群落特征影响较大,而海拔高度及温湿度的测量和分析较为方便。     

Facilitative interactions do not wane with warming at high elevations in the Andes

Positive interactions between species are known to play an important role in the structure and dynamics of alpine plant communities. The balance between negative and positive interactions is known to shift along spatial and temporal gradients, with positive effects prevailing over negative ones as the environmental stress increases. Thus, this balance is likely to be affected by climate change. We hypothesized that increases in temperature (a global warming scenario) should decrease the importance of positive interactions for the survival and growth of alpine plant species. To test this hypothesis, we selected individuals of the native grass species Hordeum comosum growing within the nurse cushion species Azorella madreporica at 3,600 m.a.s.l. in Los Andes (Chile), and performed nurse removal and seedling survival experiments under natural and warmer conditions. For warmer conditions, we used open-top chambers, which increased the temperature by 4 °C. After two growing seasons, we compared the effect of nurse removal on the survival, biomass, and photochemical efficiency of H. comosum individuals under warmer and natural conditions. Nurse removal significantly decreased the survival, biomass, and photochemical efficiency of H. comosum, demonstrating the facilitative effects of nurse cushions. Seedling survival was also enhanced by cushions, even under warmer conditions. However, warmer conditions only partially mitigated the negative effects of nurse removal, suggesting that facilitative effects of cushions do not wane under warmer conditions. Thus, facilitative interactions are vital to the performance and survival of alpine species, and these positive interactions will continue to be important in the warmer conditions of the future in high-alpine habitats.     

Altitudinal gradients,plant hybrid zones and evolutionary novelty

Altitudinal gradients are characterized by steep changes of the physical and biotic environment that present challenges to plant adaptation throughout large parts of the world. Hybrid zones may form where related species inhabit different neighbouring altitudes and can facilitate interspecific gene flow and potentially the breakdown of species barriers. Studies of such hybrid zones can reveal much about the genetic basis of adaptation to environmental differences stemming from changes in altitude and the maintenance of species divergence in the face of gene flow. Furthermore, owing to recombination and transgressive effects, such hybrid zones can be sources of evolutionary novelty. We document plant hybrid zones associated with altitudinal gradients and emphasize similarities and differences in their structure. We then focus on recent studies of a hybrid zone between two Senecio species that occur at high and low altitude on Mount Etna, Sicily, showing how adaptation to local environments and intrinsic selection against hybrids act to maintain it. Finally, we consider the potential of altitudinal hybrid zones for generating evolutionary novelty through adaptive introgression and hybrid speciation. Examples of homoploid hybrid species of Senecio and Pinus that originated from altitudinal hybrid zones are discussed.     

Stable isotope analysis reveals variation in trophic niche depending on altitude in an endemic alpine gecko

Interspecific competition is considered a major determinant of ecological niche. It is hypothesized that increased competition should reduce niche breadth. However, there are scarce field tests on this hypothesis. Here, we test this central hypothesis in ecology by using the Atlas day gecko Quedenfeldtia trachyblepharus. This alpine gecko faces fewer competitors as altitude increases, and thereby, we predict that this species should increase niche breadth and relevant fitness parameters with altitude. We tested this prediction by analysing the isotopic signature of carbon (δ¹³C) and nitrogen (δ¹⁵N). Our results reveal that specimens from higher altitudes showed higher values for both carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes, had better body condition and a greater isotopic breadth when compared to specimens from lower altitudes. Altitudinal variation in carbon values was not explained by variation in isotopic concentration in the baseline of the trophic chain. Therefore, our findings support the prediction that relaxed interspecific competition favours increased trophic niche breadth. These results also suggest that global warming may represent an important threat for this species, as it may provoke the ascent in altitude of competitors, with negative consequences for the conservation of this endemism.     

Genetic Pool Information Reflects Highly Suitable Areas: The Case of Two Parapatric Endangered Species of Tuco-tucos (Rodentia: Ctenomiydae)

Conservation of small mammals requires knowledge of the genetically and ecologically meaningful spatial scales at which species respond to habitat modifications. Conservation strategies can be improved through the use of ecological niche models and genetic data to classify areas of high environmental suitability. In this study, we applied a Maxent model integrated with genetic information (nucleotide diversity, haplotype diversity and Fu''s Fs neutrality tests) to evaluate potential genetic pool populations with highly suitable areas for two parapatric endangered species of tuco-tucos (Ctenomys minutus and C. lami). Our results demonstrated that both species were largely influenced by vegetation and soil variables at a landscape scale and inhabit a highly specific niche. Ctenomys minutus was also influenced by the variable altitude; the species was associated with low altitudes (sea level). Our model of genetic data associated with environmental suitability indicate that the genetic pool data were associated with highly suitable areas for C. minutus. This pattern was not evident for C. lami, but this outcome could be a consequence of the restricted range of the species. The preservation of species requires not only detailed knowledge of their natural history and genetic structure but also information on the availability of suitable areas where species can survive, and such knowledge can aid significantly in conservation planning. This finding reinforces the use of these two techniques for planning conservation actions.     

Effects of wind and thermal conditions on timberline formation in central Japan: a lattice model

The upper distribution limit of tall tree species Abies mariesii is the timberline in central Japan, and dwarf pine Pinus pumila dominates above the timberline to near the summit. My previous studies suggested that the main cause of the timberline formation is the increase in mortality due to strong wind in winter rather than low growth due to low summer temperature. This study evaluated how wind velocity affects timberline formation and if the altitude of timberline moves upward due to high thermal conditions, by using a lattice model. Increase in wind velocity throughout the altitude lowered the altitudes of upper distribution limits of the two species. On the contrary, prolonged growth period due to high thermal conditions increased the upper distribution limit of P. pumila, and the upper distribution limit of A. mariesii was hardly affected by the change of growth period. However, the upward shift of the upper distribution limit of P. pumila due to the prolonged growth period in the model would not be realistic because P. pumila had already distributed up to near the summit. This study concludes that A. mariesii is a superior competitor to P. pumila at low altitudes with low wind velocity, but dwarf pine P. pumila can dominate at higher altitudes because A. mariesii suffers severe mechanical damage due to strong wind in winter, and that the altitude of the timberline does not move upward even under high thermal conditions due to global warming.     

Identifying factors determining the altitudinal distribution of the invasive pest leafminers Liriomyza huidobrensis and Liriomyza sativae

Abiotic and biotic factors that change with altitude can influence the distribution of herbivorous insects. We examined factors influencing the distribution of the generalist leafminers Liriomyza sativae Blanchard and Liriomyza huidobrensis Blanchard (Diptera: Agromyzidae), two pests of agricultural crops, in the tropical Dieng mountainous area of Central Java, Indonesia. Liriomyza huidobrensis predominated at altitudes above 700 m a.s.l. and was the only species collected above 1 400 m a.s.l. In contrast, L. sativae predominated below 600 m a.s.l. and was not found above 1 200 m where the average temperature was 20.7 °C. Parasitoid diversity decreased with altitude, but parasitism did not change. The distribution of neither species was affected by parasitoids; L. huidobrensis predominant at high altitudes was preferred by Opius parasitoids common at these altitudes. Intra‐ and interspecific competition was detected in laboratory experiments where larval density was high, but led to coexistence rather than species displacement. No competition was detected in a field experiment when larval density was low. However, L. sativae failed to reproduce at the highest altitude, whereas L. huidobrensis established at all altitudes. Host composition varied with altitude and one host (faba beans) preferred by L. huidobrensis was common at high altitudes. By relating published data on the performance of the leafminer species to altitudinal temperature changes, we were partly successful in predicting the altitude at which the dominant species switched. Temperature plays an overriding influence on the altitudinal distribution of leafminers.     

Germination response to various temperature regimes of four Mediterranean seeder shrubs across a range of altitudes

In dry-summer seasonal climates, the beginning of the rainy season can prompt germination under different temperatures, depending on altitude. Understanding germination responses to temperature with altitude is important in fire-prone environments for species regenerating after fire from seeds (seeders), particularly under changing climate. Here we investigated the role of temperature in four Mediterranean seeder shrubs from Central Spain. Seeds from 17 sites (285–1,253 m altitude), of two hard-seeded nanophanerophytes (Cistus ladanifer and C. salviifolius) and two soft-seeded chamaephytes (Lavandula pedunculata and Thymus mastichina) were investigated. Intact and heat shock treated seeds were set to germinate under four temperature regimes, including a treatment simulating future warming. GLM with binomial or gamma functions were used to test treatment effects using altitude as a covariate. Altitude was a significant covariate only in L. pedunculata. Temperature did not affect final germination in either Cistus, but it significantly affected T. mastichina, and interacted with altitude in L. pedunculata, whereby the higher the altitude the less it germinated with decreasing temperature. Germination speed (T₅₀) was lower at colder temperatures in all but C. salviifolius that was insensitive to our treatments. Heat shock significantly increased final germination in both Cistus and T. mastichina, but did not interact with temperature or altitude. We conclude that germination response to temperature, including varying sensitivity with altitude, differed among these species; thus, changes in the timing of the onset of the rainy season will diversely affect populations at various altitudes. We discuss our results in a context of changing climate and fire.     

Effects of altitude of origin on trichome-mediated anti-herbivore resistance in wild Andean potatoes

The diversity and abundance of herbivores often decreases with increasing altitude. Plants are expected to respond to reduced herbivore pressure by allocating fewer resources to anti-herbivore resistance at higher altitudes. This study uses a greenhouse experiment and parallel bioassay to examine population variability in trichome-mediated defenses across a range of accessions of Solanum berthaultii Hawkes and S. tarijense Hawkes that originated from different altitudes. S. berthaultii bears two types of glandular trichomes, type A and type B, whereas S. tarijense has type A trichomes only, and hairs resembling type B trichomes that are eglandular. Both type A and type B glandular trichomes on S. berthaultii deterred ovipositing female Phthorimaea operculella (Zell.) (Lepidoptera: Gelechiidae). The density of type A, but not type B trichomes decreased with increasing altitude of origin in S. berthaultii populations. The ratio of type A to type B trichomes on the abaxial surface of S. berthaultii negatively affected oviposition and was inversely related to altitude of origin. In S. tarijense, type A trichomes deterred, but eglandular leaf hairs stimulated oviposition. Consequently, the ratio of type A trichomes to eglandular leaf hairs was negatively related to the number of eggs laid. The total numbers of trichomes per leaf generally increased with increasing altitude of origin in S. tarijense whereas the ratio of type A trichomes to eglandular hairs declined. In the oviposition bioassay, we found no direct relationship between resistance and altitude of origin, which may have been due to differences in leaf area at the time of the bioassay. Nevertheless, the results suggest that populations of both plant species that originated from higher altitudes were generally more susceptible to ovipositing P. operculella.     

Continuum or zonation? Altitudinal gradients in the forest vegetation of Mt. Kilimanjaro

Based on the analysis of 600 vegetation plots using the method of Braun-Blanquet (1964) the composition of the whole vascular forest plant flora with about 1220 species was studied in the forests of Mt. Kilimanjaro. The altitudinal distribution of all strata (trees, shrubs, epiphytes, lianas and herbs) along a transect of 2400 m is discussed with respect to altitudinal zonation and ecological factors. With uni-dimensionally constraint clustering significant discontinuities were revealed that occurred simultaneously in the different strata. Thus even in structurally highly complex, multilayered tropical montane forests distinct community units exist that can be surveyed and classified by the Braun-Blanquet approach. This observed zonation was significantly correlated with altitude, temperature and soil acidity (pH); rainfall was of importance in particular for the zonation of epiphytes. Other key factors were humidity (influenced by stable cloud condensation belts) and minimum temperature (in particular the occurrence of frost at 2700 m altitude upslope). The contrary results of other transect studies in East Africa in respect to continuity of change in floristic composition appear to be caused by different sampling methods and intensities or mixing of data from areas with different climate conditions, whereas species richness did not influence the clarity of floristic discontinuities on Kilimanjaro and other parts of East Africa.     

长白山北坡土壤动物群落物种共有度的海拔梯度变化

群落间各生活型土壤动物科及所有土壤动物科的共有度,一定程度上反映了群落间的相关性及沿环境梯度的物种替代关系。应用梯度格局法在长白山北坡海拔550～2650m地段,每隔100m海拔高度设置一块样地,共计22块样地,并应用Jaccard指数,对各海拔不同土壤动物群落间物种共有度的海拔梯度变化进行了分析。结果表明,不同海拔土壤动物群落间物种的共有度,无论以哪一生活层的土壤动物科来表达,基本都以与其最相邻的群落间为最高,群落间物种共有度峰值与谷点反映了群落类型随海拔具有间断性变化。线虫、甲螨、弹尾等各生活型土壤动物的共有度以及所有土壤动物科的共有度,均随着海拔差的增加呈下降趋势,但并非与植被情况完全吻合。     

广西猫儿山不同海拔常绿树种和落叶树种光合速率与氮的关系

分析广西猫儿山不同海拔常绿和落叶树种的光合作用-氮关系,探讨光合氮利用效率(PNUE)是否受到叶片习性和海拔的影响。落叶树种的PNUE都显著高于常绿树种,这与前者有较低的比叶重(LMA)和较高的单位叶重光合速率(Amass)、氮含量和气孔导度(gs)有密切关系。高海拔树种的PNUE显著低于中低海拔树种的PNUE,这与前者较高的LMA和较低的Amass和gs相关。PNUE和相关的叶片特征的主成分分析表明常绿-落叶树种和低海拔-中海拔-高海拔树种的分布是一个自然过渡的过程。此外,PNUE与土壤碳:氮比没有显著相关性,但与年均温正相关,这表明温度气候是调节PNUE沿海拔变化的主要环境因素。因此,这种叶片习性和温度气候调节的PNUE变化可能是调节猫儿山常绿树种沿海拔形成双峰分布的一种机制。     

Alien seedling recruitment as a response to altitude and soil disturbance in the mountain grasslands of central Argentina

Climate and disturbance are considered key factors in explaining plant invasion, mainly by their effect on alien seedling recruitment. We tested whether soil disturbance by digging and altitude, as a sub-set of climate, affected the seedling recruitment of two alien species (Cirsium vulgare and Melilotus alba) in the mountain grasslands of central Argentina. We performed a seed-addition factorial experiment with altitude as a fixed factor co-variable (6 levels), time since sowing (6 levels) as an autoregressive co-variable, and soil disturbance by digging (two levels) and seed addition (two levels) as fixed factors. Seeds of the studied species were sown in replicated undisturbed and disturbed soil plots, in grassland stands located every 200 m from 1200 to 2200 m a.s.l. The number of emerging seedlings at each plot was recorded during a period of 8 months, at intervals of 30–60 days. Seedlings of both alien species emerged at all the studied altitudes, but only the recruitment of C. vulgare increased linearly with altitude. Moreover, the time window of seedling recruitment for this species increased with altitude. Soil disturbance produced a significant decrease in overall seedling emergence of both species; however, an increase in emergence was observed for C. vulgare at an altitude over 2000 m a.s.l. Natural emergence inside control plots without seed addition was only observed for Cirsium vulgare, which indicates that this species was not seed limited. Our results show that altitude and the absence of disturbance do not restrict the recruitment of C. vulgare and M. alba in these natural mountain grasslands. Moreover, these alien species seem to be tolerant to and even favoured by conditions occurring at higher altitudes and in absence of micro-site soil disturbances in the Córdoba mountain grasslands.     

A study of the mosquito Culex pipiens (Diptera, Culicidae) population structure in the transcaucasia using molecular identification methods

The mosquito species Culex pipiens consists of two forms, or ecotypes: the typical pipiens form and the molestus form. These forms are similar morphologically but have significant ecophysiological differences. The Cx. pipiens population structure was studied in open and underground habitats (house basements) in several localities in Abkhazia, Georgia, and Azerbaijan. The pipiens and molestus forms were identified by a molecular method, namely by restriction analysis of the amplification products with HaeIII endonuclease. Altogether, 90 individuals from 10 local populations were studied. The pipiens form was found only in surface water bodies (8 cases); in two cases both forms were found in one habitat. Homogeneous molestus populations were recorded in two underground habitats. Analysis of the Cx. pipiens population structure by the molecular method agrees with the results of biological and ecological studies of this mosquito in Georgia by Sh. Sichinava (1974, 1978, 1989). Thus, molecular diagnostics of intraspecific forms in Cx. pipiens populations is doubtlessly reliable.     

Ecology of Hippophae salicifolia D. Don of temperate and sub-alpine forests of North Sikkim Himalayas—a case study

Hippophae salicifolia D. Don or Sea buckthorn is one species that has great potential for improving ecological and economic developments in the mountain areas of North Sikkim. The plant is found naturally at altitudes ranging from 2,377 to 3,093 m. in riverine (river side or torrential side) and non-riverine (non river or non torrential side) areas where geo-physico-chemical parameters like aerial temperature, aerial moisture, soil pH, soil temperature, and soil moisture varied significantly. We analyzed ecological parameters and species association using standard quadrates. Actinorhizal plants invade nitrogen-poor soils because of their ability to form root nodule symbioses with N₂-fixing actinomycetes known as Frankia. Variation in nodulation with respect to plant height was also observed. This was higher in riverine areas compared to non riverine sites at higher altitudes.