Effects of rock climbing on plant communities on exposed limestone cliffs in the Swiss Jura mountains

Abstract. Exposed cliffs of the Swiss Jura mountains harbour a highly diverse flora with many rare and endangered plant species. Many cliffs are frequently visited by rock climbers. We examined the impact of rock climbing on vascular plants in the lower part of four cliffs of the Gerstelflue (NW Switzerland) by comparing the vegetation of climbed and unclimbed areas. In climbed areas plant cover and species density were reduced. Similarly, the density of forbs and shrubs decreased, whereas the density of ferns tended to increase. In addition, rock climbing caused a significant shift in plant species composition and altered the proportions of different plant life forms. Rock climbing can be a threat to sensitive plants of the limestone cliff community.     

Factors affecting breeding habitat selection in a cliff-nesting peregrine Falco peregrinus population

We studied factors affecting breeding habitat selection in a population of cliff-nesting peregrines Falco peregrinus across multiple spatial levels (cliff site, cliff context and land-use of the surrounding landscape), over a 2,100 km² study area in the Alps (Italy and Switzerland). We detected 30 breeding pairs (density: 1.43 territorial pairs/100 km²), whose territories were uniformly distributed over the study area. We compared 15 habitat features characterising occupied cliffs and 30 randomly selected unoccupied cliffs by means of stepwise forward logistic regression and hierarchical partitioning. The logistic regression analysis showed that occupied cliffs were longer (horizontal length), steeper, and had a greater extension of urban areas in the surrounding landscape compared to non-occupied cliffs. The model had a discrimination ability of 0.95. Hierarchical partitioning indicated that the logistic regression model was appropriate. Neither the distance of cliffs to the nearest site occupied by a potential competitor and predator, the eagle owl Bubo bubo, which occurred at a low density (0.67 pairs/100 km²), nor the distance to the nearest site occupied by peregrines had any effect on the cliff suitability model. Therefore, habitat selection in cliff-nesting peregrines was mainly influenced by cliff site features and surrounding landscape characteristics, whereas the proximity to conspecifics and to an intraguild predator had no apparent effects on peregrine settlement in our study population.     

The role of spontaneous selfing in the pioneer species Saxifraga aizoides

A large variety of reproductive patterns is present among alpine plants to ensure the persistence of populations in such harsh environments. In the present study, the role of spontaneous selfing and its contribution to the actual reproductive success of an alpine pioneer plant was investigated. The results showed that Saxifraga aizoides is clearly self-compatible. Open-pollinated flowers exhibited higher seed numbers per capsules than selfing flowers, albeit the difference was not significant. Although seed weight seemed to be independent from the kind of pollination, open-pollinated flowers had a significantly higher proportion of germinated seeds than selfed ones. Furthermore, the ability of fast germination found in S. aizoides enables the seeds to take advantage of all possible opportunities for germination. In summary, S. aizoides exhibits a successful recruitment strategy for an alpine pioneer species.     

Multi-scale comparisons of cliff vegetation in Colorado

We developed a nested vegetation sampling protocol to sample the plant diversity on south-facing cliffs and cliff bases in Jefferson County, Colorado. The multi-scale plots included three nested spatial scales, 1 m², 20 m², and 40 m². We compared plant species richness and species diversity among large cliffs, medium cliffs, small cliffs, and non-cliff sites using Hill's diversity numbers (N ₀, N ₁, and N ₂) for the 1-m² quadrats. Species richness (N ₀) was calculated for the 20-m² and 40-m² plots. Our results indicate that plant species diversity on the cliff faces did not increase with increasing cliff area. This pattern was consistent at all three sampling scales. A model selection was run to determine if plant species diversity values on the cliff faces were associated with cliff variables. None of the cliff variables measured were good predictors of diversity at the 1-m² scale. However, at the 20-m² scale, canyon differences and a positive relationship with increasing cliff surface roughness explained 70% of the variability in species richness. Although most plant species sampled on the cliff faces were also found in the base plots, 13 species were sampled only on the cliff faces. Additionally, dry south facing cliffs support a mix of xeric and mesic plants indicating that cliffs may provide unique microenvironments for plant establishment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Breeding density,habitat selection and reproductive rates of the Peregrine Falcon Falco peregrinus in Álava (northern Spain)

A population of 33–35 pairs of Peregrine Falcon in álava (north Spain) was studied with the aim of assessing the habitat attributes that influence breeding density, habitat selection and breeding success. A strong relationship was found between density of the species in each UTM (Universal Transverse Mercator) square of 10 × 10 km and cliff availability. Habitat selection was analysed by comparing 15 variables in 33 occupied and 25 unoccupied cliffs located at least 2 km from the nearest Peregrine pair. Significant differences were found in five variables: cliff dominance, distance to the nearest Golden Eagle Aquila chrysaetos nesting cliff, steepness and altitude (all showing larger values in occupied cliffs), and cliff orientation, with occupied cliffs facing preferably south and east. Orientation, dominance and distance to the nearest Golden Eagle pair, and the distance to the nearest Eagle Owl Bubo bubo nesting cliff, were included in a discriminant analysis which classified 82.76% of the cliffs correctly. The productivity of the studied population was 1.44 young/territorial pair (n = 45), and no consistent relationship was found between breeding success and habitat variables.     

Environmental factors affecting patterns of distribution and co‐occurrence of two competing raptor species

Habitat selection is a complex process, that is affected by several factors, including habitat characteristics, environmental conditions, and both intra‐ and interspecific interactions. We analysed habitat preferences of two top avian predators, Peregrine Falcon Falco peregrinus, a medium‐sized diurnal raptor, and Eagle Owl Bubo bubo, a large nocturnal raptor. These two species are known to compete for preferred nest‐sites, and proximity to cliffs with Eagle Owls may reduce Peregrine breeding output through predation of young Falcons. We investigated the environmental factors affecting occurrence and coexistence of the two species and the potential role of habitat suitability in favouring co‐occurrence in 3519 km² of the central pre‐Alps of Italy, where the two species breed on cliffs and sometimes co‐occur on the same cliff. Peregrines settled on long, steep and favourably orientated cliffs in woodland landscapes close to urban areas. Eagle Owls settled on topographically similar cliffs, but in lower rainfall areas compared with cliffs occupied by Peregrines and cliffs unoccupied by either species. Sites where the two species co‐occurred were characterized by more horizontally extended cliffs compared with sites of exclusive occurrence of each species. An analysis of relative habitat suitability revealed that sites where the two species co‐occurred had the highest predicted probability of occupancy for both species, suggesting that those sites should be regarded as high‐quality sites. Breeding productivity of Eagle Owls was negatively affected by the co‐occurrence of Peregrines, whereas the effect of Eagle Owl proximity on Peregrine productivity varied according to cliff suitability for the Peregrines. Habitat selection had fitness consequences for Eagle Owls because breeding productivity increased with cliff length. Environmental conditions, particularly climatic factors, could allow the widespread coexistence of these competing raptors at the landscape scale, whereas at the local scale co‐occurrence could take place only on larger cliffs. These were preferred sites for both species, presumably because breeding at such sites offsets the costs of settling close to the competitor species.     

Phylogenetic Relationships of the Silver Saxifrages (Saxifraga, Sect. Ligulatae Haworth): Implications for the Evolution of Substrate Specificity, Life Histories, and Biogeography

The silver saxifrages (Saxifraga sect. Ligulatae Haworth; Saxifragaceae) exhibit remarkable variation of substrate specialization, with strictly calcicole to calcifuge species, as well as life histories which range from semelparity to iteroparity. They occur almost exclusively in the European mountain ranges and display high levels of endemism. Sequences from chloroplast and nuclear ribosomal DNA were obtained to resolve phylogenetic relationships among the silver saxifrages and related taxa and to gain insight into the evolution of substrate specificity, life history, and biogeography. The resulting phylogenies suggested that (1) Saxifraga sect. Ligulatae, as traditionally defined, does not constitute a monophyletic group; (2) lime-secreting hydathodes in calcifuge species apparently represent a secondary nonaptation; (3) semelparity evolved independently two or three times in the silver saxifrages and allied sections, possibly in response to climatic changes that occured during the Pleistocene; and (4) narrow endemics, for example S. cochlearis, likely evolved from the fragmentation of the widespread S. paniculata into refugial populations that became isolated during the glacial maxima of the Pleistocene.     

Environmental relations and ecological responses of some higher plant species on rock cliffs in northern Tasmania

Abstract Gravity, moisture availability and shade exert a major control over the distributions of vascular plant species on sandstone cliffs in the Great Western Tiers, northern Tasmania. The vegetation is dominated by ferns, some of which are almost always restricted to cliffs, with invasion of shrub and tree species where soil has been able to accumulate. Although it might be expected that species commonly found over a wide range of vegetation types could displace obligate cliff dwellers, this potential is rarely fulfilled. The vertical nature of the environment precludes the loss of cliff obligates, as soil-vegetation mats peel off in part of a truncated and cyclic successional process. Experimental work and field monitoring of growth and mortality in single species and mixed stands showed that the cliff obligate fern Blechnum vulcanicum (Blume) Kuhn is slower growing, more drought resistant and less responsive to fertilizer than its non-obligate associate, B. wattsii Tind.     

Impact assessment of intense sport climbing on limestone cliffs: Response of rock-dwelling land snails

Exposed limestone cliffs in the Swiss Jura Mountains harbour a diverse gastropod community with some rare species. Sport climbing has recently increased in popularity on these cliffs. We examined the effects of sport climbing and microtopographical features of rock faces on terrestrial gastropods by assessing species diversity and abundance on climbing routes and in unclimbed areas of seven isolated cliffs in the Northern Swiss Jura Mountains. We considered exclusively living individuals resting attached to rock faces. In total, 19 gastropod species were recorded. Six of them were specialized rock-dwelling species, whose individuals spend their entire life on rock faces, feeding on algae and lichens. Plots along climbing routes harboured fewer species of rock-dwelling snails as well as other gastropod species (usually living in the leaf litter layer at the cliffs’ base) than plots in unclimbed control areas. Similarly, both the density of individuals and frequency of occurrence in plots were reduced in both groups of snails on climbing routes. The complexity of the rock surface had little influence on the species richness and abundance of gastropods. Pyramidula pusilla, the species with the smallest shell and a preference to rest underneath overhangs, was less affected by sport climbing than snail species with larger shells and a preference to rest on exposed smooth rock surface. Our findings indicate land snail diversity and abundance are suitable indicators for impact assessment in rocky habitats. Future management plans and actions should therefore not only rely on plants; they ought to consider also gastropods and other invertebrates. Any management plan should include a comprehensive information campaign to show the potential impact of intensive sport climbing on the specialized flora and fauna with the aim of educating the climbers and increasing their compliance with such measures.     

郑州郊区崖沙燕(Riparia riparia)的巢址选择

2014年3—8月,采用样方法对分布于河南省郑州市郊区的4处崖沙燕(Riparia riparia Linnaeus 1758)营巢地的13个生境因子进行了调查,测定了巢址样方和对照样方各160组数据,并利用Mann-Whitney U检验对这些数据的显著水平进行了检验;最后采用逐步判别分析(stepwise discriminate analysis)方法确定影响崖沙燕巢址选择的主要生态因子,并建立了标准化的典则判别函数。结果表明,1)崖沙燕洞巢的洞口横径(8.46±0.79)cm,纵径(8.30±0.79)cm,巢洞深(87.80±11.97)cm,巢室长径(11.25±1.33)cm,短径(10.43±1.33)cm,巢室高(9.99±1.33)cm;2)巢址样方与对照样方在植被均高、植被密度、坡位、坡度、崖高、崖长、崖龄及干扰度等8个方面差异达到显著水平,崖沙燕偏好选择的洞巢周围植被较高(97.24±8.42)cm,植被密度较大(64.34±7.15)株/m~2,坡度较大(83.78±0.35)°、上坡位(1.41±0.04)、崖较高处(7.54±0.13)m、崖壁较长(51.18±3.54)m、崖龄较短(2.41±0.05)月及人类干扰较低(1.94±0.06);3)逐步判别分析结果表明,影响崖沙燕营巢地选择的主要生境因子是坡位、崖龄和干扰度,标准化的典则判别函数为y=0.753×坡位+0.681×崖龄+0.288×干扰度。建议在城市化建设过程中针对崖沙燕的巢址选择特性建立合理的保护措施。     

Evidence for the widespread occurrence of ancient forests on cliffs

Aim The objective of this work was to determine if the existence of ancient forests on cliffs was specific to the Niagara Escarpment, Canada, or part of a globally widespread pattern. Location Sixty‐five cliff sites were visited in five countries in the temperate climatic zone, and trees were sampled for age and growth rate on forty‐six of these. Methods Two hundred and twenty‐four core samples or cross‐sections were taken from trees on cliffs that varied in height, aspect, rock‐type, and exposure. General observations were also made of regeneration of the tree species forming the mature canopy, and other habitat conditions. Results The evidence shows that ancient slow‐growing forest occurs on most cliffs. Age and growth rate distributions were similar at all treed sites. Small‐statured Thuja, Juniperus, or Taxus stems with age estimates in excess of 1000 years were found in the United States, the United Kingdom and France, and small Pinus and Quercus stems nearly 400 years in Germany. There was a high rate of recurrence of plants in the genera Polypodium, Asplenium, Cystopteris, Campanula, Rosa, Prunus, Hedera, and Sorbus. Most of the sites appear to be habitats of completely natural origin. Conclusions We conclude that ancient natural forest is a normal feature of cliffs, at least in the temperate zone.     

广西扶绥白头叶猴对栖息地的选择与利用

为了解白头叶猴（Trachypithecus leucocephalus）的栖息地利用规律及其影响因素,2016年2月至2017年1月,采用瞬时扫描取样法对广西崇左白头叶猴国家级自然保护区一群白头叶猴的栖息地利用进行了研究。结果表明,白头叶猴对山体不同部位的利用存在显著性差异（χ² =39.467,df=3,P<0.001）,其中,对崖壁（56.75±9.55）%的利用比例最大,其次是对山坡（39.42±10.93）%和山顶（2.98±2.54）%的利用,而对山脚（0.84±1.47）%的利用频率最低。白头叶猴对不同微生境类型的利用存在差异（χ²=27.709,df=3,P<0.001）,其中对乔木（49.37±12.31）%的利用比例最大,其次是裸岩（24.05±13.61）%,随后依次为藤本（15.48±8.01）%和灌木（10.87±5.45）%。白头叶猴主要在山坡上觅食,利用崖壁移动、休息,进行社会活动;主要利用裸岩进行社会活动,觅食、移动、休息主要发生在乔木上。从整体来看,白头叶猴在雨季对乔木的利用频率显著大于旱季（Z=-2.680,n=12,P=0.007）;雨季在山坡觅食频率显著大于旱季（Z=-2.517,n=12,P=0.012）,而在崖壁觅食频率刚好相反（Z=-2.842,n=12,P=0.004）;白头叶猴雨季在乔木休息的频率显著大于旱季（Z=-2.355,n=12,P=0.019）。白头叶猴对栖息地的利用受到温度的影响。白头叶猴对乔木的总体利用频率随着平均温度的升高而增加（r=0.664,n=12,P=0.018）;觅食时,对崖壁、裸岩的利用频率均与平均温度成负相关关系（崖壁：r=-0.685,n=12,P=0.014;裸岩：r=-0.600,n=12,P=0.039）;休息时,对乔木的利用频率与平均温度呈正相关关系（r=0.650,n=12,P=0.022）。不同季节,白头叶猴对栖息地的利用方式不同。白头叶猴的栖息地利用模式可能是在觅食利益和捕食风险之间作出的权衡,并受到环境温度的影响。     

Sidera, a new genus in Hymenochaetales with poroid and hydnoid species

A new genus, Sidera (Hymenochaetales), is described to accommodate the dimitic polypores Skeletocutis lenis and S. vulgaris, the monomitic polypore Ceriporiopsis lowei, and the monomitic, hydnoid Athelopsis lunata. The genus is characterised by white-rot, whitish resupinate fruiting bodies, crystal rosettes on specialised hyphae, and allantoid spores. Cinereomyces (Polyporales) is confirmed to be a good genus including C. lindbladii, separate from Diplomitoporus as defined by its type species D. flavescens and the closely related D. crustulinus. Fabisporus is a taxonomic synonym of Diplomitoporus. Poriodontia subvinosa belongs to the Hymenochaetales.     

Localized hybridization betweenSaxifraga aizoides andS. mutata (Saxifragaceae): Reproductive ecology ofS. ×hausmannii and implications for conservation

The roles of hybridization and of introgression as potential threats to small native populations are current topics of interest both for evolutionary and conservation biologists. Two factors are mainly involved in the threat to natural populations through hybridization: demographic swamping and genetic assimilation. At prealpine locations, twoSaxifraga species with very different growth habits,S. aizoides andS. mutata, form a hybrid,S. ×hausmannii. Investigation of meiosis and crossing experiments revealed regular pairing of the 26 chromosomes and development of viable seeds in the hybrid. Analysis of hybrid indices, GPI allozymes, germination experiments and seed sizes showed an intermediate position ofS. ×hausmannii between its parental species.Saxifraga aizoides andS. mutata were separated by only weak hybridization barriers. Introgression occurred mostly in direction ofS. aizoides, although to a small amount. There is no actual threat to small, prealpine populations ofS. aizoides andS. mutata through interspecific gene flow because hybridization is localized, resulting from different flowering phenologies of the parental species and from the narrow habitat requirements ofS. ×hausmannii, which are similar to those ofS. aizoides. The major threat to local, prealpine populations ofS. aizoides andS. mutata is the loss of landscape dynamics. Both species need open erosion slopes for successful recruitment.     

Across the southern Andes on fin: glacial refugia,drainage reversals and a secondary contact zone revealed by the phylogeographical signal of Galaxias platei in Patagonia

We employed DNA sequence variation at two mitochondrial (control region, COI) regions from 212 individuals of Galaxias platei (Pisces, Galaxiidae) collected throughout Patagonia (25 lakes/rivers) to examine how Andean orogeny and the climatic cycles throughout the Quaternary affected the genetic diversity and phylogeography of this species. Phylogenetic analyses revealed four deep genealogical lineages which likely represent the initial division of G. platei into eastern and western lineages by Andean uplift, followed by further subdivision of each lineage into separate glacial refugia by repeated Pleistocene glacial cycles. West of the Andes, refugia were likely restricted to the northern region of Patagonia with small relicts in the south, whereas eastern refugia appear to have been much larger and widespread, consisting of separate northern and southern regions that collectively spanned most of Argentinean Patagonia. The retreat of glacial ice following the last glacial maximum allowed re‐colonization of central Chile from nonlocal refugia from the north and east, representing a region of secondary contact between all four glacial lineages. Northwestern glacial relicts likely followed pro‐glacial lakes into central Chilean Patagonia, whereas catastrophic changes in drainage direction (Atlantic → Pacific) for several eastern palaeolakes were the likely avenues for invasions from the east. These mechanisms, combined with evidence for recent, rapid and widespread population growth could explain the extensive contemporary distribution of G. platei throughout Patagonia.     

Full light and soil drought constrain plant growth in Mediterranean cliffs: the case of Primula palinuri Petagna

In any biogeographical region, cliffs shelter a larger number of relict species than do the surrounding areas. To develop effective conservation strategies, experimental studies are needed. We focused on Mediterranean coastal cliffs and on Primula palinuri Petagna, an endangered narrow-endemic plant, included in the IUCN Red List and chosen as a key species. The hypothesis was that light intensity and soil water availability could affect survival and growth of juvenile plants. To avoid the bias of other cliff environmental factors and to overcome problems of cliff site inaccessibility, we performed an ex-situ experiment. The effect of two light regimes and two soil water availability conditions on survival and morpho-functional traits were analysed. Plants grown under reduced light intensity and wet soil showed no reduction in survival and performed very well in terms of growth and storage accumulation. By contrast, full light and dry soil proved to be detrimental conditions. Overall results suggested that cliff environmental conditions play an important role in conservation of narrow-endemic species. The decreasing trend of the already severely fragmented populations of P. palinuri and the ongoing increasing aridity is likely to contribute to a further reduction of the already narrow distribution this cliff relict species.     

Burrows of Paragnathia (Crustacea: Isopoda) and bledius (Arthropoda: Staphylinidae) Enhance cliff erosion

In coastal cliffs consisting of Pliocene fine‐grained clastic sediments at El Rompido, Huelva, Spain we discovered around the high water line an extensive zone of about 1.5 m in height burrows that were made by up to 4 mm long isopods Paragnathia formica (Hesse) and small 5 mm long staphylinid beetles (Bledius cf. corniger Rosenhauer). The burrowing zone was crowded with irregular burrows 2–5 mm in diameter and extended some 5 cm deep in the cliff. The base of the cliff showed a clear (bio)erosional notch due to this burrowing activity, comparable to those produced by marine boring and scraping organisms in, for example, calcareous rocks. This indicates that the activity of Paragnathia and Bledius helps in eroding the cliffs, a phenomenon not published earlier as far as we know. A comparable but less extensive burrowed zone was found farther upstream near Cartaya. Their presence around the high tide zone also might prove these burrows to be good markers of former high tide lines in older cliffs.     

Nurse effect of the native cushion plant Azorella monantha on the invasive non-native Taraxacum officinale in the high-Andes of central Chile

Positive interactions among native plant species are common in alpine habitats, particularly those where one species (nurse plant) generates microclimatic conditions that are more benign than the surrounding environment, facilitating the establishment of other species. Nonetheless, these microclimatic conditions could facilitate the establishment of non-native species as well. A conspicuous component of the alien alpine flora of the central Chilean Andes is the perennial herb Taraxacum officinale agg. (dandelion). In contrast to other alien species that are restricted to human-disturbed sites, T. officinale is frequently observed growing within native plant communities dominated by cushion plants. In this study we evaluated if T. officinale is positively associated with the cushion plant Azorella monantha. Via seedling survival experiments and gas-exchange measurements we also assessed the patterns of facilitation between cushions and dandelions, and explore the potential mechanisms of invasion by dandelions. T. officinale grows spatially positively associated with cushions of A. monantha. Survival of seedlings, as well as their net-photosynthetic rates and stomatal conductance, were higher within cushions than in open areas away from them, suggesting that the microclimatic modifications generated by this native cushion facilitates the establishment and performance of a non-native invasive species. Our results, as well as other recent studies, highlight the role of native communities in facilitating rather than constraining non-native plant invasions, particularly in stressful habitats such as alpine environments.     

Host relationships ofDasineura ingeris: oviposition preferences and juvenile survival on clones ofSalix viminalis, otherSalix species and hybrids

In the present study we assessed the extent to which oviposition preferences and larval survival rates of the gall midge Dasineura ingeris Sylvén & Lövgren (Diptera: Cecidomyiidae) can explain variation in its use of Salix viminalis phenotypes as hosts. Furthermore, based on oviposition preferences and juvenile survival rates we estimated the host potential of various Salix species and species hybrids. Egg distribution patterns explained 67% of the intraspecific variation in damage to S. viminalis shoots in the field. Juvenile survival, on the other hand, was not related to field damage or egg distribution patterns among the S. viminalis clones studied. Clones of complete or partial S. viminalis origin ranked highest in suitability for D. ingeris survival, and four out of these six clones also received the highest egg rankings. Species belonging to the subgenus Salix were not oviposited on at all, whereas others belonging to the same subgenus as S. viminalis, i.e. Vetrix, were intermediate in terms of oviposition rates. Although there was a significant correlation between D. ingeris egg distributions and larval survival rates on the clones in the Salix species/hybrid set, mismatches were evident.