Comparative Skin Histology of Frogs Reveals High-elevation Adaptation of the Tibetan Nanorana parkeri

Adaptations to extreme environmental conditions are intriguing. Animal skin, which directly interacts with external environment, plays diverse and important roles in adaptive evolution. The thin and bare skin of amphibians is sensitive to external environmental conditions and, thus, it facilitates investigations into adaptations for living in extreme environments. Herein, we compare the structures of skin in four anuran species living at elevations ranging from 100 m to 4500 m to assess phenotypic innovations in the skin of Nanorana parkeri, which lives at extremely high elevations. Analyses reveal similar basic skin structures, but N. parkeri differs from the other species by having more epidermal capillaries and granular glands, which correlate highly with responses to hypoxia and/or ultraviolet(UV) radiation. Further intraspecific comparisons from frogs taken at ~4500 m and ~2900 m reveal that all of the changes are fixed. Changes occurring only in the higher elevation population, such as possessing more skin pigments, may represent local adaptations to coldness and/or UV radiation. These results provide a morphological basis for understanding further the molecular adaptations of these frogs.

High-elevation Adaptation of Motion Visual Display Modifications in the Toad-Headed Agamid Lizards(Phrynocephalus)

Understanding the process of adaptation is a key mission in modern evolutionary biology.Animals living at high elevations face challenges in energy meta bolism due to several environmental constraints(e.g., oxygen supply, food availa bility,and movement time). Animal behavioral processes are intimately related to energy meta bolism, and therefore, behavioral modifica tions are expected to be an important mechanism for high-elevation adaptation. We tested this behavioral adaptation hypothesis using va ria tions of motion visual displays in toad-headed agamid lizards of the genus Phr ynocephalus. We predicted tha t complexity of visual motion displays would decrease with the increase of elevation, because motion visual displays are energetically costly. Displays of 12 Phr ynocephalus species were collected with elevations ranging from sea level to 4600 m. We quantified display complexity using the number of display components, display duration, pathways of display components, as well as display speed for each species. Association between display complexity and elevation was analyzed using the phylogenetic generalized least squares(PGLS)model. We found that both the number of display components and the average value of tail coil speed were negatively correlated with elevation, suggesting that toad-headed lizards living at high-elevation areas reduced their display complexity to cope with the environmental constraints. Our research provides direct evidence for high-elevation adaptation from a behavioral aspect and illustrates the potential impacts of environment heterogeneity on motion visual display diversification.

Massive Molecular Parallel Evolution of the HSP90AA1 Gene between High-elevation Anurans

HSP90 AA1 is part of the heat shock protein 90 gene family and has important functions against heat stress. We report a case of molecular level parallel evolution of the HSP90 AA1 gene in high elevation amphibians. HSP90 AA1 gene sequences of four high-elevation anurans, Bufo gargarizans, Nanorana parkeri, Rana kukunoris, and Scutiger boulengeri, were compared along with five of their low-elevation relatives. A total of 16 amino-acid sites were identified as parallel evolution between N. parkeri and R. kukunoris. We generated both model based（Zhang and Kumar＇s test） and empirical data based（parallel/divergence plotting） null distributions for non-parallel evolution, and both methods clearly determined that the observed number of parallel substitutions were significantly more than the null expectation. Furthermore, on the HSP90 AA1 gene tree, N. parkeri and R. kukunoris formed a strongly supported clade that was away from their respective relatives. This study provides a clear case of molecular parallel evolution, which may have significant implications in understanding the genetic mechanisms of high-elevation adaptation.

High-elevation hypoxia impacts perinatal physiology and performance in a potential montane colonizer

Climate change is generating range shifts in many organisms,notably along the elevational gradient in mountainous environments.However,moving up in elevation exposes organisms to lower oxygen availability,which may reduce the successful reproduction and development of oviparous organisms.To test this possibility in an upwardcolonizing species,we artificially incubated developing embryos of the viperine snake(Natrix maura)using a split-clutch design,in conditions of extreme high elevation(hypoxia at 2877 m above sea level;72%sea-level equivalent O2 availability)or low elevation(control group;i.e.normoxia at 436 m above sea level).Hatching success did not differ between the two treatments.Embryos developing at extreme high elevation had higher heart rates and hatched earlier,resulting in hatchlings that were smaller in body size and slower swimmers compared to their siblings incubated at lower elevation.Furthermore,post-hatching reciprocal transplant of juveniles showed that snakes which developed at extreme high elevation,when transferred back to low elevation,did not recover full performance compared to their siblings from the low elevation incubation treatment.These results suggest that incubation at extreme high elevation,including the effects of hypoxia,will not prevent oviparous ectotherms from producing viable young,but may pose significant physiological challenges on developing offspring in ovo.These early-life performance limitations imposed by extreme high elevation could have negative consequences on adult phenotypes,including on fitness-related traits.

Distribution assessment of soil erosion with revised RUSLE model in Tianshan Mountains

aSoil degradation caused by soil erosion is one of the world's most critical environmental issues.Soil erosion in the Tianshan Mountains has caused various environmental problems in the surrounding areas.This study used remote sensing data to analyze the distribution of the factors influencing soil erosion,and the revised universal soil loss equation(RUSLE)to calculate the total amount and distribution characteristics of soil erosion in the Tianshan Mountains in 2019.Due to the large error of RUSLE in soil erosion estimation in mountainous areas,this study modified RUSLE equation based on the characteristics of snow cover in the Tianshan Mountains.The results show that the average soil erosion was 1690.3 t/(km^(2)·year),of which insignificant erosion,slight erosion and moderate erosion accounted for 42,8%,22.4%and 9.9%,respectively.Severe erosion and above accounted for 13.3%.The accuracy of the soil erosion modulus calculated by the RUSLE was only 61.9%,with an average error of 1631.9 t/(km^(2)·year).The average error of the double-coefficient correction method was 1259.1 t/(km^(2)·year),and the average error of the modified formula method was reduced by 40.3%compared with the RUSLE,reaching 973.7 t/(km^(2)·year),and its accuracy reached 76.2%.Very severe erosion and catastrophic erosion are distributed on mountain ridges with higher elevation and on the northern area with higher precipitation.Snow cover has a certain inhibitory effect on soil erosion,and snow cover in alpine mountains is a factor that cannot be ignored in soil erosion research.