Abstract
Fan life forms are bryophytes with shoots rising from vertical substratum that branch repeatedly in the horizontal plane to form flattened photosynthetic surfaces, which are well suited for intercepting water from moving air. However, detailed water relations, gas exchange characteristics of fan bryophytes and their adaptations to particular microhabitats remain poorly understood. In this study, we measured and analyzed microclimatic data, as well as water release curves, pressure–volume relationships and photosynthetic water and light response curves for three common fan bryophytes in an Asian subtropical montane cloud forest (SMCF). Results demonstrate high relative humidity but low light levels and temperatures in the understory, and a strong effect of fog on water availability for bryophytes in the SMCF. The facts that fan bryophytes in dry air lose most of their free water within 1 h, and a strong dependence of net photosynthesis rates on water content, imply that the transition from a hydrated, photosynthetically active state to a dry, inactive state is rapid. In addition, fan bryophytes developed relatively high cell wall elasticity and the osmoregulatory capacity to tolerate desiccation. These fan bryophytes had low light saturation and compensation point of photosynthesis, indicating shade tolerance. It is likely that fan bryophytes can flourish on tree trunks in the SMCF because of substantial annual precipitation, average relative humidity, and frequent and persistent fog, which can provide continual water sources for them to intercept. Nevertheless, the low water retention capacity and strong dependence of net photosynthesis on water content of fan bryophytes indicate a high risk of unbalanced carbon budget if the frequency and severity of drought increase in the future as predicted.
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References
Bates JW (1998) Is’ life-form’a useful concept in bryophyte ecology? Oikos 82:223–237
Beckett RP (1997) Pressure–volume analysis of a range of poikilohydric plants implies the existence of negative turgor in vegetative cells. Ann Bot 79:145–152
Chen L, Liu W-Y, Wang G-S (2010) Estimation of epiphytic biomass and nutrient pools in the subtropical montane cloud forest in the Ailao Mountains, south-western China. Ecol Res 25:315–325
Coxson DS, McIntyre DD, Vogel HJ (1992) Pulse release of sugars and polyols from canopy bryophytes in tropical montane rain forest (Guadeloupe, French West Indies). Biotropica 24:121–133
Dilks TJK, Proctor MCF (1979) Photosynthesis, respiration and water content in bryophytes. New Phytol 82:97–114
Duell R (1984) Computerized evaluation of the distribution of European liverworts. J Hattori Bot Lab 56:1–5
Fan Z-X, Thomas A (2013) Spatiotemporal variability of reference evapotranspiration and its contributing climatic factors in Yunnan Province, SW China, 1961–2004. Clim Change 116:309–325
Frahm J-P (1994) A contribution to the bryoflora of the Chocó region, Colombia. I. Mosses. Trop Bryol 9:89–110
Frahm J-P, Ohlemüller R (2001) Ecology of bryophytes along altitudinal and latitudinal gradients in New Zealand. Studies in austral temperate rain forest bryophytes 15. Trop Bryol 20:117–137
Green TGA, Sancho LG, Pintado A (2011) Ecophysiology of desiccation/rehydration cycles in mosses and lichens. In: Lüttge U, Beck E, Bartels D (eds) Plant desiccation tolerance, vol 215. Springer, Heidelberg, pp 89–120
Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A (2005) Very high resolution interpolated climate surfaces for global land areas. Int J Climatol 25:1965–1978
Kürschner H, Frey W, Parolly G (1999) Patterns and adaptive trends of life forms, life strategies and ecomorphological structures in tropical epiphytic bryophytes––a pantropical synopsis. Nova Hedwig 69:73–100
León-Vargas Y, Engwald S, Proctor MC (2006) Microclimate, light adaptation and desiccation tolerance of epiphytic bryophytes in two Venezuelan cloud forests. J Biogeogr 33:901–913
Li S, Liu W-Y, Li D-W (2013) Epiphytic lichens in subtropical forest ecosystems in southwest China: species diversity and implications for conservation. Biol Conserv 159:88–95
Ma W (2009) The composition and biomass of epiphytic materials and their relationships with ecological factors in Xujiaba region from Ailao Mountain, Yunnan. Unpublished Ph.D. Thesis, Graduate School of the Chinese Academy of Sciences, Beijing
Ma W-Z, Liu W-Y, Li X-J (2009) Species composition and life forms of epiphytic bryophytes in old-growth and secondary forests in Mt. Ailao, SW China. Cryptogam Bryol 30:477–500
Marschall M, Proctor MCF (2004) Are bryophytes shade plants? Photosynthetic light responses and proportions of chlorophyll a, chlorophyll b and total carotenoids. Ann Bot 94:593–603
Miller HA (1982) Bryophyte evolution and geography. Biol J Linn Soc 18:145–196
Norris D (1990) Bryophytes in perennially moist forests of Papua New Guinea: ecological orientation and predictions of disturbance effects. Bot J Linn Soc 104:281–291
Proctor MCF (1990) The physiological basis of bryophyte production. Bot J Linn Soc 104:61–77
Proctor MCF (1999) Water-relations parameters of some bryophytes evaluated by thermocouple psychrometry. J Bryol 21:263–270
Proctor MCF (2000) The bryophyte paradox: tolerance of desiccation, evasion of drought. Plant Ecol 151:41–49
Proctor MCF (2001) Patterns of desiccation tolerance and recovery in bryophytes. Plant Growth Regul 35:147–156
Proctor MCF (2002) Ecophysiological measurements on two pendulous forest mosses from Uganda, Pilotrichella ampullacea and Floribundaria floribunda. J Bryol 24:223–232
Proctor MCF (2004a) How long must a desiccation-tolerant moss tolerate desiccation? Some results of 2 years’ data logging on Grimmia pulvinata. Physiol Plant 122:21–27
Proctor MCF (2004b) Light and desiccation responses of Weymouthia mollis and W. cochlearifolia, two pendulous rainforest epiphytes from Australia and New Zealand. J Bryol 26:167–173
Proctor MCF (2012) Light and desiccation responses of some Hymenophyllaceae (filmy ferns) from Trinidad, Venezuela and New Zealand: poikilohydry in a light-limited but low evaporation ecological niche. Ann Bot 109:1019–1026
Proctor MCF, Tuba Z (2002) Poikilohydry and homoihydry: antithesis or spectrum of possibilities? New Phytol 156:327–349
Proctor MCF, Nagy Z, Csintalan Z, Takács Z (1998) Water-content components in bryophytes: analysis of pressure-volume relationships. J Exp Bot 49:1845–1854
Proctor MCF, Oliver MJ, Wood AJ, Alpert P, Stark LR, Cleavitt NL, Mishler BD (2007) Desiccation-tolerance in bryophytes: a review. Bryologist 110:595–621
Qiu J (2010) China drought highlights future climate threats. Nature 465:142–143
Qiu X-Z, Xie S-C (1998) Studies on the forest ecosystem in Ailao Mountains, Yunnan, China. Yunnan Sciences and Technology Press, Kunming
Ramirez J, Jarvis A (2008) High resolution statistically downscaled future climate surfaces. International Center for Tropical Agriculture (CIAT); CGIAR Research Program on Climate Change, Agriculture and Food Security (CCAFS) Cali, Colombia. http://wwwccafs-climateorg/
Roberts SW, Strain BR, Knoerr KR (1981) Seasonal variation of leaf tissue elasticity in four forest tree species. Physiol Plant 52:245–250
Romero C, Putz FE, Kitajima K (2006) Ecophysiology in relation to exposure of pendant epiphytic bryophytes in the canopy of a tropical montane oak forest. Biotropica 38:35–41
Santarius KA (1994) Apoplasmic water fractions and osmotic potentials at full turgidity of some Bryidae. Planta 193:32–37
Shi J-P, Zhu H (2009) Tree species composition and diversity of tropical mountain cloud forest in the Yunnan, southwestern China. Ecol Res 24:83–92
Song L, Liu W-Y, Ma W-Z, Tan Z-H (2011) Bole epiphytic bryophytes on Lithocarpus xylocarpus (Kurz) Markgr. in the Ailao Mountains, SW China. Ecol Res 26:351–363
Song L, Liu W-Y, Ma W-Z, Qi J-H (2012a) Response of epiphytic bryophytes to simulated N deposition in a subtropical montane cloud forest in southwestern China. Oecologia 170:847–856
Song L, Liu W-Y, Nadkarni NM (2012b) Response of non-vascular epiphytes to simulated climate change in a montane moist evergreen broad-leaved forest in southwest China. Biol Conserv 152:127–135
Song L et al (2015) Bole bryophyte diversity and distribution patterns along three altitudinal gradients in Yunnan, China. J Veg Sci (published online). http://onlinelibrary.wiley.com/doi/10.1111/jvs.12263/abstract
Tyree M, Karamanos A (1981) Water stress as an ecological factor. In: Grace J, Ford E, Jarvis PG (eds) 21st symposium of British Ecological Society. Blackwell Scientific Publications, Oxford
Wagner S, Zotz G, Allen NS, Bader MY (2013) Altitudinal changes in temperature responses of net photosynthesis and dark respiration in tropical bryophytes. Ann Bot 111:455–465
Wang X-J (2010) System thinking for drought management in Southwest China. China Water Resour 7:11–13
Wolf JH (1993) Diversity patterns and biomass of epiphytic bryophytes and lichens along an altitudinal gradient in the northern Andes. Ann Mo Bot Gard 928–960
Wu Z-Y (ed) (1983) Research of forest ecosystem on Ailao Mountains, Yunnan. Yunnan Science and Technology Press, Kunming
Xu H-Q, Liu W-Y (2005) Species diversity and distribution of epiphytes in the montane moist evergreen broad-leaved forest in Ailao Mountain, Yunnan. Biodivers Sci 13:137–147
Ye Z-P (2007) A new model for relationship between irradiance and the rate of photosynthesis in Oryza sativa. Photosynthetica 45:637–640
Ye Z-P, Yu Q (2008) Comparison of new and several classical models of photosynthesis in response to irradiance. J Plant Ecol (Chinese Version) 32:1356–1361
Zhang Y-J, Yang Q-Y, Lee DW, Goldstein G, Cao K-F (2013) Extended leaf senescence promotes carbon gain and nutrient resorption: importance of maintaining winter photosynthesis in subtropical forests. Oecologia 173:721–730
Zotz G, Schweikert A, Jetz W, Westerman H (2000) Water relations and carbon gain are closely related to cushion size in the moss Grimmia pulvinata. New Phytol 148:59–67
Acknowledgments
This work was supported by the National Natural Science Foundation of China (31300382, U1133605, U1202234), CAS135 Program (XTBG-F01), West Light Foundation and the Research Foundation for members of the Youth Innovation Promotion Association of the Chinese Academy of Sciences to Liang Song. We thank Ailaoshan Station for Subtropical Forest Ecosystem Studies and Central Laboratory of Xishuangbanna Tropical Botanical Garden for Granting permission and facilitating this research. We are also grateful to Dr. Kouki Hikosaka, Mr. Steven Hunter, and the two anonymous reviewers for their constructive comments and corrections to the manuscript.
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Song, L., Zhang, YJ., Chen, X. et al. Water relations and gas exchange of fan bryophytes and their adaptations to microhabitats in an Asian subtropical montane cloud forest. J Plant Res 128, 573–584 (2015). https://doi.org/10.1007/s10265-015-0721-z
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DOI: https://doi.org/10.1007/s10265-015-0721-z