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Ecophysiological importance of cloud immersion in a relic spruce–fir forest at elevational limits, southern Appalachian Mountains, USA

  • Physiological ecology - Original research
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Abstract

Climate warming predicts changes to the frequency and height of cloud-immersion events in mountain communities. Threatened southern Appalachian spruce–fir forests have been suggested to persist because of frequent periods of cloud immersion. These relic forests exist on only seven mountaintop areas, grow only above ca. 1,500 m elevation (maximum 2,037 m), and harbor the endemic Abies fraseri. To predict future distribution, we examined the ecophysiological effects of cloud immersion on saplings of A. fraseri and Picea rubens at their upper and lower elevational limits. Leaf photosynthesis, conductance, transpiration, xylem water potentials, and general abiotic variables were measured simultaneously on individuals at the top (1,960 m) and bottom (1,510 m) of their elevation limits on numerous clear and cloud-immersed days throughout the growing season. The high elevation sites had 1.5 as many cloud-immersed days (75 % of days) as the low elevation sites (56 % of days). Cloud immersion resulted in higher photosynthesis, leaf conductance, and xylem water potentials, particularly during afternoon measurements. Leaf conductance remained higher throughout the day with corresponding increases in photosynthesis and transpiration, despite low photon flux density levels, leading to an increase in water potentials from morning to afternoon. The endemic A. fraseri had a greater response in carbon gain and water balance in response to cloud immersion. Climate models predict warmer temperatures with a decrease in the frequency of cloud immersion for this region, leading to an environment on these peaks similar to elevations where spruce–fir communities currently do not exist. Because spruce–fir communities may rely on cloud immersion for improved carbon gain and water conservation, an upslope shift is likely if cloud ceilings rise. Their ultimate survival will likely depend on the magnitude of changes in cloud regimes.

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References

  • Arthur FH, Hain FP (1986) Water potential of Fraser fir infested with balsam wooly adelgid (Homoptera, Adelgidae). Environ Entolmol 15:911–913

    Google Scholar 

  • Bader M, Geloof I, Rietkerk M (2007) High solar radiation hinders tree regeneration above alpine treeline in northern Ecuador. Plant Ecol 191:33–45

    Article  Google Scholar 

  • Berry ZC, Smith WK (2012) Cloud pattern and water relations in Picea rubens and Abies fraseri, southern Appalachian Mountains, USA. Agric For Meteorol 162:27–34

    Article  Google Scholar 

  • Braun EL (1964) Deciduous forests of eastern North America. Hafner, New York

    Google Scholar 

  • Brewer CA, Smith WK (1997) Patterns of leaf surface wetness for montane and subalpine plants. Plant Cell Environ 20:1–11

    Article  Google Scholar 

  • Brient F, Bony S (2012) Interpretation of the positive low-cloud feedback predicted by a climate model under global warming. Clim Dynam 29:501–520. doi:10.1007/s00382-011-1279-7

    Google Scholar 

  • Brodribb TJ, Holbrook NM (2004) Diurnal depression of leaf hydraulic conductance in a tropical tree species. Plant Cell Environ 27:820–827

    Article  Google Scholar 

  • Bruck RI, Robarge WP (1988) Change in forest structure in the boreal montane ecosystem of Mount Mitchell, North Carolina. Eur J For Pathol 18:357–366

    Article  Google Scholar 

  • Bruijnzeel LA, Veneklaas EJ (1998) Climatic conditions and tropical montane forest productivity: the fog has not lifted yet. Ecology 79:3–9

    Article  Google Scholar 

  • Bruijnzeel LA, Scatena FN, Hamilton LS (2010) Tropical montane cloud forests: science for conservation and management. Cambridge University Press, New York

    Google Scholar 

  • Burgess SSO, Dawson TE (2004) The contribution of fog to the water relations of Sequoia sempervirens (D. Don): foliar uptake and prevention of dehydration. Plant Cell Environ 27:1023–1034

    Article  Google Scholar 

  • Busing RT, Clebsch EEC, Eagar CC, Pauley EF (1988) Two decades of change in a Great Smoky Mountains spruce–fir forest. B Torrey Bot Club 115:25–31

    Article  Google Scholar 

  • Campbell GS, Norman JM (1998) An Introduction to environmental biophysics. Springer, New York

    Book  Google Scholar 

  • Clark JS, Beckage B, Camill P, Cleveland B, HilleRisLambers J, Lichter J, McLachlan J, Mohan J, Wyckoff P (1999) Interpreting recruitment limitation in forests. Am J Bot 86:1–16

    Article  PubMed  CAS  Google Scholar 

  • Cogbill CV, White PS (1991) The latitude–elevation relationship for spruce–fir and treeline along the Appalachian mountain chain. Vegetation 94:153–175

    Article  Google Scholar 

  • Croke MS, Cess RD, Hameed S (1999) Regional cloud cover change associated with global climate change: case studies for three regions of the United States. J Climate 12:2128–2134

    Article  Google Scholar 

  • Day ME, Greenwood MS, White AS (2001) Age-related changes in foliar morphology and physiology in red spruce and their influence on declining photosynthetic rates and productivity with tree age. Tree Physiol 16:1195–1204

    Article  Google Scholar 

  • Delcourt HR, Delcourt PA (1984) Late-Quaternary history of the spruce–fir ecosystem in the southern Appalachian Mountain region. In: White PS (ed) The southern Appalachian spruce–fir ecosystem: its biology and threats. Research/resource management report SER-71. USDI, National Park Service, Southeast Regional Office, Atlanta, pp 22–35

  • Dengel S, Grace J (2010) Carbon dioxide exchange and canopy conductance of two coniferous forests under various sky conditions. Oecologia 164:797–808

    Article  PubMed  Google Scholar 

  • Foster P (2001) The potential negative impacts of global change on tropical montane cloud forests. Earth-Sci Rev 55:73–106

    Article  Google Scholar 

  • Germino MJ, Smith WK, Resor AC (2002) Conifer seedling distribution and survival in an alpine-treeline ecotone. Plant Ecol 162:157–168

    Article  Google Scholar 

  • Goelz JCG, Burk TE, Zedaker SM (1999) Long-term growth trends of red spruce and Fraser fir at Mt. Rogers, Virginia, and Mt. Mitchell, North Carolina. For Ecol Manage 115:49–59

    Article  Google Scholar 

  • Graham EA, Mulkey SS, Kitajima K, Phillips NG, Wright SJ (2003) Cloud cover limits net CO2 uptake and growth of a rainforest tree during tropical rainy seasons. Proc Natl Acad Sci USA 100:572–576

    Article  PubMed  CAS  Google Scholar 

  • Greenwood MS, O’Brien CL, Schatz JD, Diggins CA, Day ME, Jacobson GL, White AS, Wagner RG (2008) Is early life cycle success a determinant of the abundance of red spruce and balsam fir? Can J For Res 38:2295–2305

    Article  Google Scholar 

  • Gregory J, Webb M (2008) Tropospheric adjustment induces a cloud component in CO2 forcing. J Climate 21:58–71

    Article  Google Scholar 

  • Gu L, Baldocchi D, Verma SB, Black TA, Vesala T, Falge EM, Dowty PR (2002) Advantages of diffuse radiation for terrestrial ecosystem productivity. J Geophys Res 107. doi:10.1029/2001JD001242

  • IPCC (2007) In: Solomon S, Qin D, Manning M, Chen Z, Marquis M, Avery KB, Tignor M, Miller HL (eds) Climate change 2007: The physical science basis contribution of working group I to the fourth assessment report of the intergovernmental panel on climate change, Cambridge University Press, Cambridge

  • Ishibashi M, Terashima I (1995) Effects of continuous leaf wetness on photosynthesis: adverse aspects of rainfall. Plant Cell Environ 18:431–438

    Article  Google Scholar 

  • Ishii H, Hamada Y, Utsugi H (2012) Variation in light-intercepting area and photosynthetic rate of sun and shade shoots of two Picea species in relation to the angle of incoming light. Tree Physiol 32:1227–1236

    Article  PubMed  CAS  Google Scholar 

  • Johnson DM, Smith WK (2006) Low clouds and cloud immersion enhance photosynthesis in understory species of a southern Appalachian spruce–fir forest (USA). Am J Bot 93:1625–1632

    Article  PubMed  Google Scholar 

  • Johnson DM, Smith WK (2008) Cloud immersion alters microclimate, photosynthesis and water relations in Rhododendron catawbiense and Abies fraseri seedlings in the southern Appalachian Mountains, USA. Tree Physiol 28:385–392

    Article  PubMed  Google Scholar 

  • Johnson DM, Woodruff DR, McCulloh KA, Meinzer FC (2009) Leaf hydraulic conductance, measured in situ, declines and recovers daily: leaf hydraulics, water potential and stomatal conductance in four temperate and three tropical tree species. Tree Physiol 29:879–887

    Article  PubMed  CAS  Google Scholar 

  • Knapp AK, Smith WK (1990) Stomatal and photosynthetic responses to variable sunlight. What’s new in plant physiology. Physiol Plant 78:160–165

    Article  Google Scholar 

  • Letts MG, Mulligan M (2005) The impact of light quality and leaf wetness on photosynthesis in north-west Andean tropical montane cloud forest. J Trop Ecol 21:549–557

    Article  Google Scholar 

  • Mark AF (1958) The ecology of the southern Appalachian grass balds. Ecol Monogr 28:293–336

    Article  Google Scholar 

  • McLaughlin SB, Anderson CP, Edwards NT, Roy WK, Layton PA (1990) Seasonal patterns of photosynthesis and respiration of red spruce saplings from two elevations in declining southern Appalachian stands. Can J For Res 20:485–495

    Article  CAS  Google Scholar 

  • Min Q (2005) Impacts of aerosols and clouds on forest–atmosphere carbon exchange. J Geophys Res 110:D06203

    Article  Google Scholar 

  • Mohnen VA (1992) Atmospheric deposition and pollutant exposure of eastern US forests. Ecol Stud An 96:64–124

    Article  Google Scholar 

  • Morison JIL, Jarvis PG (1983) Direct and indirect effects of light on stomata. II. In Commelina communis L. Plant Cell Environ 6:103–109

    Article  Google Scholar 

  • Niinemets U (2002) Stomatal conductance alone does not explain the decline of foliar photosynthetic rates with increasing tree age and size in Picea abies and Pinus sylvestris. Tree Physiol 22:515–535

    Article  PubMed  Google Scholar 

  • Oosting HJ, Billings WD (1951) A comparison of virgin spruce–fir forest in the northern and southern Appalachian system. Ecology 32:84–103

    Article  Google Scholar 

  • Pittillo JD, Smathers GA (1979) Phytogeography of the Balsam Mountains and Pisgah Ridge, southern Appalachian Mountains. Ver Geobot 68:206–245

    Google Scholar 

  • Pounds JA, Fogden MPL, Campbell JH (1999) Biological response to climate change on a tropical mountain. Nature 398:611–615

    Article  CAS  Google Scholar 

  • Ramseur GS (1960) The vascular flora of high mountain communities of the southern Appalachians. J Elisha Mitch Sci S 76:81–112

    Google Scholar 

  • Reed JE, Smith WK (2012) Stomatal frequency, distribution, and needle hydrophobicity in cloud forest spruce and fir, southern Appalachian mountains. Rurals Rev Undergrad Res Agric Life Sci 7:Article3

    Google Scholar 

  • Reinhardt K, Smith WK (2008a) Impacts of cloud immersion on microclimate, photosynthesis, and water relations of Abies fraseri (Pursh.) Poiret in a temperate mountain cloud forest. Oecologia 158:229–238

    Article  PubMed  Google Scholar 

  • Reinhardt K, Smith WK (2008b) Leaf gas exchange of understory spruce–fir saplings in relict cloud forests, southern Appalachian Mountains, USA. Tree Physiol 28:113–122

    Article  PubMed  CAS  Google Scholar 

  • Richardson AD, Denny EG, Siccama TG, Lee X (2003) Evidence for a rising cloud ceiling in eastern North America. J Climate 16:2093–2098

    Article  Google Scholar 

  • Ruddiman WF (2008) Causes of warming over the last 125 years. In: Earth’s climate past and future. Freeman, New York, pp 325–342

  • Simonin KA, Santiago LS, Dawson TE (2009) Fog interception by Sequoia sempervirens (D. Don) crowns decouples physiology from soil water deficit. Plant Cell Environ 32:882–892

    Article  PubMed  Google Scholar 

  • Smith WK, McClean TM (1989) Adaptive relationship between leaf water repellency, stomatal distribution and gas exchange. Am J Bot 76:465–469

    Article  Google Scholar 

  • Smith WK, Schoettle AW, Cui M (1991) Importance of leaf area measurement to the interpretation of gas exchange parameters of complex shoots. Tree Physiol 8:121–127

    Article  PubMed  Google Scholar 

  • Smith WK, Germino MJ, Hancock TE, Johnson DM (2003) Another perspective on the altitudinal limits of alpine timberline. Tree Physiol 23:1101–1112

    Article  PubMed  Google Scholar 

  • Still CJ, Foster PN, Schneider SH (1999) Simulating the effects of climate change on tropical montane cloud forests. Nature 398:608–610

    Article  CAS  Google Scholar 

  • Urban O, Janouš D, Acosta M, Czerný R, Marková I, Navrátil M, Pavelka M, Pokorný R, Šprtová M, Zhang R, Špunda V, Grace J, Marek ML (2007) Ecophysiological controls over net ecosystem exchange of mountain spruce stand. Comparison of the response in direct vs. indirect diffuse solar radiation. Glob Change Biol 13:157–168

    Article  Google Scholar 

  • Urban O, Klem K, Ač A, Havránková K, Holišová P, Navrátil M, Zitová M, Kozlová K, Pokorný R, Šprtová M, Tomášková I, Špunda V, Grace J (2012) Impact of clear and cloudy sky conditions on the vertical distribution of photosynthetic CO2 uptake within a spruce canopy. Func Ecol 26:46–55

    Article  Google Scholar 

  • White PS (1984) The southern Appalachian spruce–fir ecosystem, an introduction. In: White PS (ed) The southern Appalachian spruce–fir ecosystem: its biology and threats. USDI, National Park Service, Southeast Regional Office, Research/Resource Management Report SER-71, Atlanta, pp 1–21

  • White PS, Cogbill CV (1992) Spruce–fir forests of eastern North America. In: Eager C, Adams MB (eds) Ecology and decline of red spruce in the Eastern United States. Springer-Verlag, New York, pp 3–39

    Chapter  Google Scholar 

  • Whittaker RH (1956) Vegetation of the Great Smoky Mountains. Ecol Monogr 26:1–80

    Article  Google Scholar 

  • Young DR, Smith WK (1983) Effect of cloudcover on photosynthesis and transpiration in the subalpine understory species Arnica latifolia. Ecology 64:681–687

    Article  Google Scholar 

  • Zar JH (1999) Biostatistical analysis. Prentice-Hall, Upper Saddle River

    Google Scholar 

Download references

Acknowledgments

Support was provided through a grant from the National Science Foundation (IOS 1122092), plus a Vecellio grant to Z. C. Berry through the Biology Department, Wake Forest University. Research was conducted under special activity permit no. R11-22 from the North Carolina Department of Environment and Natural Resources. Thanks to Lisa Crane and Brian Wilder of Mount Mitchell State Park for project insight and cooperation, to Jenny Reed for field assistance, and to Katherine D. Hitzhusen for manuscript advice.

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Correspondence to Z. Carter Berry.

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Communicated by Ram Oren.

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Berry, Z.C., Smith, W.K. Ecophysiological importance of cloud immersion in a relic spruce–fir forest at elevational limits, southern Appalachian Mountains, USA. Oecologia 173, 637–648 (2013). https://doi.org/10.1007/s00442-013-2653-4

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