The Botanical Review 65(1)The Botanical Review 65(2)
Interpreting Botanical Progress
April-June 1999

Desert Rocks - Plant Refugia in the Near East
    Avinoam Danin.................................................................93

N:P Balance in Wetland Forests: Productivity Across A 
  Biogeochemical Continuum
    B.G. Lockaby and W.H. Conner.................................................171

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Desert Rocks - Plant Refugia in the Near East
Department of Evolution, Systematics, and Ecology
The Alexander Silberman Institute of Life Sciences
The Hebrew University of Jerusalem
Jerusalem, Israel 91904

   I. Abstract
  II. Introduction
      A. Environmental Conditions in the Study Area 
         1. Topography
         2. Geomorphology and Edaphic Conditions
         3. Climate
      B. Flora
      C. Vegetation of Israel, Jordan, and Sinai
 III. Smooth-faced Rock Outcrops in Desert, Distribution
      and Significance as a Habitat for Plants
A. Fine-grained Particles in the Soil Pockets and Their Origin
      B. Limestone and Dolomite Rocks
      C. Magmatic and Metamorphic Rocks
      D. Sandstones
         1. Rock Varnish
         2. Cryptoendolithic Cyanobacteria and Green Algae
         3. Epilithic Crustose Lichens and Their Role in 
            Smooth-face Development
         4. Mosses and Their Role in the Sandstone 
      E. Chert (Flint)
  IV. The Impact of Rock, Soil, and Climate on Floristic 
      A. Number of Individuals
      B. Species Diversity 
      C. Rarity Index of Rock Species 
      D. Phytogeography
      E. Endemic Species
      F. Conclusions
   V. Major Key Refugia in the Near Eastern Deserts
      A. Nahal Elot, the Negev Highlands
      B. Gebel Halal, Northern Sinai
      C. Gebel Serbal, Southern Sinai
      D. Tafila - Petra - Ras en Naqb Area, Southwestern 
      E. Wadi Rum Area, Jordan 
  VI. Concluding Remarks
      A. Fitness of Population Structure of Rock Plants with Extinction Theories
      B. Climatic Changes and Floristic Disjunctions in the Saharan and Near Eastern Areas 
         1. Climatic Changes in Africa and Relicts in Central Saharan Mountains 
         2. Climatic Oscillations in the Near East since the Pleistocene 
      C. Conclusions
 VII. Acknowledgments
VIII. Literature Cited
    IX. Appendix: Plant Name Index
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I. Abstract
The outcrops of smooth-faced rocks in the Near Eastern deserts function as refugia 
for plants that do not fit the present local climate. They survived in the area from 
periods when the continuous extensions of the moister climate enabled their 
penetration from the Mediterranean zone. The largest Mediterranean enclave in the 
Near Eastern deserts and steppes is the sandstone and limestone outcrops at the 
upper escarpments of the southwestern Jordanian plateau, between At Tafila and Ras 
en Naqb, including the famous Petra and Wadi Dana. Hundreds of Mediterranean relict 
species and dozens of endemic species coexist in the crevices of these rocks with 
steppe and desert species. This paper discusses the ecology, phytogeography, and 
distribution of this special habitat in Jordan, Sinai, and Israel. Climatic 
oscillations during the Pleistocene and Holocene in the Near East and Africa are 
compared in their influence on past and present floras.
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N:P Balance in Wetland Forests: Productivity Across A Biogeochemical Continuum

School of Forestry
108 M.W. Smith Hall
Auburn University
Auburn, AL.  36849-5418 U.S.A. 

Baruch Forest Science Institute
P.O. Box 596
Clemson University
Georgetown, SC  29442 U.S.A.

*To whom all correspondence should be addressed.

   I. Abstract/Résumé
  II. Introduction
 III. Elemental Ratios as Indicators of Biogeochemical Balance
  IV. Oligotrophic vs Eutrophic Wetland Forests
   V. Aboveground Productivity vs Hydrology
  VI. N:P Balance and Deficiency in Wetland Forests
 VII. The N:P Balance Continuum in Wetland Forests
VIII. Concluding Remarks
  IX. Literature Cited

I. Abstract
The nature of and driving forces behind variation among wetland forests in terms of 
biogeochemistry and vegetation production are not well understood.  We suggest that 
insight into biogeochemical and productivity differences may be gained by examining 
the degree to which nitrogen and phosphorus are balanced within wetland vegetation.  
Based on examinations of data related to N/P balance and nutrient use efficiencies, 
vegetation productivity in both depressional and riverine forests appears to be 
primarily N limited.  In contrast to some current theories of wetland biogeochemistry, 
these data suggest that, when P deficiency occurs at all, it represents a secondary 
productivity constraint in comparison to N.  Similarly, a biogeochemical continuum 
is suggested for wetland forests based on the relationship between N/P ratios in 
senesced foliage vs. annual litterfall mass.  We theorize that the position of a 
particular wetland forest on this continuum reflects the integration of its geomorphic 
position and biogeochemical history.  In addition, the position of a particular 
system on the continuum may have predictive value with regard to net primary 
productivity and nutrient transformation capabilities.
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La nature et les forces agissantes à l'origine de la variation parmi les forêts 
marécageuses en ce qui concerne la biogéochimie et la production de la végétation 
ne sont pas bien comprises. Nous suggérons que la compréhension des différences 
entre la biogéochimie et la productivité peut être atteinte en examinant le degré 
auquel l'azote et le phosphore sont tenus en équilibre dans la végétation des 
marécages. Basé sur les études des données qui sont lieés à l'équilibre entre l'
azote et le phosphore (A/P) dans le feuillage et les efficacités de l'utilisation 
des substances nutritives, la productivité de la végétation dans les forêts 
dépressionelles et les forets fluviales semble être pour la plupart restreinte à l'
azote. Par opposition à quelques théories courantes de la biogéochimie des marécages, 
ces données suggèrent que, lorsque un manque de phosphore se produit, il représente 
une contrainte secondaire sur la productivité par rapport à l'azote. Et de même, un 
continuum biogéochimique est suggéré pour les forêts marécageuses fondé sur le 
rapport entre les proportions (A/P) pour les masses de feuillage tombées automnal 
et annuel. Nous émmetons l'hypothèse que la position d'une forêt marécageuse particulière 
sur ce continuum montre bien l'intégration entre sa position géomorphique et son 
histoire biogéochimique. En plus, la position d'un système particulier sur ce 
continuum pent avoir une valeur prévisible en ce qui concerne la productivité primaire 
nette et les capacités des substances nutritives de se transformer. 
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