TABLE OF CONTENTS
PREFACE X111
ACKNOWLEDGMENTS xv
CHAPTER 1. THE DESERT BlOME ....................................................................... 1
1.1. The Negev Desert of Israel: location, rains and phytogeographic regions ....... 1
1.1.1. SHRUBS AND ANNUALS ....................................................................... 6
1.1.2. GEOPHYTES AND HEMICRYPTOPHYTES (PERENNIAL
PLANTS) ................................................................................................. 7
1.1.3. ANNUAL PLANTS ................................................................................... 9
1.1.4. PORCUPINES IN THE NEGEV DESERT AND THE KAROO
DESERTS OF SOUTH AFRICA ............................................................ 10
1.1.5. THE INFLUENCE OF IBEX WALLOWS ON THE
VEGETATION ........................................................................................ 11
1.2 Amounts and distribution of rain in the Negev Desert highlands ...................... 11
1.3. Rains and runoff water ......................................................................................... 12
1.3.1 RAIN AND WATER INFILTRATION IN THE SOIL ............................. 12
1.3.2. DESERT CRYPTOGAMIC CRUST, RUNOFF WATER AND
SOIL EROSION ....................................................................................... 14
1.3.3. SLOPE ANGLE AND RUNOFF ............................................................... 15
1.3.4. DEPRESSIONS, MICRO-TOPOGRAPHY, RUNOFF, SEED
ACCUMULATION AND SEED GERMINATION ............................... 15
1.4. Evaporation rates, relative humidity and dew .................................................... 16
1.5. Seasonal temperatures ........................................................................................... 17
1.6. The temperature on the soil surface and in depressions as a regulator
for post maturation and germination of seeds ................................................. 17
1.7. Temperatures on the soil surface and in simulated porcupine diggings
during the year ................................................................................................... 18
l.7.l. THE LOCATIONS OF THE MICRO SENSORS ..................................... 18
1.7.2. TEMPERATURES DURING DAYS WITH RAIN .................................. 18
1.7.2.1. Temperatures in the digging in comparison with the soil
sUlface (locations 1 & 2) during a day with rain and one
day later . ................................................................................... 19
1.7.2.2. Temperatures in the diggings and in soil 10 em below the
soil surface (Locations 2 &4) ................................................... 20
1.7.2.3. Temperatures on soil sUrface and southern slope
(Locations 1 & 3) ...................................................................... 20
1.7.3. TEMPERATURES ON A HOT DAY ("HAMSIN") ................................ 20
1.7.3.1. Temperatures on the matrix in comparison with in the
digging (Locations 1 & 2) ......................................................... 20
1.7.3.2. Temperatures in the diggings and in soil 10 cm below the
soil swface (Locations 2 & 4) ................................................... 21

1.7.3.3. Temperatures on soil suiface and southern slope
(Locations 1 & 3) ...................................................................... 21
1.7.4. SOIL TEMPERATURES DURING THE HOT PERIOD OF THE
YEAR ....................................................................................................... 21
1.7.4.1. Temperatures on the matrix in comparison with in the
digging (Locations 1 & 2) ......................................................... 21
1.7.4.2. Temperatures in the diggings and in soil 10 em below the
soil suiface (Locations 2 & 4) ................................................... 24
1.7.4.3. Temperatures on the soil suiface and southern slope
(Locations 1 &3) ....................................................................... 24
1.7.5. SOIL TEMPERATURES DURING THE SHORTEST DAy .................. 24
1.7.6. TEMPERATURES AT TIME OF FIRST EFFICIENT RAINFALL
IN 1996/1997 SEASON ........................................................................... 26
1.8. Evaporation rates, temperatures and minimum amounts of rain for
germination ......................................................................................................... 26
1.9. The dynamics of depression filling by soil particles in different habitats ..•..... 29
1.10. Influence of the filling of the diggings by soil and organic matter on
water and seed accumulation, seed germination and seedling
establishment ...................................................................................................... 29
1.11. Depressions as wind traps for the accumulation of organic matter and
seeds. The 'treasure' effect ................................................................................. 31
1.12. Strategies of seed dispersal and the efficiency of the strategy of
mucilaginous seeds of some common annuals of the Negev that adhere
to the soil crust to delay collection by ants ....................................................... 32
1.13. Porcupine diggings and their influences on the vegetation in the Negev
Desert highlands ................................................................................................. 36
1.14. Ibex wallows and their influences on the annual vegetation in the
Negev Desert ....................................................................................................... 37
1.15. Hazeva Research Area ........................................................................................ 38
1.16. Small animal disturbances .................................................................................. 39
1.17. Ant disturbances .................................................................................................. 39
1.18. Isopods (Hemilepistus reaumuri) and porcupine diggings as a source of
erodible soil ......................................................................................................... 39
1.19. Footprints in muddy loess as long term depressions ........................................ 40
1.20 Main habitats of the Central Negev and their influence on vegetation
and porcupine activity ....................................................................................... 40
1.21. Inter-relations and ecological significance of diggings and open burrow
systems ...................................................................................................................... 41
1.21.1. DIGGINGS .............................................................................................. 41
1.21.2. ECOLOGY AND OPEN BURROW SYSTEMS IN DESERTS
AND ARID ENVIRONMENTS .............................................................. 43
CHAPTER 2. DIGGINGS AND CONSUMPTION OF SUBTERRANEAN
PLANT ORGANS ................................................................................ 43
2.1 . The porcupine and its habitat ............................................................................. 43
2.2. The influences of porcupine activity on species of geophytes and
hemicryptophytes ............................................................................................... 43
2.2.l. PLANTS THAT ARE DESTROYED BY PORCUPINE
CONSUMPTION AND RENEWED BY SEED GERMINATION
IN THE PORCUPINE DIGGING (GROUP 1) ........................................ 52
2.2.1.1. Colchicum tunicatum ................................................................... 52
2.2.1.2. Bellevalia desertorum .................................................................. 61
2.2.1.3. Erodium crassifolium ............................................ ....................... 65
2.2.2. PLANT SPECIES PARTIALLY CONSUMED THAT ARE
RENEWED VEGET A TIVEL Y IN THE PORCUPINE DIGGING
(GROUP 2) ............................................................................................... 68
2.2.2.1. Bellevalia eigii ............................................................................. 69
2.2.2.2. Tulipa systola ............................................................................... 72
2.2.2.3. Malabaila secacul ....................................................................... 75
2.2.2.4. Scorzonera papposa ............................................... ...................... 78
2.2.2.5. Leontice leontopetalum ............................................................... 80
2.2.3. PLANTS OCCASIONALLY TOTALLY CONSUMED AND
RENEWED BY SEED GERMINATION IN PORCUPINE
DIGGINGS (GROUP 3) ........................................................................... 80
2.2.3.1. Tulipa polychroma ....................................................................... 80
2.2.3.2. Allium rothii ................................................................................. 81
2.2.3.3. Gynandriris sisyrinchium ............................................................ 82
2.2.4. PLANTS OCCASIONALLY PARTIALLY CONSUMED AND
RENEWED IN DIGGINGS (GROUP 4) ................................................. 83
2.2.4.1. Urginea undulata .............................................. ........................... 83
2.2.4.2. Asphodelus ramosus .................................................................... 83
2.2.4.3. Anemone coronaria ..................................................................... 83
2.2.5. PLANT SPECIES RARELY CONSUMED AND VERY FEW OF
THEM FOUND IN PORCUPINE DIGGINGS (GROUP 5) ................... 84
2.2.5.1. Scilla hanburyi ............................................................................. 84
2.2.5.2. Narcissus tazetta .......................................................................... 84
2.2.5.3. Sternbergia clusiana ..................................................................... 87
2.2.5.4. Colchicum richii .......................................................................... 90
2.2.6. PLANT SPECIES FOUND IN PORCUPINE DIGGINGS BUT NOT
YET OBSERVED AS BEING CONSUMED BY PORCUPINES ......... 91
2.2.6.1. Ixiolirion tataricum ..................................................................... 91
2.2.6.2. Eryngium cretiCUI11 ....................................................................... 93
2.2.6.3. Ranunculus asiaticus ................................................................... 93
2.2.6.4. Crocus damascenus ..................................................................... 93
2.2.6.5. Iris regis-uzziae ........................................................................... 93
2.2.6.6. Allium pallens .............................................................................. 94
2.2.6.7. Ornithogalum trichophyllum ....................................................... 94
2.2.6.8. Ornithogalum narbonense ........................................................... 95
2.2.6.9. Leopoldia longipes ..................................................................... 95

2.2.7. SEASONS IN WHICH THE SUBTERRANEAN ORGANS OF
DIFFERENT GEOPHYTES AND HEMICRYPTOPHYTES ARE
CONSUMED AND REAPPEAR IN THE DIGGINGS .......................... 95
2.2.7.1. Group 1 ........................................................................................ 96
2.2.7.2. Group 2 ........................................................................................ 97
2.2.7.3. Group 3 ........................................................................................ 97
2.2.7.4. Groups 4 and 5 ............................................................................ 97
2.2.8. HABITAT AND SITE DISTRIBUTION OF GEOPHYTES ................... 97
2.2.9. STRATEGIES OF CONSUMPTION: THE PERCENTAGE OF
PORCUPINE DIGGINGS AND PLANT CONSUMPTION IN
PA TCHES OF DIFFERENT SPECIES OF GEOPHYTES AND
HEMICRYPTOPHYTES ......................................................................... 98
2.2.10. THE DYNAMICS OF FILLING-IN OF DIGGINGS ............................ 99
2.2.10.1. Long term filling-in ................................................................... 100
2.2.10.2. Intermediate tennfilling-in ....................................................... 100
2.2.10.3. Short termfilling-in .......................................... ......................... 101
2.2.11. DIGGINGS, FILLING-IN AND THEIR INFLUENCE ON THE
VEGETATION ......................................................................................... 101
2.3. Porcupines and the damage they cause to agricultural crops in desert
areas ..................................................................................................................... 102
2.4. The digging of the African porcupine in deserts ................................................. 102
2.5. Gazelles dig for geophytes ..................................................................................... 102
2.6. Ibex wallows ......................................................................................................... 104
2.7. Sand partridges and geophyte consumption ....................................................... 104
2.8. Subterranean herbivorous rodent activity and its influence on geophytes ...... 104
2.9 The ecological interrelations between porcupine consumption strategies
and the survival of geophytes and hemicryptophytes in the Negev Desert
highlands ............................................................................................................. 105
CHAPTER 3. ANNUALS IN PORCUPINE DIGGINGS AND IBEX
W ALLOWS IN THE NEGEV DESERT HIGHLANDS ................. 107
3.1. The annual plant species and their occurrence ................................................... 107
3.2. Annuals and perennials in the Negev Desert environment ................................ 109
3.2.1. PERENNIALS AND THE NEGEV DESERT ENVIRONMENT ........... 109
3.2.2. LIFE FORM DIVERSITY AND RAIN AMOUNTS ............................... 109
3.2.3. ANNUALS OCCURRING IN SMALL DEPRESSIONS ........................ 109
3.3. Depressions as micro-habitats in a desert biome ................................................ 110
3.3.1. DEPRESSIONS AS SEED TRAPS .......................................................... 110
3.3.1.1. The shape, depth and direction of digging .................................. 113
3.3.1.2. Distances between diggings and patches .................................... 114
3.3.1.3. The structure of the bottom of the digging, plant remnants
and orientation .......................................................................... 114
3.3.1.4. Influence of the topography and the plant or stone cover
near the digging ........................................................................ 114
3.3.1.5. Protected aerial and soil seed banks located in the diggings ..... 114
3.3.2. THE WATER CONTENT IN DIGGINGS AND THE
OCCURRENCE OF ANNUALS ............................................................. 117

3.3.3. TOPOGRAPHY, LOCATIONS AND SEEDLING NUMBERS ............. 119
3.3.3.1. General events ............................................................................. 119
3.3.3.2 Particular plant species diversity in diggings in comparison
with the n1atrix .......................................................................... 119
3.3.4. DIGGINGS AS FAVOURABLE MICRO-HABITATS FOR
ANNUALS ............................................................................................... 120
3.4. Digging depth, shape, topography, location and habitat, the species that
are consumed and the emergence of annuals ................................................... 122
3.5. The succession of annual plants in porcupine diggings - a system of
disturbance and the stages of recovery ............................................................ 124
3.5.1. CHANGES IN ANNUAL PLANT NUMBERS PER UNIT AREA
IN THE DIGGINGS ................................................................................. 125
3.5.2 CHANGES IN PLANT BIOMASS ........................................................... 128
3.5.3. PLANT COMPETITION AND CHANGES IN SEED YIELD ............... 129
3.5.4. CHANGES IN SPECIES DIVERSITy .................................................... 129
3.5.5. CHANGES IN PLANT SPECIES DOMINANCE ................................... 129
3.6. Density of annuals and species diversity in porcupine diggings along a
watershed according to runoff water gradients .............................................. 130
3.7. The influences of four artificial digging densities and slope directions on
annual plant density in three growing seasons ................................................ 137
3.7.1. ESTIMATE OF SEED NUMBERS IN A DIGGING ............................... 137
3.7.2. PLANT DENSITY .................................................................................... 138
3.7.3. PLANT BIOMASS ................................................................................... 139
3.7.4. EFFECT ON BIOMASS OF SLOPE DIRECTION AND
POSITION OF DIGGING IN A PATCH ................................................. 139
3.7.5. SLOPE DIRECTION AND FREQUENCY OF OCCURRENCE
OF ANNUAL PLANT SPECIES IN DIGGINGS AND ON THE
MATRIX IN DIFFERENT YEARS ......................................................... 140
3.7.5.1. On Loess soil ............................................................................... 140
3.7.5.2. On stabilized sand ....................................................................... 141
3.8. The influence of depressions made by ibex on the annual vegetation
along cliffs of the Zin valley in the Negev Desert highlands ........................... 143
3.9. Footprints in muddy soil as favourable micro-habitats for annuals in
deserts .................................................................................................................. 152
3.10. Environmental influences on life cycles and survival strategies of
annual plant species ........................................................................................... 156
3.10.1. FLOWERING STRATEGIES OF SOME WINTER ANNUALS ......... 156
3.10.1.1. Daylength,flowering time and life span offacultative
long-day plants for flowering .................................................... 156
3.10.1.2. Soil water content and life span of day-neutral plants for
flowering ................................................................................... 157
3.10.2. PHENOTYPIC PLASTICITY EFFECTS ON GERMINATION
DURING SEED DEVELOPMENT AND MATURATION ................... 157
3.10.2.1. Daylength influences ................................................................. 157
3.10.2.2. Position effect ............................................................................ 158

3.10.3. SEED POST MATURATION TEMPERATURES ON THE SOIL
SURFACE OR IN DEPRESSIONS, AND THE REGULATION OF
GERMINATION DURING THE HOT AND DRY SUMMERS ............ 166
3.10.4. SEED DISPERSAL STRATEGIES AND MASSIVE SEED
CONSUMPTION IN DESERTS .............................................................. 169
3.10.4.1. Subterranean synaptospermic protected inflorescences or
propagules and germination in situ . ......................................... 174
3...10.4.2. Serotinous seed dispersal strategy ............................................. 175
3.10.4.3. Dispersal offree mucilaginous seeds and single seed
protection in lignified beak .. ..................................................... 179
3.10.4.4. Mucilaginous seeds that float into depressions ......................... 180
3.10.4.5. Aerial seed bank with serotiny protection strategy
followed by soil seed bank with escape strategy ......... .............. 185
3.10.4.6. The decrease in the "treasure effect" by local protected
seed banks in depressions ......................................................... 186
3.10.4.7. Synaptospemlic seed protection strategies by lignified
dispersal units or inflorescences on lignified dry plants .......... 186
3.10.4.8. 'Escape' seed dispersal strategies and mechanisms of
plant species producing large numbers of dust-like seeds ........ 187
3.10.4.9. Escape' seed dispersal ofplant species producing tiny
mucilaginous seeds that are dispersed by wind . ....................... 190
3.10.5. SEED GERMINATION, DIGGINGS AND DEPRESSIONS AS
MICRO SITES ......................................................................................... 190
3.10.5.1. Water and germination ............................................ .................. 191
3.10.5.2. Temperatures and germination of annuals in depressions ........ 191
3.10.5.3. The temperatures during the season with rain and the
germination of annuals in the Negev Desert highlands ............ 192
3.10.5.4. Light, seed cover-over and germination .................................... 192
3.10.5.5 Light inhibition of germination and regulation of the
germination by the time of the beginning of seed wetting by
rain that engenders germination .. ............................................. 193
3.10.5.6. Slow germinating seeds ............................................................. 194
3.10.5.7 Fast germinating seeds . .............................................................. 194
3.10.5.8. Intermediate time of seed germination ...................................... 194
3.10.6. SEEDLING DROUGHT TOLERANCE ............................................... 195
3.11. Biopedturbation in deserts .................................................................................. 196
CHAPTER 4. EFFECTS OF HUMAN MADE DISTURBANCES AND
AGRICULTURE ON FLOOD AND RUNOFF WATER IN
THE NEGEV ......................................................................................... 199
4.1. Influences of human-made disturbances in the soil on annuals in the
Negev Desert ....................................................................................................... 199
4.2. Human made pits and mounds, and desert plant communities ........................ 199
4.2.1. THE SOIL WATER CONTENT OF THE PITS, MOUNDS AND
MATRIX ................................................................................................... 201
4.2.2. PLANT DENSITY .................................................................................... 201

4.2.3. PLANT SPECIES DIVERSITy ............................................................... 202
4.2.4. PLANT BIOMASS ................................................................................... 202
4.2.5. PROPAGULE SIZE AND PLANT APPEARANCE ............................... 204
4.2.6. CHANGES IN NUMBER OF PLANT SPECIES OVER
SEVERAL yEARS .................................................................................. 204
4.2.6.1. The amount of seasonal rain and number of plant species
in pits, mounds and matrix ........................................................ 206
4.2.6.2. Plant density affected by colonization during the previous
year ........................................................................................... 206
4.2.6.3. Seed dispersal strategies and plant occurrence .......................... 206
4.2.7. CONCLUSION ......................................................................................... 206
4.3. From small depressions to runoff agriculture using flood and runoff
water in the Negev .............................................................................................. 206
4.3.1. PREHISTORIC BACKGROUND ............................................................ 207
4.3.2. ANCIENT TERRACES IN WADIS USING FLOOD WATER
FOR AGRICULTURE ............................................................................. 208
4.3.3. FLOOD AND RUNOFF AGRICULTURE .............................................. 209
4.3.4. RUNOFF WATER AGRICULTURE IN MACRO-CATCHMENTS
IN ANCIENT NABATEAN FAMILY FARMS IN WIDE
VALLEYS ................................................................................................ 209
4.3.5. RUNOFF AGRICULTURE ON MODERATE LOESSIAL HILL
SLOPES IN MICRO-CATCHMENTS (NEGARIN) .............................. 210
4.3.6. FLOOD WATER AGRICULTURE IN V ALLEYS ALONG WADI
BANKS USING BY-PASS CHANNELS FROM THE WADI ............... 213
4.3.7. CONCLUSION ......................................................................................... 213
REFERENCES
PLANT INDEX
SUBJECT INDEX
ANIMALS AND INSECT INDEX
 

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