Shrubland Patch Formation Controlled by Shrub Development States

Project Number: 
Project Duration: 
24 months
May 1, 2001 to April 30, 2003
Institution of Principle Investigator while on this project: 
Ben-Gurion University of the Negev

Investigators (most current known information)

Professor, Ben-Gurion University of the Negev, Mitrani Center for Desert Ecology, Institute for Desert Research, Sede Boqer 84990, ISRAEL
TEL: +972-8-659-6786, FAX: +972-8-659-6772, Email:
Senior Lecturer, Department Head, Department of Evolution, Systematics and Ecology, Hebrew University of Jerusalem, Givat-Ram Campus, Jerusalem 91904, ISRAEL
TEL: +972-7-658-4305, FAX: +972-7-658-4741, Email:
Associate Professor, Biology, University of New Mexico, Albuquerque NM 87131
TEL: +1-505-277-2604, FAX: +1-505-277-6079, Email:

Proposal Abstract

In this report, the part of the research investigating patterns of shrub distribution within S. spinosum patch in relation to shrub and clone demography, is presented. Other parts of the study investigated the different developmental stages of S. spinosum and focused on the development and establishment of seedlings. Main findings showed that S. spinosum has a potential to reproduce vegetatively and this capability increases with age. Field measurements, field manipulations and pots experiment showed that moderate shading increases seedling survivorship in comparison to no shading, while strong shading inhibits seedling germination and decreases their survivorship. Moderate shading ameliorates seedling development, while with the increase of shading levels; seedlings develop shade tolerance and shade avoidance morphology. There are some indications, that the shade avoiding seedlings have an advantage in surviving the dry season.

These results imply that during the process of patch formation sexual recruits will establish within patch as long as the canopy of patch is not providing strong shade.

Since, developed canopy is related to the older stages of development plus vegetative reproduction might be related to the older stages too, there might be interplay between the possible patch expansion by clonal means and its expansion by sexual means. This part of the study aims to infer processes of patch formation from the spatial distribution of the different stages of development of individual shrubs and clones. Therefore, it also involves the study of S. spinosum clonal structure and expansion.


Please note that this list includes outcomes from related projects 94R-15, 96R-18, and 97D-03.

Articles in Journals

Reisman-Berman, O. 2007. Age related canopy traits shift facilitation to interference in semi-arid shrubland. Ecography 30: 459-470.

Reisman-Berman, O. and Z. Henkin. 2007. Sarcopterium Spinosum : revisiting shrub development and its relationship to space occupation with time. Israel Journal of Plant Sciences 55: 53-61.

Reisman-Berman, O., R. Kadmon, and M. Shachak. 2006. Spatio-temporal scales of dispersal limitations in the recolonization of a semi-arid Mediterranean old-field. Ecography 29(3); 418-426 IF3,4.

Gilad, E., J. von Hardenberg, A. Provenzale, M. Shachak and E. Meron. 2004. "Ecosystem engineers: From pattern formation to habitat creation." Phys. Rev. Lett. 93:098105(1-4).

Zaady, E., R. Levacov and M. Shachak. 2004. "Application of a chemical disturbance to biological soil crusts and its effect on resource flow in a patchy desert landscape." Arid Land Research and Management 18:397-410.

Meron, E., E. Gilad, J. von Hardenburg, M. Shachak and Y. Zarmi. 2003. "Vegetation patterns along a rainfall gradient."Chaos, Solitons and Fractals 19:367-376.

Zaady, E., P.M. Groffman, M. Shachak and A. Wilby. 2003. "Consumption and release of nitrogen of harvester termite Anacanthotermes ubachi navas in the northern Negev desert, Israel." Soil Biology and Biochemistry 35:1299-1303.

Eldridge, D.J., E. Zaady and M, Shachak. 2002. "Microphytic crusts, shrub patches and water harvesting in the Negev Desert: The Shikim system."Landscape Ecology 17:587-597.

Karnieli, A., A. Gabai, C. Ichoku, E. Zaady and M. Shachak. 2002. "Temporal dynamics of soil and vegetation spectral responses in a semiarid environment."International Journal of Remote Sensing 23:4073-4087.

Boeken, B. and D. Orenstein. 2001. "The effect of plant litter on ecosystem properties in a patchy Mediterranean semi-arid shrubland."Oikos 92:436-444.

von Hardenburg, J., E. Meron, M. Shachak and Y. Zarmi. 2001. "Diversity of vegetation patterns and desertification." The Physical Review Letters 87:198101-198104.

Wilby, A., M. Shachak and B. Boeken. 2001. "Integration of ecosystem engineering and trophic effects of herbivores." Oikos 92:436-444.

Zaady, E., Y.Z. Offer and M. Shahchak. 2001. "The content and contribution of the accumulated aeolian organic matter in a dry ecosystem." Atmospheric Environment 35:769-776.

Zaady, E., R. Yonatan, M. Shachak and A. Perevolotsky. 2001. "The effects of grazing on abiotic and biotic parameters in a semiarid ecosystem: a case study from the northern Negev desert, Israel." Arid Soil Research and Management 15:245-261.

Eldridge, J. D., Zaady, E. and M. Shachak. 2000. "Infiltration through three contrasting soil crusts in patterned Negev Desert landscapes."Catena 40:323-336.

Shachak,M. and G.M. Lovett. 1998. "Atmospheric deposition to a desert ecosystem and its implication for management". Ecological Applications 8:455-463.

Boeken, B. and M. Shachak. 1998. "Colonization by annual plants of an experimentally altered desert landscape: source-sink relationships."Journal of Ecology 86:804-814.

Boeken, B. and M. Shachak. 1998. "The dynamics of abundance and incidence of annual plant species during colonization in a desert." Ecography21:63-73.

Shachak, M., M. Sachs and I. Moshe. 1998. "Ecosystem management of desertified shrublands in Israel." Ecosystems 1:475-483.

Zaady, E., P. Groffman and M. Shachak. 1998. "Nitrogen fixation in macro- and microphytic patches in the Negev desert." Soil Biology & Biochemistry30:449-454.

Jones, C.G, J.H. Lawton and M. Shachak. 1997. "Positive and negative effects of organisms as ecosystem engineers." Ecology 78:1946-1957.

Offer Z. Y., E. Zaady and M. Shachak. 1997. "Aeolian particles input to the soil surface at the northern limit of the Negev desert." Arid Soil Research and Rehabilitation 2:55-62.

Karnielli, A., M. Shachak, H. Tsoar, E. Zaady, Y. Kaufman, A. Danin and W. Porter. 1996. "The effect of microphytes on the spectral reflectance of vegetation in semi-arid regions." Remote Sensing Environment57:88-96.

Zaady, E., P. Groffman and M. Shachak. 1996. "Litter as a regulator of nitrogen and carbon dynamics in macrophytic patches in Negev desert soils." Soil Biology & Biochemistry 28:39-46.

Zaady, E., P. Groffman and M. Shachak. 1996. "Release and consumption of snail feces in Negev desert soils." Biology & Fertility of Soils 23:399-404.

Boeken, B., M. Shachak, Y. Gutterman and S. Brand. 1995. "Patchiness and disturbance: plant community responses to porcupine diggings in the Central Negev." Ecography 18:410-422.

Boeken, B. and M. Shachak. 1994. "Desert plant communities in human made patches - implications for management." Ecological Applications4:702-716.

Zaady, E. and M. Shachak. 1994. "Microphytic soil crust and ecosystem leakage in the Negev Desert." American Journal of Botany 81:109.


Reisman-Berman, O., R. Kadmon, Z. Henkin, and M. Shachak. 2004. From seeding to landscape: an integrated study of Sarcopterium Spinosum . In MEDECOS X. proceedings of the 10th international conference on Mediterranean climate ecosystems. 25 April - 2 May 2004. ed. M. Arianoutsou. CD-ROM.

Gilad, E., J. von Hardenberg, E. Meron, M. Shachak and Y. Zarmi. 2004. "A dynamical system approach to aridity and desertification." In Continuum models and discrete systems, eds. D. Bergman et al, 405-418.

Zaady, E., M. Shachak and Y. Moshe. 2001. "Ecological Approach for Afforestation in Arid Regions of the Northern Negev Desert, Israel." In Deforestation, environment and sustainable development, a comparative analysis, ed. D. Vajpeyi, 219-238. Greenwood Publishing Group.

Eldridge, J. D., E. Zaady, M. Shachak and C. and Myers. 1999. "Control of desertification by microphytic crusts in a Negev desert shrubland."In Desertification and soil processes, the sixth international Rangeland Congress proceedings, 1:111-113.

Pickett, S. T. A., M. Shachak, B. Boeken and J. J. Armesto. 1999. "Management of ecological systems." In Arid lands management - Toward ecological sustainability, eds, T.W. Hoekstra and M. Shachak Hoekstra, 8-17. Urbana, IL. The Illinois University Press.

Saltz, D., M. Shachak, M. Caldwell, S.T.A. Pickett, J. Dawson, C. Tsoar, Y. Yom-Tov, M. Weltz and R. Farrow. 1999. "The study and management of dryland population systems." In Arid lands management - Toward ecological sustainability, eds. T.W. Hoekstra and M. Shachak, 75-96. Urbana, IL. The Illinois University Press.

Shachak, M., S. T. A. Pickett, B. Boeken and E. Zaady. 1999. "Managing patchiness, ecological flows, productivity and diversity in the Negev."In Arid lands management - Toward ecological sustainability, eds. T.W. Hoekstra and M. Shachak, 254-263. Urbana IL. The Illinois University Press.

Shachak, M. and S.T.A. Pickett. 1997. "Linking ecological understanding and application: patchiness in dry land system." In The ecological basis for conservation, eds. S.T.A. Pickett, R.S. Ostfeld, M. Shachak and G.E. Likens. New York. Chapman & Hall.

M.S Theses

Linde, L. 2000. The role of Thymelea hirsuta in redistribution of water on a watershed scale. M.S. Thesis. Leibzig, Germany.

Levacov, R. 1997. The effect of herbicides on resource flows in a patchy desert landscape. M.S. Thesis.

Orenstein, D. 1996. The effects of leaf litter on ecological systems in a dryland environment. M.S. Thesis.


Brandwine, S. 2004. The effect of water enriched patches on desert invertebrates. Ph.D. Dissertation.

Reisman, O. 2004. Colonization and patch formation by the desert shrub Sarcopoterium spinosum. Ph.D. Dissertation.

Kahn, B. 2003. The effect of livestock grazing on patchiness and ecosystem function in the northern Negev. Ph.D. Dissertation. University of Wisconsin. Madison WI.

Oren, Y. 1999. Experimental studies on the effect of grazing on a semi arid ecosystem in the northern Negev, Israel. Ph.D. Dissertation.


"Lehavim – bedouin demonstration and research site." 28 January 2005. Electronic Style. <> (28 January 2005).

"Ecology and the environment." 28 January 2005. Electronic Style. <> (28 January 2005).


Support for this project came from the USDA Cooperative State Research, Education, and Extension Service