Tree-shrub interactions in a subtropical savanna parkland: competition or facilitation?


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Abstract. Prosopis glandulosa var. glandulosa has played a central role in the encroachment of woody plants in southern Texas, grasslands and savannas by acting as a nurse plant for various shrubs that establish in its understory. To test for continued facilitation of established understory shrubs by Prosopis and to determine if established shrubs compete with the Prosopis nucleus, selective removal experiments were conducted and monitored over a 2–5 yr period. Short-term (1–3 days) and long-term (2 yr) growth and physiological activities (midday net photosynthesis and leaf/shoot water potential) of two common understory shrubs, Zanthoxylum fagara and Berberis trifoliolata, growing with Prosopis, were generally comparable to those of individuals occurring in clusters where Prosopis was removed. Shrubs growing with an intact Prosopis occasionally showed significantly higher leaf-[N] and pre-dawn water potentials than those in clusters lacking a live Prosopis, especially under drought conditions; however, these differences did not translate into greater midday leaf gas exchange or shoot growth. By comparison, removal of understory shrubs elicited large increases in Prosopis net photosynthesis, annual trunk growth in each of the 5 yr monitored, and seed pod production in three of the four years monitored. Seven of 26 Prosopis plants in experimental clusters with an intact understory died over a 5-yr period, compared to only two of the 26 plants in clusters with the cleared understory.

Results indicate that (1) the founding overstory Prosopis plant may continue to facilitate understory shrubs following their establishment, but these beneficial effects appear to be small and transitory, and (2) the understory shrubs have a pronounced negative effect on Prosopis, such that competition between overstory and understory woody plants is strongly asymmetrical. These findings suggest that understory shrubs will likely persist despite changes in microclimate and soils (potentially) that occur after the Prosopis plant, which facilitated their ingress or establishment, has died. Soil resource depletion by shallow-rooted understory shrubs appears to be a primary factor contributing to the demise of the deeply rooted, overstory Prosopis plants, especially on upland sites with duplex soils where below-ground competition is accentuated.