Impediments to forest recovery in bracken-dominated clearings in the African Highlands

Philosophiae Doctor (PhD) Thesis 2018:70

Fredrick Ssali

Impediments to forest recovery in bracken-dominated clearings in the African Highlands

Einstape i afrikanske høylandsområder:

hvordan bregner hindrer gjenvekst av skog

Norwegian University of Life Sciences Faculty of Environmental Sciences and Natural Resource Management

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Paper I

Contents lists available atScienceDirect

Forest Ecology and Management

journal homepage:www.elsevier.com/locate/foreco

A first look at the impediments to forest recovery in bracken-dominated clearings in the African Highlands

Fredrick Ssali^⁎, Stein R. Moe, Douglas Sheil

Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life Sciences, P.O. Box 5003, 1432Ås, Norway

A R T I C L E I N F O

Keywords:

Afro-tropical forest Arrested succession Bwindi Competition Forest-bracken edge Invasive species Pteridium aquilinum Regeneration

A B S T R A C T

Areas dominated by bracken,Pteridium aquilinum, occur on every continent except Antarctica. These fern thickets appear to retard forest recovery and thus reduce forest diversity and carbon values. We examined how bracken inhibits forest recovery in the Bwindi Impenetrable National Park, Uganda. We established 40 50 × 5-m transects across the forest-bracken interface. On average, we recorded 596 ± 64 large trees (stems ha⁻¹± 1 SE), 1440 ± 159 saplings, 33580 ± 8860 large seedlings and 31003 ± 8854 small seedlings in the forest and 45 ± 16 large trees, 114 ± 28 saplings, 7015 ± 2268 large seedlings and 6317 ± 2240 small seedlings in bracken. All bracken clearings had been affected byfire. The density of bracken was 4.9 ± 0.3 fronds m⁻². Our results suggest that distance limitation, lack of perches, damage by vertebrates and suppression by climbers all offer potential explanations of impeded regeneration in bracken. The sparser woody vegetation that occurred in bracken typically had smaller-seeded and thicker-barked tree species than the nearby forest and also included more pioneers and fewer animal dispersed species. Interestingly we detected a negative relationship between proximity to bracken plants and woody regeneration within the forest. Several key explanatory variables in- cluding canopy cover, litter depth, distance to forest and bracken density are correlated and hard to separate.

1. Introduction

Areas dominated by brackenPteridium aquilinum(L.) Kuhn occur on every continent except Antarctica and appear to impede forest recovery (Der et al., 2009; Marrs and Watt, 2006; Marrs et al,. 1997, Page 1976).

In the East African Highlands such fern thickets tend to result from droughts,fires and other disturbance processes (landslides, abandoned cultivation etc.), and already cover hundreds of square kilometres of previously forested habitat (DS pers. obs.,Holm et al., 1997; TGC, 2011; Verdcourt, 1999). Bracken thickets are likely to expand in tro- pical Africa in future as droughts andfire are predicted to increase (IPCC, 2014).

Temperate studies indicate that common bracken,Pteridium aqui- linum(L.) Kuhn, interferes with forest regrowth (Den Ouden, 2000;

Dolling, 1996; Facelli and Picket, 1991; Gaudio et al., 2011; Priewasser, 2013). However, various interference mechanisms have been suggested and it is unclear which of these, if any, are involved in the highlands of equatorial Africa. In the Bwindi Impenetrable National Park (Bwindi), a UNESCO World Heritage site, bracken thickets are already locally abundant and appear persistent: retarding forest recovery and reducing forest biodiversity and carbon values.

There have been few ecological studies of how bracken and forest

interact in the tropics. Most publications relating to bracken in Africa focus on taxonomy or control (Masozera, 2004; Thomson et al., 2005;

TGC, 2011, but seeAdie et al., 2011). Even outside Africa, forest boundaries adjacent to bracken-dominated clearings have received little attention despite a considerable literature on forest-edge dynamics (seeHaddad et al., 2015; Harper et al., 2005; Porensky and Young, 2013for reviews). In this paper, we examine how bracken interferes with forest recovery at the forest-bracken interface in and around Bwindi in the equatorial highlands of Western Uganda. To our knowl- edge, this is thefirst ecological study of bracken in tropical Africa.

Our objectives were to: (1) describe vegetation structure across the forest-bracken boundary, (2) determine the potential factors affecting the ability of woody species to re-colonise bracken, and thus (3) provide an initial exploration of mechanisms that might explain how bracken slows forest regrowth. Our observations and review of the literature suggested many mechanisms might be operating and that bracken might be both the cause or the consequence of impeded forest recovery (Fig. 1; see alsoMarrs and Watt, 2006for a review). Crucially, these mechanisms might be operating alone or in combination so we sought to avoid misleading simplifications (seeEvans et al., 2013; Hilborn and Stearns, 1982). For this exploratory investigation, we developed simple hypotheses, and predictions, associated with a subset of the candidate

http://dx.doi.org/10.1016/j.foreco.2017.07.050

Received 20 March 2017; Received in revised form 23 July 2017; Accepted 28 July 2017

⁎Corresponding author.

E-mail addresses:[email protected],[email protected](F. Ssali).

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