Rendimiento de la extracción supercrítica

Jan Bokdam, J. Maurits Gleichman, Sipke E. van Wieren & Michiel F. WallisDeVries

Submitted to Journal of Applied Ecology

Cattle as opportunistic exploiters. Frangula alnus (photo) and other woody and herbaceous understory species were suppressed by consumption, trampling and head-rubbing. Wolfhezerheide, July 1987 (photo Maurits Gleichman).

Free-ranging cattle as opportunistic exploiters of a successional woodland-grassland-heathland mosaic

Jan Bokdam, J. Maurits Gleichman, Sipke E. VanWieren & Michiel F. WallisDeVries

Summary

Large herbivores are increasingly used as a conservation management tool. Using herbivores in this way requires knowledge of their habitat and diet preferences. We studied the habitat and diet preferences of cattle in woodland-grassland-heathland mosaics using direct observations and bite counts from three study areas. The aim was to test differential preferences for successional habitats and species (grasses, early and mid-seral woody species) as causal factors driving cyclic succession. Our results provided evidence for differential habitat preference during foraging and non-foraging.

Foraging time was spent predominantly (83 %) in open communities, non-foraging time mainly (56 %) in woodland, despite a low proportion (33 %) of this habitat in the mosaic. The findings implicated a substantial nutrient redistribution from lawns to woodland. Deschampsia flexuosa was the dominant species in the diet. This grass species and a group of other ”short grasses” (i.a Holcus spp, Agrostis capillaris) were significantly selected. Molinia caerulea was consumed during summer only. Mid-seral shrubs and trees (e.g. Quercus robur, Sorbus aucuparia, Frangula alnus and Prunus serotina) were selected, especially during summer, but not significantly so. Rubus fruticosus was the exception and was avoided. The early-seral dwarf shrubs Erica tetralix and Calluna vulgaris (during summer only) and trees (Pinus sylvestris and Betula spp) were avoided. The opportunistic use of the mosaic components suggests that successional mosaics with at least woodland and grassland are optimal habitats for cattle. The results confirmed that cattle may act as a driving force for mosaic cycling but they also revealed variation in the palatability within successional species groups of grasses, early-seral and late-seral woody species. Cattle can suppress palatable woody undergrowth in woodland and create and deplete lawns until resistant shrubs or

trees invade. Implications of the findings for the use of cattle as a conservation tool are briefly discussed.

Key-words: nature conservation, cyclic vegetation succession, grazing, browsing, habitat preference, diet selection, soil nutrients.

INTRODUCTION

Livestock and wild herbivores are increasingly used as a management tool in nature conservation areas (Oosterveld 1975; Bakker 1989, 1998; Gordon et al. 1990;

Severson & Urness 1994; VanWieren 1995; Piek 1998). Their use as a tool presumes a key role of herbivores in the functioning of ecosystems and requires knowledge of their habitat and diet preferences (Coughenour 1991). Open, nutrient-poor habitats are important for biodiversity and in many areas traditional extensive livestock grazing is continued or reintroduced (Gimingham 1992; Bignal, McCracken & Curtis 1994;

WallisDeVries, Bakker & VanWieren 1998, Bokdam & Gleichman 2000). Livestock grazing as a management policy is challenged by the wilderness grazing policy (Bunzel-Drüke, Drüke & Vierhaus 1993/1994; Vera 1997, 2000; Svenning 2002).

Cattle, horses and other grazers may play an important role in both strategies, either as livestock or as feral species. We studied the habitat and diet preferences of free-ranging cattle as a determinant for open, nutrient-poor habitats in successional woodland-grassland- heathland mosaics.

The effects of grazing on plant succession depend on the involved herbivores (Hoffman 1989, Gordon, Illius & Milne 1996), plant species (Briske 1996; Bakker 1998; Bullock et al. 2002; Milchunas & Noy Meir 2002), and environmental factors e.g. soil fertility, fire and man (Ormerod 2002). Openness is the result of replacement of woodland by short vegetation and by prevention of re-invasion of trees.

Replacement of woodland by grassland requires tree mortality, suppression of woody undergrowth and grass invasion. In W-Europe, mature trees are rarely killed by large herbivores. They usually die by ageing, storm, fire, insects, fungi or cutting by man.

But many large herbivores are able to suppress woody undergrowth (shrubs, saplings) and to introduce grass seeds. Common undergrowth species on nutrient-poor acid soils that are suppressed are the mid-seral Quercus spp, Sorbus aucuparia, Frangula alnus, Prunus serotina and Vaccinium myrtillis and late-seral species Fagus sylvatica, Ilex aquifolium and Taxus baccata (nomenclature follows Van der Meijden 1990).

Openness of grassland or heathland habitats may be lost by invasion of early-seral trees (Pinus sylvestris, Betula spp) (Gimmingham 1992). These early-early-seral woody species differ from the mid-seral ones by smaller seeds, anemochorous (in stead of zoochorous) dispersal, smaller seedling, higher requirements for light and lower requirement for nutrients and moisture (Bazzaz 1979; Ellenberg et al. 1992).

Diet selection (Putman 1986; Van Wieren 1996) and plant chemistry (Bokdam &

WallisDeVries 1992; Van Genderen, Schoonhoven & Fuchs 1996) suggest that early-seral woody species – being more resistant than mid-early-seral ones – might be able to outcompete ungulates.

Herbivores can create or maintain nutrient-poor soils by net nutrient removals (offtake > return) and by secondary losses due to volatilisation, erosion, leaching, etc., if the losses are not compensated by inputs (fertilisation) or by accelerated cycling (Milchunas & Lauenroth 1993; Frank, Kuns & Guido 2002). Net removal is enhanced by differential habitat use (foraging and non-foraging in different habitats) redistributing nutrients from foraging habitats to resting habitats. Cattle and other social large ruminants are superior nutrient movers. Their consumption is aggregated and they spend less time foraging and more time on resting sites than non-ruminants (e.g. horses) (Arnold & Dudzinski 1978). Large herbivores with large home ranges may transport nutrients over longer distances.

Until now, models predicting grazing effects on vegetation succession have been predominantly based upon differential defoliation of established plants and competitive displacement for light (Coughenour 1993; Jorritsma et al. 1997; Vera 1997, 2000; Van Oene, Van Deursen & Berendse 1999, Groot Bruinderink et al. 1999;

Olff et al. 1999). Differential dispersal and establishment and effects mediated by changing soil fertility have either been ignored or incompletely included. Recently, long-term botanical changes in sub-alpine pastures grazed by red deer (Krüsi et al.

1995; Schütz et al. 2000) and wood encroachment in Deschampsia heaths grazed by free-ranging cattle (Bokdam 1996; Bokdam & Gleichman 2000) were attributed to lawn nutrient depletion. Differential habitat and differential diet preference (preference for mid-seral woody species, avoidance of early-seral species) have recently been integrated in a conceptual Resource-mediated Successional Grazing Cycle (RSGC) model (Bokdam, Cornelius & Krüsi 2001; Bokdam 2001). The name refers to grazing as the driving force for successional grazing cycles (as opposed to circadian, intra-seasonal and annual grazing cycles, Drent & Van der Wal 1999) and to mediation by nutrients and light. The RSGC model (Fig 1) plots biomass (and light availability) on the y-axis and soil nutrients (total or available pools) on the x-axis. The three essential grazing influences to be tested in this paper are (i) suppression of mid-seral woody undergrowth species, (ii) soil nutrient depletion of the grass lawn and (iii) avoidance of early-seral dwarf shrubs and trees invading in the nutrient-depleted grass lawn.

Domestic cattle (Bos taurus) are generally considered to be typical grass consumers and grassland exploiters. This suggests a low pressure on browse (woody food plants). The adaptation of cattle to grasses and grasslands involve a wide lower incisor arcade (Illius & Gordon 1987), an adapted digestive tract (Hofmann 1989), a high capacity to digest cell wall material (Van Wieren 1996) and an aggregative behaviour (McNaughton 1984, Phillips 1983). Although grazers (‘Grass & Roughage

BIOMASS RSGC

Fagus

TREE III BUILDING WOODLAND

I DEGENERATING

Quercus WOODLAND

Betula Pinus

SHRUB Rubus

DWARF Calluna SHRUB

FORB

ES MS

GRASS Deschampsia Holcus

II DEPLETING LAWN

SOIL NUTRIENTS

Figure 1: Resource-mediated Successional Grazing Cycle (RSGC). During woodland degeneration (I) soil fertility increases by input of nutrients in dead biomass and increased mineralisation. Grazers facilitate the establishment of grass lawns by suppressing mid-seral woody undergrowth (MS), grass seed dispersal, defoliation and treading. Nutrient and light availability increase. During the lawn stage (II) the soil nutrient pool is depleted by biomass removal. Nutrients ingested on the grazing lawns are excreted in non-foraging habitats. Nutrient depletion followed by reduced food production, food quality and grazing intensity enhance the invasion of light demanding unpalatable early-seral (ES) dwarf shrubs (Calluna vulgaris, Erica tetralix) and trees (Pinus sylvestris and Betula pendula). Abandonment by the grazer initiates a new woodland phase and nutrient replenishment (III).

Feeders’ sensu Hofmann 1989) co-evolved during the Tertiary and Quaternary with grasses (Stebbins 1981; McNaughton 1984) actual wild cattle and spiral-horned antelope species are predominantly found in woodland habitats with varying proportions of grasses (Sinclair, Cock & Hall 2001; Hall & Underwood 2001). The last free-roaming Aurochs (Bos primigenius) also lived in woodland (Jaktorow Forest, Poland, 15-16^th century) and were reported to consume grasses during summer, acorns in autumn and twigs in winter (Van Vuure 2001). It may be assumed that habitat choice and woody diets of these wild species have been affected by predators. This assumption would imply that domestication, breeding and selection since 9000 years BP (Clutton-Brock 1999) might have enforced the preference of cattle for open landscapes and grasses.

Studies on free ranging domestic cattle in woodland-grassland mosaics do not confirm this hypothesis but point to a sustained opportunistic use of woodland and grassland across foraging and non-foraging periods, seasons and environmental conditions (Arnold & Dudzinski 1978; Senft, Rittenhous & Woodmansee 1985 a,b;

Pratt et al 1986; Putman 1986; Bokdam 1987; Van Wieren 1988; Phillips 1993;

WallisDeVries 1994). The average composition (± sd) of 121 diets of free-ranging cattle, mainly from North America showed 72 (± 25) % grass (and graminoids), 15 (±

16) % forbs ( dicotyledonous herbs) and 13 (± 21) % woody plants (‘browse’) (Van Dyne et al. 1980). In some cases, the low browse content may have been due to a low availability. Ten out of the 121 diets contained > 50 % browse. It suggests that domestic cattle might be capable of creating and maintaining openness by browsing.

Cattle select not only between but also within the broad categories of grasses, forbs and browse (Van Dyne et al. 1980; Putman 1986). Based on empirical observations, pasture plants of W-Europe have been ranked in 10 classes of Forage Value ranging from –1 (poisonous and avoided), 0 (without any value, avoided) to 8 (highest quality, strongly preferred) (Klapp 1965).

Our aim was to test the following hypotheses:

1. Cattle display differential habitat preference. They prefer grassland to non-grassland (woodland and heathland) during foraging and prefer woodland to open habitats during non-foraging.

2. Cattle display diet preferences. They prefer lawn grasses to woody species, and prefer mid-seral to early-seral woody species.