Grazing Exclosures Reveal Divergent Patterns of Change in Bunchgrass Grasslands of Western Canada
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Botany Grazing Exclosures Reveal Divergent Patterns of Change in Bunchgrass Grasslands of Western Canada Journal: Botany Manuscript ID cjb-2020-0083.R1 Manuscript Type: Article Date Submitted by the 22-Jul-2020 Author: Complete List of Authors: Bradfield, Gary; University of British Columbia, Cumming, Preston; University of British Columbia Newman, Reg; British Columbia Ministry of Forests Lands and Natural Resource Operations Krzic, Maja;Draft University of British Columbia, Faculty of Land and Food Systems and Faculty of Forestry grassland plant communities, temporal change vegetation, rough fescue, Keyword: Kentucky bluegrass, non-metric multidimensional scaling Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : https://mc06.manuscriptcentral.com/botany-pubs Page 1 of 40 Botany Grazing Exclosures Reveal Divergent Patterns of Change in Bunchgrass Grasslands of Western Canada Gary E. Bradfield, W.F. Preston Cumming, Reg F. Newman, and M. Krzic Gary E. Bradfield. Department of Botany, The University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada. W.F. Preston Cumming. Department of Botany, The University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada. Current Address: Center for Teaching & Learning, The University of Colorado,Draft Boulder, CO, 80309, USA. Reg F. Newman. British Columbia Ministry of Forests, Lands, and Natural Resource Operations, Kamloops, British Columbia, V2C 2T3, Canada. Maja Krzic. Faculty of Land and Food Systems and Faculty of Forestry, The University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada. Corresponding author: Gary Bradfield (email: [email protected]) https://mc06.manuscriptcentral.com/botany-pubs Botany Page 2 of 40 Abstract Thirty-six long-term (14-83 years) cattle grazing exclosures and adjacent grazed pastures spanning a climatic gradient from cooler-wetter to warmer-drier growing seasons in south- central British Columbia were compared for temporal vegetation change. Trajectories of temporal vegetation change from non-metric multidimensional scaling were mostly scattered for grazed areas, but more directed toward the dominant grasses, primarily rough fescue (Festuca campestris) or Kentucky bluegrass (Poa pratensis), for exclosures. Plant community differences, detected only after 10 years of grazing exclusion, were primarily due to structural shifts in overall species cover related to growth increases of the dominant grasses inside exclosures. Species richness remained unchangedDraft between the first and last sampling dates in both grazed areas and exclosures, with both treatments showing moderate degrees (15-30%) of turnover in species composition. Shannon diversity declined in both treatments as a result of the structural changes in species cover. The results highlight the value of repeated monitoring of long-term exclosures for assessment of grassland resiliency to grazing. Further studies of the interaction of grazing and climate are needed for a more complete understanding of the ongoing vegetation change. Key words: Grassland plant communities, temporal change vegetation, rough fescue, bluebunch wheatgrass, Kentucky bluegrass, non-metric multidimensional scaling. https://mc06.manuscriptcentral.com/botany-pubs Page 3 of 40 Botany Introduction Temporal change in plant species richness and community composition has long been a central focus in ecological research. A SpringerLink search for the key words “temporal change vegetation” identified 46,334 articles published in the period 1990–2020 (search date 17 April 2020). Vegetation composition and change over time play key roles in setting priorities for grassland management, especially in balancing the pasture requirements of domestic grazing animals with the conservation goals of sustaining native plant biodiversity (Brown et al. 2008; Sanderson et al. 2015). Despite its central importance, knowledge of vegetation change remains speculative for most terrestrial ecosystems owing to the lack of repeated long-term monitoring. As a result, management decisions are oftenDraft hampered by an incomplete understanding of the vegetation changes actually taking place. The grasslands of south-central British Columbia (BC) are recognized for the forage and ecosystem services they provide to the BC cattle ranching industry as well as their high biodiversity values (Delesalle et al. 2009; Lee et al. 2014). Following the introduction of domestic grazing animals (primarily horses and cattle) in the mid-to-late 1800s, large areas of grassland experienced major shifts in ecological conditions (Tisdale 1947; McLean and Tisdale 1972; McLean 1982; Wikeem and Ross 2002). For example, increases in year-round grazing pressures led to severe declines in the dominant native bunchgrasses, particularly, bluebunch wheatgrass (Pseudoroegneria spicata (Pursh) A. Löve), Idaho fescue (Festuca idahoensis Elmer), and rough fescue (Festuca campestris Rydb.), and increases in other, more ‘grazing tolerant’ plant species, both native and introduced (Tisdale 1947; Gayton 2003). In particular, the naturalized invasive, Kentucky bluegrass (Poa pratensis L.), has become a cause of concern https://mc06.manuscriptcentral.com/botany-pubs Botany Page 4 of 40 in the mesic (wetter) grasslands of BC owing to its high grazing tolerance and ability to suppress native grasses (Gayton 2004). By the 1930s, the serious state of overgrazing in many areas led to initiatives by a small group of rangeland ecologists to establish a number of grazing exclosures (fenced areas of <0.01 to 1 ha) that would provide ‘rest’ from overgrazing and demonstrate the potential for degraded grasslands to recover back to their natural state (Tisdale 1947). While the exclosures were designed primarily to prevent cattle entry, they did not block access from other native grassland fauna (e.g., birds, mice, voles, deer, elk). More exclosures were added over the years, particularly from the 1960s to 1990s, with a present-day total of over 360 exclosures scattered throughout the BC grasslands (Province of BC 2019). Owing to the challenges of follow-upDraft monitoring, a consistent record of vegetation change for many of the exclosures and adjacent grazed areas was not maintained. Fortunately, repeated monitoring records are available for a sub-set of exclosures representative of the geographic and climatic variability of the south-central BC grasslands. Some of these exclosures served as valuable case studies for early seminal work on grassland classification and management in the province (van Ryswyk et al. 1966; McLean and Marchand 1968; McLean and Tisdale 1972). Subsequent studies have reported on detailed changes in species cover at some of the exclosures (Gayton 2003; Newman et al. 2011; Wikeem et al. 2012; Krzic et al. 2014; Cumming et al. 2016). Results have indicated the potential for recovery of native bunchgrasses following either grazing removal (i.e., inside exclosures), or after a change from spring to fall grazing (outside exclosures), with recovery rates largely dependent on the seral stage of the plant community at the time of exclosure establishment or management change. In some cases, improvements in soil conditions inside exclosures over time also have been reported (Krzic et al. 2014; Richardson et al. 2017). https://mc06.manuscriptcentral.com/botany-pubs Page 5 of 40 Botany Here we report on a comparative analysis of the subset of repeatedly monitored exclosures and adjacent grazed areas to examine the broader patterns of plant community change across the south-central BC grasslands. Given the wide range of spatiotemporal variability inherent in the dataset, we focused our assessment on the following four, general questions relating to overall patterns of change: (1) To what extent have species composition and diversity changed over the course of monitoring? (2) How have the vegetation changes differed between grazed areas and exclosures? (3) Is there evidence for recovery of native bunchgrass communities? and (4) What recommendations can be drawn from this study for rangeland management? Draft Methods Assembly of the long-term dataset Vegetation data from 36 grassland exclosures in three Natural Resource Districts as defined by the BC Ministry of Forests, Lands and Natural Resource Operations (2017) in south- central BC were used in this study (Table 1; Fig.1). Exclosures ranged in age from 14 to 83 years at the time of the most recent sampling (1996-2015). Most exclosures had been sampled 2- 3 times since establishment, with two exclosures (Prudens and Goose Lake) having been sampled 7 and 8 times, respectively. Intervals between the first and last monitoring periods ranged from 5 years (Minnie Lake East) to 55 years (Goose Lake). In all cases, a nearby grazed area had been selected as a ‘matched pair’ for comparative monitoring of plant community change. The majority of exclosures were fenced areas (ca 1.5 m high fencing) whose size and https://mc06.manuscriptcentral.com/botany-pubs Botany Page 6 of 40 location were determined by ‘expert judgement’ of the local rangeland ecologist (size range 0.03 to >1 ha; three exclosures (Clapperton and two sites at Long Lake South) were actually large, ungrazed pastures. At each site, vegetation sampling occurred along permanent transects (25-50 m length) using fifty, 20 cm × 50 cm quadrats to record canopy cover of individual plant species (Daubenmire 1959); the same quadrat locations (random or contiguous depending on total transect length)