Diurnal behavior of Kiko wethers in southern-pine silvopastures planted with warm-season forages

Diurnal behavior of Kiko wethers in southern-pine silvopastures planted with warm-season forages

Accepted Manuscript Title: Diurnal Behavior of Kiko Wethers in Southern-pine Silvopastures Planted with Warm-season Forages Author: Sanjok Poudel PII:...

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Accepted Manuscript Title: Diurnal Behavior of Kiko Wethers in Southern-pine Silvopastures Planted with Warm-season Forages Author: Sanjok Poudel PII: DOI: Reference:

S0921-4488(18)30227-X https://doi.org/10.1016/j.smallrumres.2019.03.005 RUMIN 5860

To appear in:

Small Ruminant Research

Received date: Revised date: Accepted date:

30 May 2018 8 January 2019 6 March 2019

Please cite this article as: Poudel S, Diurnal Behavior of Kiko Wethers in Southern-pine Silvopastures Planted with Warm-season Forages, Small Ruminant Research (2019), https://doi.org/10.1016/j.smallrumres.2019.03.005 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

Diurnal Behavior of Kiko Wethers in Southern-pine Silvopastures Planted with Warmseason Forages

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Mr Sanjok Poudel Virginia Tech [email protected]

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Grazing was the predominant diurnal behavior shown by Kiko wethers in southern-pine silvopasture during the warm-season grazing period. Kiko wethers did not show any pine debarking behavior throughout the observation period. Grazing was the predominant diurnal behavior shown by Kiko wethers during post-midday period while lying was predominant diurnal behavior during midday period. Kiko wethers were found safe to integrate in southern-pine silvopasture consisting of 12-yrsold trees during the warm-season grazing period.

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Highlights

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1. Introduction

Silvopasture is a common form of agroforestry system that involves deliberate integration

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and management of timber, livestock, and forages under a common management unit (Clason and Sharrow, 2000). It can be established in existing pastures by planting single or double rows of trees with wide alleys between tree rows or sets of tree rows for continuously growing forages in alleys. It can also be established in existing stands of trees by thinning out the number of trees to reduce the tree density to a desirable level for supporting forage production (Walter, 2015).

The Southeastern United States presents a considerable potential for developing the silvopasture system due to its mild and moist climatic condition suitable for the production of all three components of this system – timber, forages, and grazing livestock (Rietveld and Francis, 2000).

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Another condition that provides the avenues for silvopasture development in this region is its large portion of land under forest cover.

Forest-use land comprises 61.9% of the total land area in the Southeast (USDA/ERS, 2017), and the major portion is owned by private landowners (Nickerson et al., 2011). This situation provides a unique opportunity to landowners for converting the existing forests into

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silvopastures by thinning out the selected trees that are defective and not good for quality timber

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production and cultivating forages in the available ground space. Once forages are established

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well in the system, suitable grazing animals need to be used to utilize these forages in order to

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generate regular short-term economic returns while waiting for the long-term returns from timber

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sale, for which landowners may have to wait for a couple of decades. As trees are the major

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components of the silvopasture system, it is crucial to identify animal species that utilize the available vegetation in the system well and do not hamper trees. Goats are one of the potential

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candidates for integration in silvopastures, as they have a distinctive ability to utilize a wide range of vegetation available in the grazing land that cannot be utilized by other grazing animals

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(Luginbuhl et al., 2000).

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Papachristou et al. (2005) highlighted goats’ diet consisting of the mixture of herbaceous

(33%) and woody species (67%). Similar findings are reported by other authors with goats’ eating woody species (69%), grasses (19%), and forbs available in the grazing system (Yayneshet et al., 2008). Karki et al. (2018), from studies conducted in southern-pine silvopastures, mentioned that Kiko wethers consumed well all the planted forages (eight different

species of cool-season grass, legumes, and forbs) and volunteer vegetation spontaneously grown in the study plots. These authors found grazing as the most dominant behavior (36-54%) of Kiko wethers, rotationally stocked in silvopastures, followed by lying (29-34%) and loafing (15-22%)

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during the cool-season grazing period. The time allocated to grazing might vary depending on diurnal and seasonal timescale, photoperiod, and temperature (Dumont et al., 2004; Rutter, 2010; Pokorna et al., 2013). When animals are integrated in the tree production system, such as silvopastures, their behavior towards trees can be a matter of concern. Karki et al. (2018) reported the debarking behavior of Kiko wethers while rotationally stocked in silvopasture plots

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containing 10-11-year-old longleaf (Pinus palustris Mill.) and loblolly (Pinus taeda L.) pine

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trees despite plenty of high-quality forages available for grazing in the study plots during the

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cool-season grazing period. However, how these animals would interact with trees present in the

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southern-pine silvopastures during the warm-season grazing period has not been known.

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Behavior and the distribution pattern of livestock are key factors to be taken into

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consideration for the successful management of any grazing system. Basic understanding of the diurnal behavior of goats in southern-pine silvopastures helps producers and land managers in

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managing these systems integrated with goats well, especially avoiding the untoward effect on trees - the main component of the silvopasture system. Understanding the behavior of grazing

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livestock also helps producers in efficiently designing an improved grazing scheme that enhance animal productivity and welfare, along with improving the forage productivity (Lin et al., 2011;

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Meier et al., 2012). Moreover, the distribution pattern of grazing livestock is a major concern for producers as uneven grazing distribution may increase grazing pressure on areas that are overused, and wastage of resources available in certain areas that are underused. Furthermore, understanding the behavior pattern of grazing livestock helps producers and land managers in

improving the distribution of animals in the grazing system. The objective of the study was to determine the diurnal behavior and distribution pattern of Kiko wethers in southern-pine silvopastures during the warm-season grazing period.

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2. Materials and Methods All procedures were approved by the Tuskegee University Institutional Animal Care and Use Committee’s protocol. 2.1 Study Site

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The study was conducted during the summer 2017 at the Atkins Agroforestry Research

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and Demonstration site, Tuskegee University, Tuskegee, Alabama, United States (32° 26'

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38.2416" N; 85° 44' 2.5152" W). The study site comprised of Uchee loamy sand and Cowarts

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loamy sand with a slope of 1-15 percent and located at 141 m above the sea level. Its average annual rainfall is 1320 mm. The study site consisted of three fenced silvopasture plots (0.4-ha

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each) with 12-year-old longleaf and loblolly pines at a tree density of 402 trees ha-1 with loblolly

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to longleaf ratio of 1.27 at the beginning of the study. The average height, diameter at the breast

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height (DBH), and basal area of loblolly pine were greater than longleaf pine (p<0.0001) (Table 1). Study plots also consisted of some spontaneous browse plants, such as American beautyberry

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(Callicarpa americana L.), yaupon (Ilex vomitoria Aiton), peppervine (Ampelopsis arborea L. Koehne), honeysuckle (Lonicera L.), greenbrier (Smilax L.), persimmon (Diospyros virginiana

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L.), and winged elm (Ulmus alata Michx.) along the fence line and close to tree bases. 2.2 Forage Establishment Each silvopasture plot was demarcated into three equal subplots and three different warm-season forages were randomly allocated in these subplots (zones) within each plot (Figure

1), and seeded with no-till drill. Forages planted in different forage zones within each study plot included bahiagrass (Paspalum notatum Flueggé), crabgrass (Digitaria sanguinalis (L.) Scop.), and sericea lespedeza (Lespedeza cuneata (Dum. Cours.) G. Don). Soil samples were collected

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from each study plot and analyzed for soil pH and nutrient content three months prior to forage plantation. Lime and fertilizers (urea, muriate of potash, and triple super phosphate) were applied to each study plot based on the soil-test recommendation. 2.3 Forage Biomass and Quality

Forage samples were collected by clipping ten random quadrats (0.25-m2 each) to four-

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inch stubble height from each forage zone within each study plot prior to stocking animals.

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Samples were dried at 60 °C for 72 h and weighed to determine the available forage dry matter.

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Dried samples were then ground to 2-mm mesh and analyzed for crude protein (CP), acid

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detergent fiber (ADF), and neutral detergent fiber (NDF). Dry combustion method (Hue and

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Evans, 1986) was used to determine the total nitrogen, which was used to calculate CP content

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(CP = %N x 6.25). Proximate analysis method (Van Soest et al., 1991) was used to determine ADF and NDF contents, and total digestible nutrients (TDN) was calculated from NDF content

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using the equation - TDN = {(105.2-0.667*NDF) *0.88} (Soil, Forage, & Water Testing

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Laboratory at Auburn University, personal communication).

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2.4 Diurnal Behavior and Distribution Pattern of Kiko Wethers Ten Kiko wethers (2.75-2.90-year-old, 62.9 ± 1.56 kg BW) were rotationally stocked in

three silvopasture plots. They had free access to portable shelters, clean drinking water, and mineral-mix (ad lib.) throughout the study. Wethers were allowed to adjust in each study plot for a day prior to monitoring their behavior. The diurnal (dawn-dusk) behavior and distribution pattern of wethers were monitored for one day when they were stocked in each study plot.

Behavior and distribution pattern of animals were monitored and recorded once every 10minutes in pre-formatted data sheets. Behavior category recorded during the study included grazing, browsing, loafing, lying, debarking, and staying in shelters. Animals’ feeding on

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herbaceous species with head down was considered grazing; animals’ feeding on tender shoots, twigs, and leaves of trees and shrubs was considered browsing; animals’ walking, standing, or playing were considered loafing; animals’ resting on the ground was considered lying; animals’ peeling off and chewing the bark of standing trees was considered debarking. Observations were taken from the observation stations, installed at strategic locations at the height of 4.5 m from the

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ground level; binoculars were used when necessary. Goats were moved from one plot to the next

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once 50% of the available forages were eaten, which was determined by measuring the forage

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height with grazing stick before grazing began and during grazing coupled with daily visual

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observation of the standing forages in the study plots. Behavior study began on June 20 and

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ended on July 12, 2017.

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2.5 Weather Data

Secondary weather data (temperature, air pressure, humidity, visibility, dew point, and

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wind speed) were retrieved from www.friendlyforecast.com recorded for Tuskegee Institute for

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each observation day.

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2.6 Data Analysis Observation dates were divided into three diurnal periods as morning (dawn-1100h),

midday (1100h-1500h), and post-midday periods (1500h-dusk) (Karki and Goodman 2010) to determine the variation in behavior and distribution patterns of wethers at different diurnal periods. Distribution Evenness Index (DEI) was quantified using the formula given by Zuo and Miller-Goodman (2004).

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DEI= (− ∑𝑖=𝑙 𝑝𝑖 ln𝑝𝑖)/In𝑍 Where pi = the proportion of wethers present in a specific zone at the observation time, Z = number of zones included in the study.

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The behavior and distribution data were analyzed using Kruskal-Wallis rank-sum test (Kruskal and Wallis, 1952) in R-package 3.3.2 that determines whether the mean score (mean rank for different groups) differs for different groups (each behavior category among different diurnal periods). Weather data were analyzed using generalized linear model (GLM) option in

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SAS 9.4. Forage biomass data were analyzed using Mixed model in SAS 9.4 with plots as the

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random factor. Forage quality data (CP, NDF, ADF, and TDN) were analyzed using Multivariate

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analysis of variance (MANOVA) option as the quality parameters were correlated. Probability

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level of alpha was set at 0.05 for rejection of the null hypothesis. 3. Results

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3.1 Forage Biomass and Quality

Forage biomass was the highest (p<0.0001) for crabgrass (1798 ± 83.1 kg ha-1) planted in

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Zone 2 followed by bahiagrass planted in Zone 1 and sericea lespedeza planted in Zone 3 (Table 2). Crude protein (CP) content was the lowest (p<0.05) in bahiagrass (7.8 ± 0.62%) as compared

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to crabgrass and sericea lespedeza. The neutral detergent fiber (NDF) was the lowest (53.2 ±

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1.55%) and the total digestible nutrient (TDN) was the highest in sericea lespedeza (61.3 ± 0.82%) planted in Zone 3 as compared to forages planted in the other two zones (p<0.05) (Table 2). 3.2 Weather

Air temperature was the lowest and relative humidity was highest in the morning (dawn1100h) (p<0.0001) among the diurnal periods. Wind speed was higher during midday (1100h1500h) versus the morning hours (dawn-1100h) (p<0.05) (Table 3).

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3.3 Animal Behavior and Distribution Pattern The average diurnal observation period lasted for 15 hours for each observation date. Kiko wethers spent 46% of diurnal time grazing, 22% time staying in shelter, and 17% time lying in the tree shades (Figure 2). Grazing was the predominant diurnal behavior of wethers during the post-midday period (57%) followed by morning (47%) and midday (29%) periods,

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while lying (24%) and staying in shelters (40%) were predominant diurnal behavior during the

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midday period (1100h-1500h) (Table 4). Wethers did not debark any pine trees throughout the

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observation period. The overall distribution evenness index (DEI) of wethers was 0.06 with no

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significant difference for any diurnal period. Overall, wethers spent the most time in Zone 3

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(42%) planted with sericea lespedeza followed by Zone 1 (31%) planted with bahiagrass and

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Zone 2 (27%) planted with crabgrass. Goats were concentrated the most in Zone 1 during the

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(Figure 3).

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post-midday period (1500h-dusk) and in Zone 3 during the midday period (1100h-1500h)

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4. Discussion

Grazing was the predominant diurnal behavior (46%) shown by wethers throughout the

study. Similar finding was reported by Karki et al. (2018) where grazing was the predominant diurnal behavior (36-54%) shown by goats in southern-pine silvopasture during the cool-season grazing period. Though goats are considered as good browsers (Wilson et al., 1975), with limited

availability of shrubs and trees within the reach of goats for browsing in the current study, they grazed well in the silvopasture plots with predominant coverage of planted forages to meet their dietary needs. Goats tend to have a special ability to change their foraging behavior according to

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the available vegetation (du Plessis et al., 2004). Grazing was the predominant diurnal behavior shown by goats during the post-midday period (1500h-dusk; 57%) compared to morning (dawn1100h; 47%) and midday (1100h-1500h; 29%), thereby showing the effect of diurnal periods on goats’ behavior (p<0.05). This finding from the current study is in agreement with Solanki (2000), who reported that goats, when allowed to graze in semi-arid grassland during summer,

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monsoon, and winter, showed two peaks for grazing: morning and evening, with evening grazing

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being more pronounced than in the morning.

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Diurnally active mammals and birds exhibit foraging activity higher in the evening hours,

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considering the long period of food deprivation during the night (Marler and Hamilton, 1968).

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Higher forage concentration of nonstructural carbohydrates during the evening hours (Sauve et

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al., 2009) may also influence goats to graze more during the evening hours. Burns et al. (2005) has also reported higher digestibility of the forages cut in the late afternoon compared to early

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morning in goats. Also, wet vegetation in grazing lands due to dews during the morning hours may have caused the decrease in grazing time of goats in the morning to minimize the possible

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intake of gastrointestinal parasites, which swim up onto the wet forage blade and get into animals as these parasites are consumed along with the wet forages (Miller 2004; Karki et al. 2018).

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Previous studies with goats have shown decreased grazing time with wet vegetation (Brindley et al.1989; Mellado et al., 2004). Goats’ spending most of the post-midday period grazing and no physical activities during the night may lead to low appetite in the morning, causing them to spend less time grazing.

Wethers spent significant diurnal time lying in tree shade and staying in shelters, especially during the midday with scorching sunlight and resulting high temperature (28.7 ± 0.87°C). In contrast, there was no significant difference in time spent lying by goats during the

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morning, midday, and post-midday periods in the southern-pine silvopastures during the coolseason grazing period (Karki et al., 2018). This contrasting behavior of goats may be because of higher temperature (28.7 ± 0.87°C) during the midday period in the current study versus the previous study. Goats may have preferred lying to performing other activities during the midday period in order to avoid the heat stress (Johnson, 1991). The shade- and shelter-seeking behavior

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of wethers also demonstrates the importance of trees and shelters in the grazing system that may

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reduce heat stress and eventually improve the performance of animals. Wethers also showed

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some browsing behavior by consuming pine needles intact on trees. Khatri et al. (2016) stated

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that goats can browse vegetation of their interest using a bipedal stance to reach up to the average height of 1.5 m. Wethers did not show any debarking behavior in the current study.

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Previous study conducted during the cool-season grazing period at the same research site

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involving the same animals as used in the current study showed some debarking behavior (2%)

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on pine trees, especially longleaf pine (Karki et al., 2018). This contrasting behavior of goats in different grazing seasons under the similar landscape may have been influenced by the change in

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physiological characteristics of pine barks during different seasons of the year. Further study is required to determine the effect of seasonal variation on the physical structure and chemical

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composition of pine barks. Wethers showed a very little value for overall distribution evenness index (DEI) (0.06)

with no significant difference among the diurnal periods. Karki et al. (2018) reported higher value (spring- 0.29 ± 0.032; winter- 0.15 ± 0.035) of DEI for Kiko wethers grazing in southern-

pine silvopastures compared to the current study. This variation in DEI of goats between these two studies could be because of the larger (0.1-0.2 ha) forage zones and fewer animals used in the current study, and difference in study season between these two studies. Despite low DEI,

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wethers went to all forage zones together and consumed the available vegetation there. Wethers were concentrated more in Zone 3 (42%) containing sericea lespedeza compared to other forage zones, indicating their preference for higher-quality forages. Sericea lespedeza had higher level of CP and total digestible nutrient compared to forages present in other zones. Goats have a unique ability to select diets higher in digestible organic matter and CP than the average of all

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available forages in the grazing land (Papachristou, 1997; Hadjigeorgiou et al., 2003).

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5. Conclusions

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Grazing was the predominant diurnal behavior (46%) shown by Kiko wethers throughout

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the study followed by staying in shelters (22%) and lying (17%). Behavior of wethers was

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influenced by the diurnal period (p<0.05), as they spent more time grazing (57%) during the

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post-midday period (1500h-dusk) and lying (24%) and staying in shelters (40%) during the midday period (1100h-1500h). Wethers did not show any damaging behavior to pine trees in the

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study plots throughout the observation, showing that these animals were safe to integrate in

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southern-pine silvopastures during the warm-season grazing period. Wethers utilized shelters and tree shades for lying (resting) during the midday period, which signifies the importance of trees

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and artificial shelters incorporated into the grazing system for animals’ comfort. Although wethers showed a very little value for the overall distribution evenness index (0.06), they went to all forage zones together and consumed the available forages well. Findings from the current study revealed that Kiko wethers are safe to integrate in southern-pine silvopastures consisting of 12-year-old longleaf and loblolly pines during the warm-season grazing period.

Acknowledgments The major funding for this study was provided by USDA NIFA AFRI Competitive Grant Number 2016-68006-24764, and a partial support was received from McIntire Stennis Forestry We would like to extend our deep gratitude to all staff at Tuskegee

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Research Program.

University George Washington Carver Agricultural Experiment Station and Cooperative Extension for their committed help and necessary support during the field and lab work

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performed during the study.

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developments

and

season.

J.

Range

Manag.

A

CC

EP

TE

D

M

A

N

U

https://doi.org/10.2307/4003970.

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Zuo, H.T., Miller-Goodman, M.S., 2004. Landscape use by cattle affected by pasture 57,

426–434.

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A

N

Fig. 1. Study plots (separated with yellow solid lines) with different forage zones (subplots; separated with the dotted line within each plot), summer 2017, Atkins Agroforestry Research and Demonstration Site, Tuskegee University, Tuskegee, Alabama, USA

A

CC

EP

TE

D

M

Zone 1 consisted of bahiagrass (Paspalum notatum Flueggé); Zone 2 consisted of crabgrass (Digitaria sanguinalis (L.) Scop.); Zone 3 consisted of sericea lespedeza (Lespedeza cuneata (Dum. Cours.) G. Don)

40 30 20 10 0 Grazing

Browsing

Loafing

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Diurnal Time Spent (%)

50

Lying

N

U

Behavior Category

Staying in Shelter

A

CC

EP

TE

D

M

A

Fig. 2. Diurnal time spent (dawn-dusk; mean ± SE) by Kiko wethers performing different behaviors in southern-pine silvopastures, summer 2017, Atkins Agroforestry Research and Demonstration Site, Tuskegee University, Tuskegee, Alabama, USA

Morning (Dawn-1100h) Midday (1100h-1500h) Post-midday (1500h-Dusk) a

60

a*

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Diurnal Time Spent (%)

80

b

b

c

40

20

c

0

Zone 2

N

Forage Zones

Zone 3

U

Zone 1

A

CC

EP

TE

D

M

A

Fig. 3. Diurnal time spent (dawn-dusk; mean ± SE) by Kiko wethers in different forage zones at different diurnal periods in southern-pine silvopastures, summer 2017, Atkins Agroforestry Research and Demonstration Site, Tuskegee University, Tuskegee, Alabama, USA (* P<0.05) Zone 1 consisted of bahiagrass (Paspalum notatum Flueggé); Zone 2 consisted of crabgrass (Digitaria sanguinalis (L.) Scop.); Zone 3 consisted of sericea lespedeza (Lespedeza cuneata (Dum. Cours.) G. Don)

Table 1 Parameters of pine trees present in the study plots, summer 2017, Atkins Agroforestry Research and Demonstration Site, Tuskegee University, Tuskegee, Alabama, USA

Loblolly (Pinus taeda L.),

Diameter at Height (cm)

16.1 ± 0.18a***

the

Breast Basal Area (m2)

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Tree Species

Numb Height (m) er of Trees LSMean ± SE 270 7.9 ± 0.07a***

0.023 ± 0.0020a ***

A

CC

EP

TE

D

M

A

N

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Longleaf (Pinus palustris 212 7.3 ± 0.08b 11.9 ± 0.21b Mill.). ab LSMeans with different superscripts in a column differ (***p<0.0001)

0.014 ± 0.0021b

Table 2 Dry matter and quality of forages from study plots, summer 2017, Atkins Agroforestry Research and Demonstration Site, Tuskegee University, Tuskegee, Alabama, USA. Dry Matter CP† ADF -1 (kg ha ) % (LSMeans ± SE) b 1 1333 ± 83.1 7.8 ± 0.62b 34.3 ± 0.64 Bahiagrass (Paspalum notatum Flueggé) 2 33.9 ± 0.64 Crabgrass (Digitaria sanguinalis (L.) Scop.) 1798 ± 83.1a* 10.3 ± 0.62a c a* 3 11.2 ± 0.62 34.1 ± 0.64 Sericea lespedeza (Lespedeza cuneata 857 ± 83.1 (Dum. Cours.) G. Don) abc LSMeans with different superscripts in a column differ (*P<0.05) † CP- Crude protein; ADF- Acid detergent fiber; NDF- Neutral detergent fiber; TDN- Total digestible nutrients Table 3 Weather variables for three different diurnal periods for the observation dates, summer 2017, Atkins Agroforestry Research and Demonstration Site, Tuskegee University, Tuskegee, Alabama, USA

N

U

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Zones Forage Species

A

CC

EP

TE

D

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A

Temperature (°C) Humidity (%) Wind Speed (MPH) Diurnal Period LSMeans ± SE Morning (Dawn-1100h) 24.5 ± 0.55b 79.3 ± 2.52a*** 5.8 ± 0.63b Midday (1100h-1500h) 28.7 ± 0.87a 58.7 ± 1.59b 8.2 ± 0.30a* Post-midday (1500h-Dusk) 30.1 ± 0.40a*** 55.8 ± 1.44b 7.2 ± 0.67ab abc LSMeans with different superscripts in a column differ (*P<0.05, ***P<0.0001) Source: Adapted from friendlyforecast.com - data recorded at Auburn-Opelika airport for Tuskegee Institute, AL † msl- mean sea level; MPH- Miles Per Hour

NDF

66.4 ± 1.38 63.9 ± 1.26 53.2 ± 1.55

Table 4 Distribution Evenness Index (DEI) and diurnal time spent by Kiko wethers on different behaviors and their corresponding mean scores (Kruskal-Wallis rank-sum test), summer 2017, Atkins Agroforestry Research and Demonstration Site, Tuskegee University, Tuskegee, Alabama, USA

138b 105c 164a*

139 143 135

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Morning (Dawn-1100h) Midday (1100h-1500h) Post-midday (1500h-Dusk) Mean Score Morning (Dawn-1100h) Midday (1100h-1500h) Post-midday (1500h-Dusk)

% (Means ± SE) 0.06 ± 0.017 47 ± 3.3 0.07 ± 0.022 29 ± 4.6 0.05 ± 0.016 57 ± 4.3

Loafing

Lying

Debarking

S s

1 ± 0.5 0 0

24 ± 1.9 7 ± 1.2 11 ± 1.7

13 ± 2.7 24 ± 4.1 16 ± 3.4

0 0 0

1 4 1

146a* 135b 135b

179a* 103b 119b

132b 156a* 132b

138 138 140

1 1 1

U

DEI

N

Diurnal Period

Behavior Category Grazing Browsing

abc

A

CC

EP

TE

D

M

A

Mean scores with different superscripts in a column within each behavior category differ (*P<0.05)

24