3.1. Animal Performance
The month when grazing was initiated (May or July) had no apparent effect on animal performance when year 2 was included in the analysis evaluating the environmental influence of year, as years 2 and 4 were similar and years 3 and 5 were similar. The previous year’s stocking density had no influence on any animal gain variable (
p < 0.14). Otherwise, both of the main effects of year and present-year stocking density and their interaction were significant for all variables (
Table 3). For the interactions, there was an increase across years within stocking density treatments and from the low- to high-stocking densities within each year that varied in magnitude. Consequently, the discussion of animal gain variables will concentrate on the main effects.
Stocker operations expect to attain an ADG of 0.91 kg d
−1 [
1]. The low stocking density approximated that level, for both ADG28 and ADGALL (
Table 3). Despite being less than the low stocking density treatment, ADG28 and ADGALL of the medium and high densities were sufficient at >0.5 kg d
−1 [
19] to support moderate weight gain and allow for more economically effective gains at the feedlot [
3]. Summer ADG on mixed native grass pastures in a previous study at this location was 1.21 and 0.63 kg d
−1, in years 1 and 2, respectively, which was no different than introduced pastures in the same study [
3].
In his review paper, Allison (1985) [
20] reported that ADG was greatest under continuous stocking and least under rotation or strip grazing of native grasses. The present study did not have a continuous stocking treatment; however, our differences in ADG28 and ADGALL among stocking density treatments being low > medium and high is consistent with ADG declining with increasing stocking densities (
Table 3).
In previous research at this location, Capitan (2004) [
3] reported summer-long total gains by yearlings grazing mixed native grasses of 44 and 21 kg ha
−1 in years 1 and 2, respectively, which is considerably less than those measured in the present study for GAINHAALL (
Table 3). The difference is likely due to stocking density. The stocking density for Capitan et al. (2004) [
3] was 0.31 hd ha
−1, using 238 kg yearlings for an equivalent season-long (mid-May through mid-August or mid-September) stocking rate of about 8 AUD ha
−1 yr
−1, compared to the 35 AUD ha
−1 yr
−1 used in the present study. Contrary to the results of the present study regarding GAINHA and GAINALL (
Table 3), Allison (1985) [
20] and others [
2] reported that, as grazing intensity increased on blue grama pastures, gain ha
−1 increased, but individual animal gains decreased. Crawford (2019) [
4] reported work by others indicating that controlled-time grazing increased cattle weight gains using greater stocking densities and concluded from their [
4] own research that continuously grazing cattle, at lower stocking densities, spent more time walking, thus burning more energy, and thereby reducing gains.
One limitation of this study is that it reports stocking density effects at the same stocking rate season-long for a single pasture. For the medium and high stocking density treatments, multiple pastures would be needed to equal the same grazing duration as the low-density treatment (i.e., 1.7 and 2.4 times the number of pastures for the medium and high stocking densities, respectively, to equal the length of the grazing season of the low-density treatment, calculated from
Table 2). This being said, the GAINALL should also be multiplied by those factors to achieve an estimated 214 and 254 kg ha
−1, respectively, for the medium and high stocking densities for the same time frame compared to 152 kg ha
−1 measured for the low stocking density (calculated from
Table 3), which would be consistent with the findings of Crawford et al. (2019) [
4].
The stocking density used by Venter et al. (2019) [
5] was similar to that of the present study. There was no difference in animal gain from various grazing management treatments in that study [
5], including season-long grazing, but they found that too frequent movements can lead to reduced animal performance due to a reduced ability to selectively graze more palatable plants, while season-long grazing allows for repeat visits to regrowth of previously grazed plants that are more palatable and have higher quality [
4,
5,
7,
20]. Perhaps, the low stocking density treatment in the present study and stocking rate combination allowed animals in that treatment to selectively graze in a small pasture similar to patch grazing of larger areas where previously grazed plants could be revisited for a higher plane of nutrition to attain greater ADG (
Table 3). The effects of frequent moving were not evaluated in the present study; however, the influence of changing diet quality due to pasture rotations and changes in environment or pasture species has been reported at the location of this study [
21,
22,
23,
24].
3.2. Forage Mass
Soil type may have influenced the establishment of blue grama [
9] in the present study, but that was overcome by reseeding with supplemental irrigation. The cause of poor establishment in the Redona soil is not well-understood, but it may well have been related to low precipitation in late summer and autumn of year 1 (
Table 1) coupled with subtle soil-type differences in water holding capacity and plant-available water.
Results for blue grama and total forage mass were consistent because blue grama constituted >88, 98, and 95% of the total grass mass in years 3, 4, and 5, respectively (calculated from
Table 4). Consequently, the discussion for all forage mass variables will concentrate on blue grama mass after year effects are discussed, which pertain equally to all grass variables. Hence, forage mass in the present study (
Table 4) was similar to that measured previously at this location as well as elsewhere. Capitan et al. (2004) [
3] reported available forage mass of mixed native grass of 0.79, 1.19, and 1.49 Mg ha
−1 in May, July, and September, respectively, of year 1, and 1.51, 1.03, and 0.66 Mg ha
−1 in May, July, and August, respectively, of year 2. Similarly, Smart et al. (2007) [
13] reported that herbage production of shortgrasses averaged 1.3 Mg ha
−1 yr
−1, while Gillen et al. (2000) [
9] measured live standing forage mass in July and September and found it to vary from 0.63 to 1.41 Mg ha
−1 annually in mixed-grass prairie in a higher precipitation zone (766 mm annually) [
9] at the same latitude as the present study. In a review of the literature, Allison (1985) [
20] reported that livestock production in native grasses is generally low due to low intake rates driven by animal body size and low forage availability, but intake was not limited when available forage mass exceeded 0.09 Mg ha
−1. Consequently, animal performance was likely never compromised by forage availability in the present study.
The year effect (
Table 4) may have been caused by available soil moisture from the combination of precipitation (
Table 1) and irrigation (applied only in year 5), which totaled 412, 243, and 492 (203 mm irrigation + 289 mm precipitation) and influenced both blue grama and other grass mass (
Table 4). Pre-grazing precipitation for year 3 (August, year 2 through June, year 3) was similar to the long-term average for the same period (
Table 1). As mentioned, growing season precipitation likely influenced results as did annual precipitation (
Table 1 and
Table 3) [
7].
The total grass value does not equal the sum of the blue grama and other grass due to rounding and the generation of lsmeans. Means within a column and treatment effect followed by similar letters are not significantly different at p < 0.05. SEM signifies the standard error of the mean.
The year × sampling period interaction for blue grama and total mass (
Table 1 and
Table 5, for blue grama mass) likely occurred because, while there was no statistical difference in blue grama mass from the beginning of grazing until dormancy in year 3, blue grama mass did not recover after grazing in year 4; however, it did increase across the season in year 5 in similar proportions to that reported by Capitan et al. (2004) [
3] for their year 1.
The greater blue grama mass at the beginning of grazing in year 3 was likely due to significant early spring/summer grazing in year 2 (
Table 2 and
Table 5) that provided a sufficient period of regrowth prior to dormancy with precipitation in the months before dormancy in year 2 and in the spring months preceding the beginning of grazing (
Table 1). The probable cause of the season-long lesser blue grama mass in year 4 was the lateness of grazing in year 3 (
Table 2) that limited time for regrowth that year, coupled with less precipitation over winter and in the spring and early summer of year 4 prior to grazing (
Table 1 and
Table 2). Bai et al. (2022) [
7] reported that spring and summer grazing exclusion increased biomass production compared to autumn grazing. Lesser blue grama mass at the beginning of grazing in year 5 (
Table 5) was also likely caused by the lateness of grazing in year 4 (
Table 2). The productivity of perennial grasses can be greatly reduced by grazing in the previous season [
10], coupled with less precipitation throughout winter and early spring (
Table 1 and
Table 5). Changes in precipitation cause changes in the condition of rangeland [
8]. The most stressful conditions for mixed-grass prairie occur when growing season precipitation was only 69% of the long-term average [
9], which occurred after grazing in year 3 until grazing began in year 4 and after grazing in year 4 until grazing began in year 5. Regarding forage mass at the beginning of grazing (
Table 5), cattle weight gains have been found to be reduced in years with greater grass production due to excessive precipitation [
10]. That may have led to the lower ADG28 and ADGALL in year 3 and greater ADG28 and ADGALL in year 5, but the reduced values of those variables in year 4 are likely due to season-long forage mass availability (
Table 3), although Allison (1985) [
20] reported that intake was not limited when available forage mass exceeded 0.09 Mg ha
−1 and Irisarri et al. (2019) [
2] found that beef production was maximized at about 0.5 Mg ha
−1 annual net primary production.
Blue grama was intensively grazed twice (early June and early July): (1) during a single year in which drought was imposed from May through October and in the year after that, (2) during the year that drought was imposed, but not in the year after that, or (3) not grazed at all [
25]. Growth rates of blue grama grazed during and after drought were greater after the first grazing bout of the second year, but growth rates were not different among grazing treatments after the second grazing bout in early July [
25]. Monthly temperature and total long-term precipitation averages during the April-through-September growing season at the 45th to 46th latitudes, where the Bai et al. [
7] and Eneboe et al. [
25] (
https://www.usclimatedata.com/climate/miles-city/montana/united-states/usmt0229, accessed on 8 May 2023) took place, are roughly equal to April through November at the location of the present study (
Table 1) and closely reflects the blue grama growing season for the present study. Nonetheless, blue grama has been observed to initiate growth earlier when temperatures and precipitation are conducive, which may have occurred in March of years 4 and 5. Blue grama aboveground net primary production (the sum of positive monthly changes in growth from April through November) did not differ due to imposed drought or grazing treatment in either of the two years of study [
25].
Seasonal grazing, which was limited to spring or summer using the high stocking density treatment in the present study (
Table 2), was beneficial compared to season-long grazing [
7], which the low stocking density treatment most closely mimicked in the present study. Low grazing intensity had no effect on aboveground biomass, but under heavier grazing intensities, aboveground biomass was increased by exclusion in spring or summer compared to autumn [
7]. This may be related to recovery time during the period of active growth caused by warmer temperatures and greater precipitation compared to recovery during cooler temperatures and less precipitation in the latter part of the native warm-season perennial grass growing season (late summer/early autumn [
9,
12] (
Table 1,
Table 2 and
Table 6). Aboveground biomass was greater under medium than low and high grazing densities for the same season of exclusion, indicating that grassland productivity is maximized by moderate grazing intensities [
7].
While blue grama mass numerically increased after grazing ended in year 3 (
Table 5), the significant increase in blue grama mass after grazing ended in year 5 was likely due to the timing of the irrigation and significant precipitation during the post-grazing period through dormancy (
Table 1 and
Table 2). Favorable precipitation after grazing can delay the conversion of standing live crop to standing dead crop [
9], allowing native warm-season grasses to restock root energy reserves [
6] in autumn and provide greater forage in the following year prior to grazing, which may also have been the case after grazing in year 2 and prior to grazing in year 3 (
Table 5). Current-year grassland productivity can be influenced by the climatic events of the previous four years [
10] in addition to previous grazing management. In another study [
25], the effects of a single year of drought on blue grama growth rates did not carry over into the subsequent year, likely due to above-average precipitation in mid-spring and that it was only after three years of persistent drought that the regrowth potential of grasses through axillary buds was reduced, indicating significant stand loss. Recovery from frequent or severe defoliation is an indication that growing conditions, including adequate moisture, were conducive to recovery [
9]. Holochek et al. (2006) [
8] reviewed grazing management research on rangeland and reported that low-to-moderately-grazed treatments had similar effects on native vegetation and that native vegetation utilization of up to 40% was sustainable because it maintained greater basal cover than either grazing exclusion or more intensive utilization by grazing. Without accounting for growth during the grazing season, <40% utilization occurred in year 3 and growth outpaced utilization in year 5, but utilization was >50% in year 4 (
Table 5), indicating a high grazing intensity [
8] that could have influenced initial forage mass in year 5. Eneboe et al. (2002) [
25] reported that the negative influence of drought, which also was mildly the case between grazing in year 4 and year 5 (
Table 1), was not as evident in stand loss until after three years.
This study was not designed to evaluate grazing initiation at different times of the year; however, the earliness of grazing in year 2 (
Table 2) followed by precipitation near the long-term average (
Table 1) may have led to greater blue grama mass in July of year 3 when grazing began. Conversely, the lateness of grazing in years 3 and 4 (
Table 2) coupled with less precipitation between the end of grazing in those years and the beginning of grazing in the next year (243 and 289 mm from year 3 to year 4 and from year 4 to year 5, respectively) likely influenced available forage mass at the beginning of grazing in the subsequent year, especially when grazing began earlier in year 5. Holochek et al. (2006) [
11] reported that perennial grass and blue grama mass measured in autumn were similar under a season-long rest after grazing from February through April or when grazing was excluded. Grazing during the growing season that removes photosynthetically active plant material likely has a greater influence on the spring growth of shortgrasses, such as blue grama, than removal of that same plant material after the onset of dormancy, but before the initiation of spring growth. Removing animals earlier in late winter could leave greater plant residues for soil cover and plant protection over late winter and spring when winds are greater [
14]. As with animal performance, when year 2 was included in the analysis to evaluate for the environmental effects of year, years 2 and 4 had similar blue grama mass and years 3 and 5 had similar mass indicating that the month when grazing was initiated (May or July) had little influence on blue grama mass.
The trend (0.05 <
p < 0.10 [
19]) toward a year x stocking density interaction for blue grama mass (
Table 4 and
Table 6) is due to differences in the magnitude of change over years within stocking densities and nonsignificant increases in blue grama mass from low to high stocking densities in years 3 and 4, followed by a numeric decrease in blue grama mass from low to high in year 5. Although not observed in the present study due to a lack of any year × stocking density interaction for GAINHAALL (
Table 3), in light of the year × stocking density interaction for forage mass (
Table 6) showing greater blue grama mass in years 3 and 5 when greater total moisture was available for growth (
Table 1, with 203 mm supplemental irrigation applied in year 5), beef production on rangeland can be increased with grazing intensity in wet years, but not dry years [
2]. Conversely, Gillen et al. (2000) [
9] reported that stocking rate did not influence shortgrass forage production and there was no interaction with year, although, in the present study, stocking rate was held constant across treatments and years. Consequently, the trend (0.05 <
p < 0.10 [
17]) toward a year × stocking density interaction would be related to environmental influences and stocking density treatments in the present study (
Table 6). Nonetheless, shortgrasses, such as blue grama, have greater grazing resistance than other native perennial grasses [
12] because they maintain a greater proportion of photosynthetic material under grazing than taller grasses, which allows them to maintain a positive net aboveground primary productivity [
25], potentially, even during low precipitation years.
As mentioned, recovery from frequent or severe defoliation is an indication that growing conditions, including adequate moisture, were conducive to recovery [
12]. Consistent with forage mass measured 28 and 56 d after grazing indicating recovery each year (
Table 4 and
Table 5), recovery periods of 30 and 60 d were also sufficient to overcome frequent or severe defoliation of sideoats grama (
B. curtipendula) [
12], which is less grazing-tolerant than blue grama as a midgrass species [
11]. Defoliating sideoats more than once in a season significantly reduced biomass production with little difference in response when it was defoliated three (60 d intervals) or six times (30 d intervals) either moderately (10 cm stubble) or severely (5 cm stubble) when grown with a competing species [
12]. It is likely that the blue grama in the present study was defoliated by grazing more than once in each treatment each year even within the 28-day high stocking density treatment because grazing animals tend to revisit previously grazed areas due to more palatable growth [
4,
5,
7,
20]. Drought in late summer and autumn limited forage production by native warm-season grasses to 50% of the production during that period in a high precipitation year in the study by Tilhou et al. (2019) [
6], potentially also limiting the production of root energy reserves for spring growth. Under moisture-limited conditions, recovery may take longer; hence, rest periods should be longer [
12]. Consequently, similarly to sideoats [
12], it would be prudent under typical precipitation conditions to allow blue grama recovery periods of 50 or 90 d during fast and slow growth periods, respectively.
Under the analyses for the effects of the previous year’s stocking density on the present year’s blue grama mass, the effect of the previous year’s stocking density was not significant (
p < 0.19); however, the effect of the current year’s stocking density was significant (
p < 0.0338), as was the previous x current year’s stocking densities interaction (
Table 7,
p < 0.0117). As previously stated, the results were similar for total grass mass as for blue grama mass; hence, only data for the blue grama mass will be discussed. When low-density, long-duration grazing (low) followed either high-density, short-duration grazing (high), or medium stocking density, blue grama mass increased compared to low following low (
Table 7). Alternatively, when medium followed high, blue grama mass was reduced, but when high followed any previous stocking density treatment, forage mass increased, although not significantly.
The effects of grazing exclusion can be positive or negative depending on the season and duration of the exclusion period and environmental conditions; hence, grazing management practices to increase grassland productivity must be determined on a regional scale [
7], including the length of rest required for recovery after defoliation [
12]. Bai et al. (2022) [
7] found that spring grazing exclusion followed by moderate grazing intensity was the best option to preserve grasslands; however, animal feeding must continue somewhere year-round and grazing is the most cost-effective form of forage production. Consequently, based on the present study results (
Table 7), high stocking density may be the best management for a short-duration grazing session at the same time of year using multiple pastures each with the same grazing period to allow for a long-duration rest in accordance with the carrying capacity of the ranch, even if fixed seasonal grazing periods may not maximize growth potential through nutrient recycling and availability for plant growth [
10]. Additionally, because beef production on native grasses can be negligible after growth slows late in the growing season [
2] and late summer/autumn is a period during which root energy is restocked for spring growth [
10], rotating cattle to actively growing annual forages during that period may be advisable until dormancy is induced [
24]. However, grazing the dormant native grass forage as soon as possible to minimize the cost of supplementation is advisable since the animals will be lighter at that time than during late winter [
23]. Otherwise, during persistent drought, a longer-duration, medium stocking density could be used for less-intensive grazing to allow for as much recovery as possible, and avoid season-long low-density grazing, if possible (
Table 7).
The stocking rate in the present study was intermediate to that used in the 7-year study by Gillen et al. [
9], who considered their stocking rates to be high [
2]. They attributed the lack of any effect on forage mass production was due to long-term (80-year) exposure of their pastures to high stocking rates leading the vegetation to an equilibrium of production under high stocking rates. In the present study, no long-term history was in place as the pastures were in their 3
rd to 5
th year after seeding. The present study was also not likely of sufficient duration to lead to potential stand degradation due to overgrazing that can occur from the low-density, long-term grazing described by Venter et al. (2019) [
5]. Longer-term studies are necessary to evaluate the longer-term effects of grazing management [
4].