Next Article in Journal
Agricultural Holdings and Slaughterhouses’ Impact on Patterns of Pathological Findings Observed during Post-Mortem Meat Inspection
Previous Article in Journal
Pre- and Post-Slaughter Methodologies to Estimate Body Fat Reserves in Lactating Saanen Goats
 
 
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

Temporal Variations in Chemical Composition, In Vitro Digestibility, and Metabolizable Energy of Plant Species Browsed by Goats in Southern Mediterranean Forest Rangeland

by
Youssef Chebli
1,2,*,
Samira El Otmani
1,2,
Mouad Chentouf
2,
Jean-Luc Hornick
1 and
Jean-François Cabaraux
1
1
Department of Veterinary Management of Animal Resources, University of Liège, Avenue de Cureghem 6, B43, 4000 Liège, Belgium
2
National Institute of Agricultural Research (INRA), 78 Bd. Mohamed Ben Abdellah, Tangier 90010, Morocco
*
Author to whom correspondence should be addressed.
Animals 2021, 11(5), 1441; https://doi.org/10.3390/ani11051441
Submission received: 23 April 2021 / Revised: 13 May 2021 / Accepted: 15 May 2021 / Published: 18 May 2021
(This article belongs to the Section Animal Nutrition)

Abstract

:

Simple Summary

Mediterranean forest rangelands constitute essential feed resources for grazing goats. The objective of this study was to evaluate the temporal variations in chemical composition, in vitro digestibility, and metabolizable energy of browsed plant species by goats on forest rangelands of the Southern Mediterranean of northern Morocco. Overall, the nutritive value of the selected plant species was highest in spring and then steadily decreased through the summer and autumn. Most of the selected plant species present high levels of crude protein than the minimum required level for maintenance. This study provides a valuable and useful database to elaborate the seasonal grazing and feeding management plan for goat herds.

Abstract

Forest rangelands contribute largely to goat diets in the Mediterranean area. Information about browsed plant quality is essential for adequate feeding management. The purpose of this study was to evaluate the temporal changes in chemical composition and in vitro digestibility of the main plant species selected by goats in the Southern Mediterranean forest rangeland during two consecutive years; these were very contrasted (dry and wet). The browsed species were composed of herbaceous, eleven shrubs, and four tree species. Overall, large variability in chemical composition, in vitro organic matter digestibility (IVOMD), and metabolizable energy (ME) was observed among species, grazing season (spring, summer, and autumn), and years within each species. Crude protein (CP) content varied from 60 to 240 g/kg dry matter (DM). The fiber fractions, except for Quercus suber, increased significantly by advancing maturity. Due to the water stress, the lignin level presented a higher value during the spring of the dry year. Condensed tannin (CT) content varied from 2 to 184 g/kg DM. CP, IVOMD, and ME showed a negative correlation with lignin and CT. Based on the results presented herein, it is concluded that the nutritive value of the browsed plant species was highest in the spring and lowest during the summer and autumn of both studied years. With a good grazing management strategy, the selected plant species by goats could guarantee high-quality feeding resources throughout the year.

1. Introduction

Mediterranean forests are composite landscapes of shrubs and trees, which constitute essential dietary resources for domestic ruminants. They also play a very important role in sustaining biodiversity [1] and provide multiple ecosystem services to local people for millennia [2]. These woodlands are characterized by heterogeneous and diversified flora [3].
Livestock, especially extensive goat farming, is one of the most important components of agricultural systems in the Southern Mediterranean Basin. Goat farming systems have not received significant investments due to their low required management cost and to the adaptation capacity of goats to harsh environments [4,5,6]. Due to goats having high metabolic efficiency and behavioral mechanisms, they are the livelihoods of poor farmers; they provide tangible (e.g., milk, meat, and manure) and intangible (e.g., savings and cultural services) benefits to mountainous societies [7,8].
Previous researches have investigated and detailed the diet composition of goats in Mediterranean forest rangelands [9,10,11]. The available studies on the nutritive value of some browse species were mainly conducted in the northern [6,12] and eastern Mediterranean countries [13,14]. In the Southside of the Mediterranean forest, most of the studies [15,16] focused solely on a few lists of plant species (less than ten). Specific parts of these plants were separately analyzed (leaves, stems, and twigs) from shrub species and collected only for one period or throughout their vegetative cycle. Nevertheless, these findings did not consider the actually consumed parts of the plant at grazing time. Furthermore, the nutritive value of browsed plant species by goats has been unexplored on forest rangelands of the Southern Mediterranean of northern Morocco.
Moreover, forage quality is characterized by seasonal variations that could affect plant selection by grazing animals and, thus, diet quality and quantity and animal performance. However, there are differences in the degree of these variations depending on each regional climate and vegetation types [17,18,19]. Extensive grazing goat production systems in northern Morocco are affected by annual dry periods, resulting in reduced animal performance, and farm profitability [7,20]. The changes in chemical composition and digestibility of plant species with grazing seasons during two consecutive years have not been investigated previously.
In northern Morocco, the existing forest vegetation, mountainous topography, and animal adaptation explain the predominance of grazing goats in forest rangelands [20]. In this area, extensive goat farming plays an important socioeconomic role and contributes from (approximately) 68% to 100% of farmer incomes [21]. Therefore, this study was carried out to follow the temporal evolution in the chemical composition, in vitro digestibility, and metabolizable energy (ME) of each plant species selected by goats in the Southern Mediterranean forest rangeland of northern Morocco over three grazing seasons of two consecutive years.

2. Materials and Methods

2.1. Description of the Sampling Area

This research was conducted in a Southern Mediterranean forest rangeland of the Western Rif (35°14′ N; 5°30′ W; 300 to 520 m a.s.l), located in northern Morocco. The climate of the region is influenced by the Atlantic Ocean, dominated by Mediterranean humid to sub-humid conditions (dry in summer and wet in winter). The site was studied for two consecutive years under contrasting climatic conditions, with 270- and 755-mm rainfalls in 2016 and 2017, respectively. The mean annual precipitation was estimated to 700 mm, with a daily temperature range of 3–14 °C (minimum) and 18–38 °C (maximum) [11]. Based on meteorological data of this last two decades, the year 2016 could be considered as dry and 2017 as a wet year. The study area is mountainous and characterized by relatively rugged topography. This forest pasture is covered mainly with shrub strata resulting from oak forest degradation. The high formation includes Quercus ilex L. and Quercus suber L. associated with shrublands dominated by Arbutus unedo L., Cistus crispus L., Cistus monspeliensis L., and Erica arborea L. [22,23].

2.2. Source of Forage Samples

The study area was covered by heterogeneous vegetation composed mainly of three distinct groups of plant species: shrubs (A. unedo L., Calicotome villosa (Poir.) Link, Cistus spp. (inclusive of C. crispus L., C. monspeliensis L., and C. salviifolius L.), E. arborea L., Lavandula stoechas L., Myrtus communis L., Phillyrea media L., Pistacia lentiscus L., and Rubus ulmifolius Schott.), trees (Quercus spp. (inclusive of Q. canariensis L., Q. ilex L., and Q. suber L.), and Olea europaea var. sylvestris (Mill) Lehr), and herbaceous (mainly Anthemis cotula L., Brachypodium distachyon L., Bromus rigidus Roth, Calamintha nepeta (L.) Kuntze, Cynodon dactylon (L.) Pers., Eryngium tricuspidatum L., Lythrum junceum Banks and Sol., Rumex bucephalophorus L.). According to Chebli et al. [11], these plant species are listed as the main dietary components of goats in Southern Mediterranean forest rangelands. Grazing in the forest rangelands of northern Morocco is practiced only over three seasons (spring, summer, and autumn). For the winter, pasture access is very limited; goats do not browse in forest pastures and graze only in fallow land around the goat shed, which explains the exclusion of this season from the study. The present research studied the chemical composition, in vitro digestibility, and metabolizable energy of all browsed species by goats. Samples were collected by hand-plucked simulation of each ingested part of the plant species similar to those consumed by goats. Diet composition and hand-plucked simulation are briefly summarized here and described fully in Chebli et al. [11]. The study concerned the botanical composition of each consumed part of plant by goats. The sampling was undertaken in the last month of each studied season (May, August, and November). Representative hand-plucked samples per plant species (a mixture of leaves and green tender stems) and herbaceous, similar to those consumed by goats, were imitated seasonally. For the thorny species, we used scissors to clip the selected parts. For herbaceous species, they were mixed into a single group because of difficulty to identify all ingested species by goats during grazing and their low selectivity. For shrubs and trees, the samples were harvested per species in special bags, with three replications, and transported to the laboratory for analysis.

2.3. Laboratory Analysis

Chemical analyses and in vitro digestibility studies were performed on three independent samples of the hand-plucked forage of each ingested plant species by goats during each grazing season of two consecutive years.

2.3.1. Chemical Analysis

Collected samples were dried at 40 °C in a ventilated oven to minimize changes in tannins content and activity until reaching constant weight [24], and then milled with a sieve mesh size of 1 mm for analysis. Dry matter (DM), organic matter (OM), crude protein (CP), and ether extract (EE) were analyzed according to the Association of Official Analytical Chemists [25]. The neutral detergent fiber (NDF) was estimated using the Mertens [26] method with α-amylase and sodium sulfite. Acid detergent fiber (ADF) was determined according to method 973.18 of AOAC [27]. Acid detergent lignin (ADL) was determined by the solubilization of cellulose with sulfuric acid, according to Robertson and Van Soest [28]. All fiber extractions were performed using ANKOM 200 Fiber Analyzer® (ANKOM Technology, Fairport, NY, USA). The NDF, ADF, and ADL values were expressed inclusive of residual ash. Condensed tannins (CT) were predicted by Porter et al. [29] method using butanol-HCl, and ferric reagents.

2.3.2. In Vitro Digestibility and Metabolizable Energy

In vitro dry matter (IVDMD) and organic matter (IVOMD) digestibility were performed using DAISYII Incubator® (ANKOM Technology, Fairport, NY, USA) as described by Mabjeesh et al. [30]. This device is essentially based on the in vivo simulation of digestion [31]. The rumen liquor for incubation was collected from five goats at a communal slaughterhouse, as described by El Otmani et al. [32]. These goats grazed in similar forest rangeland of the study area. The collected ruminal fluid was maintained in a thermos at 39 °C to keep rumen microflora alive. A weight of 0.5 g of each sample was placed in ANKOM filter bags (F57) and was put in jars (24 bags/jar). The inoculum, mixture containing 4/5 volume of artificial saliva, and 1/5 of rumen liquor was added in jars and incubated at 39.5 °C for 48 h. IVDMD and IVOMD were estimated by quantifying residuals DM and OM comparing to incubated initial quantities.
The metabolizable energy (ME; MJ/kg DM) of each consumed plant species was calculated using the equation [27]:
ME = 0.17 × DMD – 2,
where DMD is the dry matter digestibility in percentage.

2.4. Statistical Analysis

Data were analyzed using SAS software® (SAS Inst. Cary, NC, USA). Chemical composition, digestibility, and ME of each plant species (n = 15) and herbaceous were analyzed using a general linear model (GLM) procedure of SAS in a factorial structure. Data were compared between seasons (i.e., spring, summer, and autumn), years (i.e., 2016 and 2017), and their interactions. Simple correlation analysis was used to establish the relationships between the chemical composition, IVOMD, and ME. The correlation plot was obtained by utilizing the “corrplot” library in the R-package [33]. For all analyses, the significance level was declared at p < 0.05. In case of significant effect, means were compared using the Tukey’s test.

3. Results

The chemical composition, IVOMD, and ME of the browsed plant species by goats at different sampling seasons and years are given in Table 1 (shrubs) and Table 2 (trees and herbaceous). Overall, these parameters of shrubs (n = 11), trees (n = 4), and herbaceous species varied seasonally in each studied year.
Across shrub species, all of them presented a higher DM content in summer, except for C. salviifoluis and M. communis, with a higher DM content in spring, and for R. ulmiformis, with a higher DM content (also) in autumn. The higher water content was observed either in spring (for five shrubs) or autumn (for six shrubs). This parameter was significantly affected by both studied factors (season and year) except for C. crispus, which was not affected by year. Their effects on OM of the studied shrub species were variable, A. unedo, E. arborea, L. stoechas, M. communis, and P. media, having the same OM throughout the year. The CP content varied significantly among seasons of both years (p < 0.05), except for the season effect of the dry year (2016) on C. villosa and the season effect of the wet year (2017) on A. unedo, C. crispus, C. salviifoluis, E. arborea, and P. media (p > 0.05). During both years, the highest and lowest CP concentrations were recorded in C. villosa (about 240 g/kg DM in the autumn) and A. unedo (about 60 g/kg DM in summer), respectively. The CT content ranged from 1.97 g/kg DM (summer 2017) in C. villosa to 191 g/kg DM (summer 2016) in P. lentiscus. The EE content ranged from 15.8 g/kg DM in C. crispus, to 90–101 g/kg DM in C. monspeliensis (summer and autumn) and E. arborea (spring and summer). The highest NDF and ADF levels of both years were observed in C. villosa, with 629 and 482 g/kg DM, respectively. Overall, the ADL contents showed a significant increase from spring to summer–autumn (except for P. media) with a range from 62.3 g/kg DM in R. ulmifolius (2016) to 324 g/kg DM in E. arborea (2017). All studied shrub species presented a higher IVOMD in spring of both consecutive years, except for R. ulmifolius and M. communis, with the highest IVOMD in summer–autumn and summer, respectively. The lower IVOMD were found in summer (for five shrubs) or autumn (for two shrubs), or there was no significant difference between summer and autumn (for two shrubs). The ME results showed the same trend as the IVOMD. The highest ME content was observed in L. stoechas (about 10 MJ/kg DM) browsed during spring and the lowest one (about 4.5 MJ/kg DM) in C. villosa (autumn) and E. arborea (summer) during both dry and wet years. The most notable changes due to advancing maturity were found in the CP, CT, ADL, and ME contents. Generally, the CP and ME contents decreased, and CT and ADL contents increased during spring to summer–autumn of both years.
Across trees species and for the two years, DM content was higher in summer and lower in spring, except for O. europaea, where it was the opposite. The DM content in autumn was the same as in spring for Q. suber, the same as in summer for O. europaea and Q. ilex, and significantly different from the two other seasons for Q. canariensis. Each oak tree species showed no variation of its OM content during a year, except for Q. suber in 2016, which presented a decrease over time. Q. ilex recorded the higher CP concentration in spring of the wet and dry years (99.7 and 114 g/kg DM, respectively). Q. suber had the highest, and O. europaea the lowest CT content during all studied seasons. Among tree species, the oak trees showed low EE content (about 24 g/kg DM). The high EE content was recorded in O. europaea during the summer of the wet year (131 g/kg DM). The highest NDF content was recorded for Q. suber and Q. ilex (about 550 g/kg DM), and the lowest for O. europaea (about 410 g/kg DM). The highest and lowest ADL levels were observed during autumn and spring of the wet year in Q. ilex (about 190 g/kg DM) and in Q. canariensis (103 g/kg DM), respectively. Quercus spp. had a high IVOMD significantly during spring and a low one in summer. The IVOMD of O. europaea was similar in all studied seasons during the dry year. Nevertheless, this similarity was not observed in the wet year, with a slight increase over time. The ME levels of the studied tree species varied slightly, being particularly low in Quercus spp. (about 6 MJ/kg DM) during the summer of the dry and wet years and highest in Q. canariensis (8.8 MJ/kg DM) during the two springs.
Comparatively to the two other groups, herbaceous also had a higher DM content in summer and a lower content in spring. The OM content was variable according to the year and season. It was higher in the spring and similar and lower in the other seasons. The CP concentrations recorded the highest value in the spring of 2016 and 2017 (156 and 142 g/kg DM, respectively; p < 0.001). The CT contents recorded the highest values during the summer of 2016 (4.17 g/kg DM; p < 0.05) but were similar among seasons of 2017 (with about 2.7 g/kg DM; p = 0.116). The NDF concentrations were similar among seasons of both years (p > 0.05). The ADF content was higher in summer than spring and autumn of both years (p < 0.01). The ADL concentrations were similar among seasons of 2016 (about 70 g/kg DM; p = 0.265) but increased in 2017 from spring to summer (p < 0.05). The IVOMD and ME contents were higher in spring than in autumn and summer (p < 0.001). Overall, the highest CP and ME contents were recorded in the herbaceous and the lowest in shrub and tree species. An opposite trend was recorded for CT and ADL levels.
The correlation values among the chemical composition, IVOMD, and ME from the studied forage species are presented in Figure 1. The CP showed a negative correlation with ADL, CT, and EE (p < 0.001). The ME was strongly correlated with IVOMD (p < 0.001). The NDF, ADF, and ADL contents were positively correlated with each other (p < 0.001). A negative correlation was observed between IVOMD and ME with CT, ADL, and ADF (p < 0.001), and with NDF (p < 0.05).

4. Discussion

The aim of this study was to assess the nutritive value of the plant species browsed by goats and their variations throughout three grazing seasons of two years. These years appeared very contrasted regarding the mean annual rainfall, with a dry year in 2016 and a wet one in 2017. According to Papachristou et al. [34], the bulk of the grazing goats’ diet includes few ligneous and herbaceous species, representing less than ten species. Ligneous species A. unedo, C. villosa, E. arborea, M. communis, P. lentiscus, and Q. suber are considered the most widespread species in the Southern Mediterranean rangelands [15,23]. As observed and described by Chebli et al. [11], plant species analyzed herein represent the all-selected diet by grazing goats in Southern Mediterranean forest rangelands. During spring of 2016, the contribution of C. monspeliensis (28.8%), C. crispus (19.8%), and C. salviifolius (17.6%) was the highest followed by L. stoechas (17.3%) and herbaceous (7%). These species contributed lowly to the diet during autumn and summer (< 3%). In the autumn and summer, the diet proportion of Quercus spp. (3–20%), M. communis (14–19.4%), P. lentiscus (8–13%), A. unedo (11–13%), E. arborea (9.5–11%), and O. europaea (2–7%) was largely significant. During spring of 2017, the contribution of C. crispus was significantly increased by 42% with the decreased rate of C. salviifolius and L. stoechas by 10 and 15%, respectively. In the autumn, the greatest increase in contribution to the diet was observed for O. europaea followed by P. lentiscus, and E. arborea. The opposite trend was observed with the diet proportion of Q. canariensis and C. villosa. In the summer, the contribution of P. lentiscus and P. media increased by 93 and 17%, respectively. On the other hand, diet contribution of A. unedo and E. arborea decreased by 35 and 17%, respectively. On average, the diet of the goats was largely composed of shrubs (64–90%) and trees (2–35%). However, the contribution of herbaceous did not exceed 8%. The contribution of trees to the diet during spring dropped from 30.3 to 3.7% and from 29.0 to 2.2% in 2016 and 2017, respectively. The diet proportion of R. ulmifolius varied from 0.01 to 3.4% [11].
The nutritive value of the hand-plucked samples, corresponding to the most tender part of the plant, appears to reflect the quality of the diet consumed by grazing goats [6]. For this study, the browsed parts of the plant by goats were analyzed, which represent a mixture of leaves, stems, and twigs.
The chemical traits of browsed species were extremely wide, which are in accordance with previous studies conducted in northern and eastern Mediterranean forest rangelands [6,12,13,14], deciduous tropical forest [35], South African rangeland [36]. In northwestern Italy, Ravetto Enri et al. [37] reported the relevant effect of the vegetative season on chemical composition and in vitro true digestibility of four tree species selected by goats. These wide variations on the nutritional proprieties of plant species could be explained by soil fertility [15,38], environmental conditions, and stage of growth or age [14,39].
The observed mean CP level found in this study varied from 60 to 240 g/kg DM. Most of the analyzed plant species present high levels of CP than the minimum level of 70–80 g/kg DM required by microorganisms for optimum rumen functioning and feed intake in ruminant livestock [40]; a lower CP content affects negatively feed intake and digestibility [41]. In the present study, the low CP content was particularly recorded in A. unedo (from 52.7 to 70.7 g/Kg DM). In the northwest of Tunisia, a value of 55 g/kg DM in A. unedo, collected in March 1998 from the uplands of Taaref, was reported [15], which is in the range of the current results. The high proportions of mature leaves and twigs in the samples could explain the low CP level in some plant species, such as A. unedo. Overall, the average CP level was higher during spring 2016 in all species because plants contain the maximum CP content during the vegetative stage [42]. The decrease of this parameter in the summer agrees with the literature [19,43] because CP drops with the physiological maturity stage of the plant [44], which explains the negative correlation of CP with ADL and EE that increase with plant maturity as found by Ammar et al. [12] in some Spanish shrub species. As expected, the CP content was higher in C. villosa as it is a leguminous plant. According to Kokten et al. [14], leaves of C. villosa could be used as protein supplements for livestock since their CP contents are high compared to the other Mediterranean shrubs. The high protein level in C. villosa could be attributed to the ability of this plant to fix atmospheric nitrogen thanks to rhizobia associated with their nodules [12,16]. Overall, the older leaves contained less CP and more fiber than the young and tender part of the selected plant species. This statement is in consistent with other studies [6,14]. In terms of CP content, many of the woody species cover the daily maintenance requirements of grazing goat but not for milk or meat production needs, which is above 130 g/kg DM. In another environment (hills of Nepal), Khanal and Subba [45] reported a good nutritional value of leaves from most of the tree fodder species, with a minimum CP of 110 g/kg DM.
Generally, trees had a higher EE content during summer, which coincides with the maturity stage of these groups of plant species. Indeed, plant species had a higher fat content (EE) in the late physiological stage that increases with maturity [12].
According to the species and sampling season, the NDF, ADF, and ADL contents in ligneous species varied from 242 to 629 g/kg, 186 to 482 g/kg, and 70 to 322 g/kg, respectively. However, herbaceous recorded the lower lignin content (64.3–89 g/kg). Overall, these contents significantly increased by advancing maturity. The results are in line with the findings of several authors [14,16,46], who indicated that cell wall content (NDF, ADF, and ADL) augmented with maturity (cell wall lignification). All analyzed samples recorded higher ADL levels during the spring of the dry year compared to the wet year. This higher concentration during the dry year could reflect the response of plant species to water stress (rarity rains), which is associated with the increased level of tannins. Khanal and Subba [45] reported a high ADL content in most of the fodder trees in the hills of Nepal, with values more than 100 g/kg DM. High fiber content, and lignin especially, means low free-nitrogen extract and soluble carbohydrates contents, which explain the observed negative correlations of fibers with IVOMD and ME. Ammar et al. [47] reported an increase of fiber content in parallel with a decrease of in vitro digestibility, with the maturity of mountain grasses, which confirm the negative correlation between fibers and digestibility.
The observed CT concentrations varied from 2.3 to 184 g/kg DM, showing significantly different and slightly higher values than those obtained by several authors [12,48] with shrub leaves from Northern Spain. These variations could be due in part to the difference in analysis methods. Moreover, it could be owing to the stage of growth and the sampled parts of the plants (leaves, stems, and twigs), to the season and to the nature of the sampling site [49,50]. In addition, the current study concerns the analysis of different parts of the plant selected by goats, not only their leaves.
A CT concentration of 20–45 g/kg DM has a negative effect on protein digestibility and proteolytic bacteria [51], and a concentration above 55 g/kg DM reduces the voluntary feed intake of grazing ruminants [52,53]. Thus, except for herbaceous (2.20–4.17 g/kg DM), C. villosa, L. stoechas, P. media, and O. europaea, all pastoral species had a CT content higher than this maximum level. However, even with a high CT content, the shrubs were highly consumed in spring and autumn [11]. These findings are consistent with Fomum et al. [36] and Mkhize et al. [54], who reported no correlation between CT and feed intake in goats. It could be explained by the ability of goat to balance their diet and dilute secondary compounds by consuming a mixture of plant species [55,56]. Moreover, grazing animals exposed to high CT feed could excrete more saliva richer in proline-rich proteins that has the ability to bind with CT to neutralize it [52,54]. Nevertheless, goats have a specificity compared to other ruminants that their ruminal microbiota is able to valorize feed with low nutritional values due to their cellulolytic bacteria and the tanninase activity [57]. Min et al. [51], reported that high CT concentrations reduced digestibility, which could explain the negative correlation between CT and IVOMD.
Bartolomé et al. [58], who studied the quality of forest resources in the undergrowth of the pine forests of Mallorca (Spain), reported that species, such as P. lentiscus, C. monspeliensis, and A. unedo, could be interesting as feed for goats, as they show protein levels above the minimum of maintenance, and at the same time, high digestibility. Nevertheless, they present a high content of secondary compounds, such as tannins.
In the present study, the higher values of digestibility, mainly observed in spring, are attributed to its negative correlation with ADF and ADL. Ammar et al. [12], also reported a negative correlation between fibers and digestibility of browsed leaves. Most of the pastoral species had low digestibility and, consequently, a low energy content (IVOMD < 550 g/kg; ME < 8 MJ/kg DM), except for herbaceous and some shrubs (A. unedo and Cistus spp.), as their values varied from medium (IVOMD: 550–700 g/kg; ME: 8–10 MJ/kg DM) to higher nutritional values (IVOMD > 700 g/kg; ME > 10 MJ/kg DM), especially during spring [59]. As reported by Paton [60], ME depends mainly on IVOMD, which could explain their high positive correlation. Overall, goats select species with high CP and digestibility and low fiber content basis, in accordance with the literature [34].

5. Conclusions

This study provides a valuable and useful database on the temporal variations in chemical composition, in vitro digestibility, and ME of the main plant species browsed by goats in the Southern Mediterranean forest rangeland. Most of these plant species showed considerable variation among grazing season. In general, the nutritive value of plant species was highest in spring, and then steadily decreased through the summer. In autumn, the nutritive value decreased, remained the same, or increased compared to summer. Most of the selected plant species presented high levels of CP than the minimum required levels for maintenance needs. All analyzed samples recorded higher lignin levels during the spring of the dry year compared to the wet year. Except for herbaceous, C. villosa, L. stoechas, P. media, and O. europaea, all analyzed species had a CT content higher than this maximum level. The high values of digestibility in spring are attributed to its negative correlation with ADF and lignin. Owing to the morphological and physiological differences between the consumed plant species, changes in chemical compositions and in vitro digestibility could be expected. Consequently, goats are forced to adapt their browsing behavior to the low-quality vegetation, typical of the Mediterranean forest. The results could be used as indicators to assess the nutritional value of the goat diets in forest rangelands throughout the year to help the grazing management strategy, and/or to eventually supplement the goats adequately, to prevent the low farmer incomes due to the animal performance decrease.
Future work would allow knowing if the practiced grazing systems in the Southern Mediterranean region could guarantee the seasonal dietary requirements of grazing goats, in terms of energy and protein, taking into account the physical and physiological conditions of grazing goats.

Author Contributions

Conceptualization, Y.C.; methodology, Y.C. and S.E.O.; formal analysis, Y.C., S.E.O., M.C., and J.-F.C.; investigation, Y.C. and S.E.O.; data curation, Y.C., S.E.O., M.C., J.-L.H., and J.-F.C.; writing—original draft preparation, Y.C.; writing review and editing, Y.C., S.E.O., M.C., J.-L.H., and J.-F.C.; project administration, Y.C. and J.-F.C.; funding acquisition, Y.C. All authors have read and agreed to the published version of the manuscript.

Funding

This project received funding from the Academy for Research and Higher Education-Development Cooperation Committee (ARES-CCD), Brussels, Belgium. This study is realized in the framework of the Research Project for Development: PRD (2013–2018).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The data presented in this study are available from the corresponding author on request.

Acknowledgments

The authors would like to thank the Research for Development Project collaborators. We thank the herders for their patience and participation in this study.

Conflicts of Interest

The authors declare no conflict of interest.

References

  1. Butchart, S.H.M.; Walpole, M.; Collen, B.; Van Strien, A.; Scharlemann, J.P.W.; Almond, R.E.A.; Baillie, J.E.M.; Bomhard, B.; Brown, C.; Bruno, J.; et al. Global biodiversity: Indicators of recent declines. Science 2010, 328, 1164–1168. [Google Scholar] [CrossRef] [PubMed]
  2. Roces-Díaz, J.V.; Vayreda, J.; De Cáceres, M.; García-Valdés, R.; Banqué-Casanovas, M.; Morán-Ordóñez, A.; Brotons, L.; de-Miguel, S.; Martínez-Vilalta, J. Temporal changes in mediterranean forest ecosystem services are driven by stand development, rather than by climate-related disturbances. Forest Ecol. Manag. 2021, 480, 118623. [Google Scholar] [CrossRef]
  3. Buira, A.; Fernández-Mazuecos, M.; Aedo, C.; Molina-Venegas, R. The contribution of the edaphic factor as a driver of recent plant diversification in a Mediterranean biodiversity hotspot. J. Ecol. 2021, 109, 987–999. [Google Scholar] [CrossRef]
  4. Silanikove, N. The physiological basis of adaptation in goats to harsh environments. Small Rumin. Res. 2000, 35, 181–193. [Google Scholar] [CrossRef]
  5. Arsenos, G.; Gelasakis, A.I.; Pinopoulos, S. Description and typology of dairy goat farms in Greece. In Proceedings of the 4th ISOFAR Scientific Conference, Istanbul, Turkey, 13–15 October 2014; pp. 571–574. [Google Scholar]
  6. Manousidis, T.; Parissi, Z.M.; Kyriazopoulos, A.P.; Malesios, C.; Koutroubas, S.D.; Abas, Z. Relationships among nutritive value of selected forages, diet composition and milk quality in goats grazing in a Mediterranean woody rangeland. Livest. Sci. 2018, 218, 8–19. [Google Scholar] [CrossRef]
  7. Chentouf, M.; Boulanouar, B.; Bister, J.L. Elevage Caprin au Nord du Maroc; INRA-Editions Press: Rabat, Morocco, 2015. [Google Scholar]
  8. Fantahun, T.; Alemayehu, A.; Abegaz, S. Characterization of goat production systems and trait preferences of goat keepers in Bench Maji zone, south western Ethiopia. Afr. J. Agric. Res. 2016, 11, 2768–2774. [Google Scholar]
  9. Glasser, T.A.; Landau, S.Y.; Ungar, E.D.; Perevolotsky, A.; Dvash, L.; Muklada, H.; Kababya, D.; Walker, J.W. Foraging selectivity of three goat breeds in a Mediterranean shrubland. Small Rumin. Res. 2012, 102, 7–12. [Google Scholar] [CrossRef]
  10. Manousidis, T.; Kyriazopoulos, A.P.; Parissi, Z.M.; Abraham, E.M.; Korakis, G.; Abas, Z. Grazing behavior, forage selection and diet composition of goats in a Mediterranean woody rangeland. Small Rumin. Res. 2016, 145, 142–153. [Google Scholar] [CrossRef]
  11. Chebli, Y.; Otmani, S.E.; Chentouf, M.; Hornick, J.-L.; Bindelle, J.; Cabaraux, J.-F. Foraging Behavior of Goats Browsing in Southern Mediterranean Forest Rangeland. Animals 2020, 10, 196. [Google Scholar] [CrossRef] [Green Version]
  12. Ammar, H.; López, S.; González, J.S.; Ranilla, M.J. Chemical composition and in vitro digestibility of some Spanish browse plant species. J. Sci. Food Agric. 2004, 84, 197–204. [Google Scholar] [CrossRef]
  13. Parlak, A.O.; Gokkus, A.; Hakyemez, B.H.; Baytekin, H. Shrub yield and forage quality in Mediterranean shrublands of West turkey for a period of one year. African J. Agric. Res. 2011, 6, 1726–1734. [Google Scholar] [CrossRef]
  14. Kokten, K.; Kaplarn, M.; Hatipoǧlu, R.; Saruhan, V.; Çinar, S. Nutritive value of Mediterranean shrubs. J. Anim. Plant Sci. 2012, 22, 188–194. [Google Scholar]
  15. Ammar, H.; López, S.; González, J.S. Assessment of the digestibility of some Mediterranean shrubs by in vitro techniques. Anim. Feed Sci. Technol. 2005, 119, 323–331. [Google Scholar] [CrossRef]
  16. Mebirouk-Boudechiche, L.; Cherif, M.; Boudechiche, L.; Sammar, F. Teneurs en composés primaires et secondaires des feuilles d’arbustes fourragers de la région humide d’Algérie. Revue Méd. Vét. 2014, 165, 344–352. [Google Scholar]
  17. Van, D.T.T.; Mui, N.T.; Ledin, I. Tropical foliages: Effect of presentation method and species on intake by goats. Anim. Feed Sci. Technol. 2005, 118, 1–17. [Google Scholar] [CrossRef]
  18. Celaya, R.; Oliván, M.; Ferreira, L.M.M.; Martínez, A.; García, U.; Osoro, K. Comparison of grazing behaviour, dietary overlap and performance in non-lactating domestic ruminants grazing on marginal heathland areas. Livest. Sci. 2007, 106, 271–281. [Google Scholar] [CrossRef]
  19. Safari, J.; Mushi, D.E.; Kifaro, G.C.; Mtenga, L.A.; Eik, L.O. Seasonal variation in chemical composition of native forages, grazing behaviour and some blood metabolites of Small East African goats in a semi-arid area of Tanzania. Anim. Feed Sci. Technol. 2011, 164, 62–70. [Google Scholar] [CrossRef]
  20. Chebli, Y.; Chentouf, M.; Ozer, P.; Hornick, J.L.; Cabaraux, J.F. Forest and silvopastoral cover changes and its drivers in northern Morocco. Appl. Geogr. 2018, 101, 23–35. [Google Scholar] [CrossRef]
  21. Chentouf, M.; Zantar, S.; Doukkali, M.R.; Farahat, L.B.; Jouamaa, A.; Aden, H. Performances techniques et économiques des élevages caprins dans le nord du Maroc. Options Méditerranéennes 2011, 100, 151–156. [Google Scholar]
  22. Mastere, M.; Van Vliet-Lanoë, B.; Ait Brahim, L. Land use mapping and its relation to mass wasting and gullying in North-Western Rif (Morocco). Géomorphologie 2013, 193, 335–352. [Google Scholar] [CrossRef]
  23. Chebli, Y.; Chentouf, M.; Hornick, J.L.; Cabaraux, J.F. Extensive goat production systems in northern Morocco: Production and use of pastoral resources. In Grassland Resources for Extensive Farming Systems in Marginal Lands: Major Drivers and Future scenarios; Porqueddu, A., Franca, C., Lombardi, A., Molle, G., Peratoner, G., Hopkins, G., Eds.; Wageningen Academic Publishers Press: Wageningen, The Netherlands, 2017; pp. 131–133. [Google Scholar]
  24. Makkar, H.P.S.; Singh, B. Effect of drying conditions on tannin, fiber and lignin levels in mature oak (Quercus incana) leaves. J. Sci. Food Agric. 1991, 54, 323–328. [Google Scholar] [CrossRef]
  25. AOAC (Association of Official Analytical Chemists). Official Methods of Analysis, 16th ed.; AOAC International: Arlington, VA, USA, 1997. [Google Scholar]
  26. Mertens, D.R. Gravimetric determination of amylase-treated neutral detergent fiber in feeds with refluxing beakers or crucibles: Collaborative study. J. AOAC Int. 2002, 85, 1217–1240. [Google Scholar] [PubMed]
  27. AOAC (Association of Official Analytical Chemists). Official Methods of Analysis, 15th ed.; AOAC International: Arlington, VA, USA, 1990. [Google Scholar]
  28. Robertson, J.B.; Van Soest, P.J. The detergent system of analysis. In The Analysis of Dietary Fiber in Food; James, W.P.T., Theander, O., Eds.; Marcel Dekker: New York, NY, USA, 1981; pp. 123–158. [Google Scholar]
  29. Porter, L.J.; Hrstich, L.N.; Chan, B.G. The conversion of procyanidins and prodelphinidins to cyanidin and delphinidin. Phytochemistry 1986, 25, 223–230. [Google Scholar] [CrossRef] [Green Version]
  30. Mabjeesh, S.J.; Cohen, M.; Arieli, A. In vitro methods for measuring the dry matter digestibility of ruminant feedstuffs: Comparison of methods and inoculum source. J. Dairy Sci. 2000, 83, 2289–2294. [Google Scholar] [CrossRef]
  31. Tassone, S.; Fortina, R.; Peiretti, P.G. In Vitro Techniques Using the DaisyII Incubator for the Assessment of Digestibility: A Review. Animals 2020, 10, 775. [Google Scholar] [CrossRef]
  32. El Otmani, S.; Chentouf, M.; Hornick, J.L.; Cabaraux, J.F. Chemical composition and in vitro digestibility of alternative feed resources for ruminants in Mediterranean climates: Olive cake and cactus cladodes. J. Agric. Sci. 2019, 157, 260–271. [Google Scholar] [CrossRef]
  33. R Core Team. R: A Language and Environment for Statistical Computing; R Foundation for Statistical Computing: Vienna, Austria, 2019. [Google Scholar]
  34. Papachristou, T.G.; Platis, P.D.; Nastis, A.S. Foraging behaviour of cattle and goats in oak forest stands of varying coppicing age in Northern Greece. Small Rumin. Res. 2005, 59, 181–189. [Google Scholar] [CrossRef]
  35. González-Pech, P.G.; de Jesús Torres-Acosta, J.F.; Sandoval-Castro, C.A.; Tun-Garrido, J. Feeding behavior of sheep and goats in a deciduous tropical forest during the dry season: The same menu consumed differently. Small Rumin. Res. 2015, 133, 128–134. [Google Scholar] [CrossRef]
  36. Fomum, S.W.; Scogings, P.F.; Dziba, L.; Nsahlai, I.V. Seasonal variations in diet selection of Nguni goats: Effects of physical and chemical traits of browse. Afr. J. Range Forage Sci. 2015, 32, 193–201. [Google Scholar] [CrossRef]
  37. Ravetto Enri, S.; Probo, M.; Renna, M.; Caro, E.; Lussiana, C.; Battaglini, L.M.; Lombardi, G.; Lonati, M. Temporal variations in leaf traits, chemical composition and in vitro true digestibility of four temperate fodder tree species. Anim. Prod. Sci. 2020, 60, 643–658. [Google Scholar] [CrossRef]
  38. Jackson, F.S.; Barry, T.N.; Lascona, C.; Palmer, B. The extractable and bound condensed tannin content of leaves from tropical tree. J. Sci. Food Agric. 1996, 71, 103–110. [Google Scholar] [CrossRef]
  39. Dzowela, B.H.; Hove, L.; Topps, J.H.; Mafongoya, P.L. Nutritional and anti-nutritional characters and rumen degradability of dry matter and nitrogen for some multipurpose tree species with potential for agroforestry in Zimbabwe. Anim. Feed Sci. Technol. 1995, 55, 207–214. [Google Scholar] [CrossRef]
  40. Van Soest, P.J.; Robertson, J.B.; Lewis, B.A. Methods for dietary fiber, neutral detergent fiber, and nonstarch polysaccharides in relation to animal nutrition. J. Dairy Sci. 1991, 74, 3583–3597. [Google Scholar] [CrossRef]
  41. Minson, D.J. Forage in Ruminant Nutrition; Academic Press: New York, NY, USA, 1990. [Google Scholar]
  42. Mero, R.N.; Udén, P. Promising tropical grasses and legumes as feed resources in central Tanzania II. In sacco rumen degradation characteristics of four grasses and legumes. Anim. Feed Sci. Technol. 1997, 69, 341–352. [Google Scholar] [CrossRef]
  43. Hassen, A.; Rethman, N.F.G.; van Niekerk, W.A.; Tjelele, T.J. Influence of season/year and species on chemical composition and in vitro digestibility of five Indigofera accessions. Anim. Feed Sci. Technol. 2007, 136, 312–322. [Google Scholar] [CrossRef]
  44. Rawnsley, R.R.; Donaghy, D.J.; Fulkerson, W.J.; Lane, P.A. Changes in the physiology and feed quality of cocksfoot (Dactylis glomerata L.) during regrowth. Grass Forage Sci. 2002, 57, 203–211. [Google Scholar] [CrossRef] [Green Version]
  45. Khanal, R.C.; Subba, D.B. Nutritional evaluation of leaves from some major fodder trees cultivated in the hills of Nepal. Anim. Feed Sci. Technol. 2001, 92, 17–32. [Google Scholar] [CrossRef]
  46. Kamalak, A. Determination of nutritive value of leaves of a native grown shrub, Glycyrrhiza glabra L. using in vitro and in situ measurements. Small Rumin. Res. 2006, 64, 268–278. [Google Scholar] [CrossRef]
  47. Ammar, H.; López, S.; Bochi-Brum, O.; Garcia, R.; Ranilla, M.J. Composition and in vitro digestibility of leaves and stems of grasses and legumes harvested from permanent mountain meadows at different stages of maturity. J. Anim. Feed Sci. 1999, 84, 599–610. [Google Scholar] [CrossRef]
  48. Alvarez del Pino, M.C.Á.; Hervás, G.; Mantecón, Á.R.; Giráldez, F.J.; Frutos, P. Comparison of biological and chemical methods, and internal and external standards, for assaying tannins in Spanish shrub species. J. Sci. Food Agric. 2005, 85, 583–590. [Google Scholar] [CrossRef]
  49. Makkar, H.P.S.; Becker, K. Vanillin-HCl method for condensed tannins: Effect of organic solvents used for extraction of tannins. J. Chem. Ecol. 1993, 19, 613–621. [Google Scholar] [CrossRef] [PubMed]
  50. Salem, A.F.Z.M. Impact of season of harvest on in vitro gas production and dry matter degradability of Acacia saligna leaves with inoculum from three ruminant species. Anim. Feed Sci. Technol. 2005, 123–124, 67–79. [Google Scholar] [CrossRef]
  51. Min, B.R.; Barry, T.N.; Attwood, G.T.; McNabb, W.C. The effect of condensed tannins on the nutrition and health of ruminants fed fresh temperate forages: A review. Anim. Feed Sci. Technol. 2003, 106, 3–19. [Google Scholar] [CrossRef]
  52. Waghorn, G. Beneficial and detrimental effects of dietary condensed tannins for sustainable sheep and goat production-progress and challenges. Anim. Feed Sci. Technol. 2008, 147, 116–139. [Google Scholar] [CrossRef]
  53. Dove, H. Ingestive behaviour, diet selection, and feed intake. In Goat Science and Production; Solaiman, S.G., Ed.; Wiley-Blackwell: Ames, IA, USA, 2010; pp. 179–192. [Google Scholar]
  54. Mkhize, N.R.; Scogings, P.F.; Nsahlai, I.V.; Dziba, L.E. Diet selection of goats depends on season: Roles of plant physical and chemical traits. Afr. J. Range Forage 2014, 31, 209. [Google Scholar] [CrossRef]
  55. Iason, G.R.; Villalba, J.J. Behavioral strategies of mammal herbivores against plant secondary metabolites: The avoidance-tolerance continuum. J. Chem. Ecol. 2006, 32, 1115–1132. [Google Scholar] [CrossRef]
  56. Rogosic, J.; Pfister, J.A.; Provenza, F.D.; Grbesa, D. Sheep and goat preference for and nutritional value of Mediterranean maquis shrubs. Small Rumin. Res. 2006, 64, 169–179. [Google Scholar] [CrossRef]
  57. Molina-Alcaide, E.; Yáñez-Ruiz, D.R. Potential use of olive by-products in ruminant feeding: A review. Anim. Feed Sci. Technol. 2008, 147, 247–264. [Google Scholar] [CrossRef]
  58. Bartolomé, J.; Baraza, E.; Rita, J.; Serrano, E.; Pareja, J.; Ibáñez, M.; Capó, M.; Alomar, D.; Domenech, O. Cabra i forest: La qualitat dels recursos del bosc per a la cabra mallorquina. Mon. Soc. Hist. Nat. Balears 2019, 28, 95–100. [Google Scholar]
  59. Tainton, N. Veld Management in South Africa; University of Natal Press: Pietermaritzburg, South Africa, 1999. [Google Scholar]
  60. Paton, D. Elaboration of a multi-variate model for the determination of the metabolizable energy of Mediterranean bushes based on chemical parameters. J. Arid Environ. 2003, 53, 271–280. [Google Scholar] [CrossRef]
Figure 1. Correlation plot between the chemical composition, IVOMD, and ME from forage species browsed by goats. Positive and negative correlation coefficients are displayed in blue and brown scale, respectively. EE, ether extract; NDF, neutral detergent fiber; ADF, acid detergent fiber; ADL, lignin; CP, crude protein; CT, condensed tannins; IVOMD, in vitro organic matter digestibility; ME, metabolizable energy. Significance level (*** < 0.001, ** < 0.01, and * < 0.05).
Figure 1. Correlation plot between the chemical composition, IVOMD, and ME from forage species browsed by goats. Positive and negative correlation coefficients are displayed in blue and brown scale, respectively. EE, ether extract; NDF, neutral detergent fiber; ADF, acid detergent fiber; ADL, lignin; CP, crude protein; CT, condensed tannins; IVOMD, in vitro organic matter digestibility; ME, metabolizable energy. Significance level (*** < 0.001, ** < 0.01, and * < 0.05).
Animals 11 01441 g001
Table 1. Chemical composition (g/kg DM), IVOMD (g/kg), and ME (MJ/kg DM) of shrub species (n = 11) browsed by goats in Southern Mediterranean forest rangeland of northern Morocco during two contrasting years.
Table 1. Chemical composition (g/kg DM), IVOMD (g/kg), and ME (MJ/kg DM) of shrub species (n = 11) browsed by goats in Southern Mediterranean forest rangeland of northern Morocco during two contrasting years.
Item2016 (Dry Year) 2017 (Wet Year)SEMp-Value (2016–2017)
SpringSummerAutumnSEMp-ValueSpringSummerAutumnSEMp-ValueSYY × S
Arbutus unedo
DM576 b660 a445 c31.6<0.001550 b647 a427 c32.3<0.00122<0.0010.0350.828
OM9629649554.70.7689749749683.720.7753.230.60.1160.99
CP69.8 a52.7 b60.3 ab2.790.0096067.370.72.240.131.840.1560.0540.003
CT91.4 b112 ab121 a5.10.01483.7 b101 ab110 a4.560.0183.53<0.0010.0280.926
EE70.3 b91.3 a67.7 b4.110.00573.3 b96.6 a72.3 b4.270.0032.93<0.0010.1380.936
NDF354 b485 a482 a23.50.004344 b491 a488 a26.10.00317<0.0010.9720.884
ADF257 c324 b363 a16<0.001243 c346 b377 a20.4<0.00112.6<0.0010.2030.056
ADL110 b172 a185 a12.70.004105 b191 a197 a15.70.0019.82<0.0010.3240.538
IVOMD603 a506 b387 c31.5<0.001617 a512 b405 c30.8<0.00121.4<0.0010.0370.637
ME9.11 a7.34 b5.13 c0.582<0.0019.23 a7.56 b5.20 c0.591<0.0010.403<0.0010.3720.899
Calicotome villosa
DM228 c487 a366 b37.6<0.001215 c471 a326 b37.1<0.00125.7<0.0010.0020.138
OM927 b984 a950 ab10.10.038937 b988 a961 ab9.330.0446.750.0020.3830.95
CP17519023210.90.059161 b201 ab238 a12.90.0168.180.0010.9440.603
CT3.812.432.970.4390.4922.871.972.090.2020.1330.2520.1780.1370.905
EE31.3 a27.0 a22.0 b1.440.00234.3 a30.3 b30.7 ab0.7770.0330.999<0.001<0.0010.019
NDF520 b619 a593 a15.3<0.001511 b629 a601 a18.1<0.00111.5<0.0010.6320.425
ADF417 b464 a417 b8.630.007406 b482 a429 b120.0027.22<0.0010.3660.212
ADL95.7 b124 a117 a4.650.00494.7 c138 a128 b7.08<0.0014.19<0.0010.0170.028
IVOMD545 a439 b351 c28.4<0.001554 a443 b362 c28.1<0.00119.4<0.0010.2390.906
ME7.74 a6.17 b4.42 c0.485<0.0017.89 a6.16 b4.49 c0.499<0.0010.338<0.0010.60770.889
Cistus crispus
DM414 b528 a344 c27<0.001399 b514 a332 c26.9<0.00118.6<0.0010.0640.983
OM945 a946 a914 b6.560.049963 a951 ab927 b6.560.0344.730.0020.0830.695
CP113 a60.3 b76.0 b8.710.00999.779.385.35.050.2664.920.0020.4510.166
CT15.1 b65.0 a61.7 a8.2<0.00113.7 b61.3 a54.0 a7.51<0.0015.42<0.0010.0830.537
EE15.8 b21.8 a17.9 ab0.9860.0111923.320.30.8730.0980.70.0010.0180.758
NDF309 b242 c384 a20.8<0.001305 b256 c393 a20.3<0.00114.1<0.0010.3180.43
ADF266 a207 b252 a10.40.022512262657.460.0766.250.0020.5290.257
ADL103 b184 a170 a12.9<0.00193.0 b195 a184 a16.4<0.00110.1<0.0010.2610.101
IVOMD642 a408 c488 b34.3<0.001652 a412 c498 b35.2<0.00123.9<0.0010.0390.676
ME9.63 a5.59 c7.12 b0.589<0.0019.76 a5.71 c7.20 b0.592<0.0010.405<0.0010.0590.891
Cistus monspeliensis
DM585 b698 a379 c46.8<0.001573 b676 a366 c45.8<0.00131.8<0.0010.0110.717
OM905 c952 a930 b6.94<0.001918 c966 a947 b7.32<0.0015.21<0.001<0.0010.832
CP98.4 a84.7 ab66.3 b5.360.01588.0 ab98.7 a73.7 b4.390.0323.390.0010.4050.088
CT46.0 c65.6 b78.0 a4.82<0.00140.7 b54.7 a65.7 a3.930.0043.23<0.0010.0010.439
EE56.4 b90.6 a96.0 a6.39<0.00160.6 b98.6 a101 a6.78<0.0014.57<0.0010.0610.845
NDF388 c492 a434 b15.2<0.001377 c503 a440 b18.3<0.00111.6<0.0010.5880.062
ADF220 c255 b314 a13.8<0.001206 c274 b325 a17.6<0.00110.9<0.0010.230.028
ADL172 b176 b205 a5.810.007162 c187 b221 a8.72<0.0015.13<0.0010.1640.04
IVOMD592 a407 c489 b26.8<0.001601 a412 c501 b27.4<0.00118.6<0.0010.0120.617
ME8.27 a5.39 c6.72 b0.417<0.0018.46 a5.49 c6.77 b0.431<0.0010.291<0.0010.0530.562
Cistus salviifolius
DM488 a441 b366 c18<.0001477 a424 b353 c18.2<0.00112.5<0.0010.0190.913
OM876 b854 b906 a80.001890 b861 c921 a9.240.0016.1<0.0010.0210.714
CP108 a80.7 b70.7 b5.73<0.00195.794832.680.0913.11<0.0010.1070.002
CT25.0 c78.0 a49.7 b7.96<0.00121.0 c61.0 a41.7 b5.91<0.0014.95<0.0010.0060.215
EE23.3 b50.3 a40.6 a4.230.00227.3 c63.6 a46.3 b5.42<0.0013.46<0.0010.0070.274
NDF417 c506 a485 b13.6<0.001406 c515 a496 b16.9<0.00110.5<0.0010.4340.025
ADF252 c341 a291 b13.23<0.001239 c360 a303 b17.7<0.00110.7<0.0010.2820.071
ADL154 c228 a206 b11.1<0.001142 b238 a224 a15.2<0.0019.14<0.0010.080.003
IVOMD602 a439 b440 b27.5<0.001611 a445 b455 b27.2<0.00118.8<0.0010.1810.877
ME8.47 a5.91 c6.32 b0.398<0.0018.60 a5.91 c6.38 b0.416<0.0010.279<0.0010.3020.725
Erica arborea
DM571 b650 a500 c21.8<0.001551 b634 a475 c23.1<0.00115.6<0.0010.0010.711
OM9479559654.690.3319609679774.150.243.410.0830.0510.999
CP88.7 a53.7 c69.7 b5.380.00273.76577.32.660.1462.92<0.0010.6730.009
CT1081071192.780.14310091.71083.210.1112.480.0190.0060.674
EE96.3 a90.0 a46.6 b7.89<0.00199.6 a93.6 a57.0 b6.77<0.0015.09<0.0010.0120.294
NDF439 c531 b578 a21<0.001428 c544 b586 a24<0.00115.4<0.0010.6110.323
ADF341 c399 b445 a15.6<0.001328 c414 b458 a19.5<0.00112.1<0.0010.4480.178
ADL217 b307 a311 a15.4<0.001207 b320 a324 a19.3<0.00112<0.0010.0420.002
IVOMD479 a343 c407 b19.9<0.001486 a348 c417 b20.3<0.00113.8<0.0010.2570.948
ME6.62 a4.58 c5.79 b0.299<0.0016.77 a4.41 c5.86 b0.347<0.0010.222<0.0010.8910.282
Lavandula stoechas
DM299 c475 a409 b25.7<0.001281 c459 a385 b25.8<0.00117.8<0.001<0.0010.491
OM9449399166.240.1339539549306.250.2174.560.0310.1160.936
CP106 a83.0 b72.7 b5.09<0.001224 a96.7 b80.0 b2.810.0142.83<0.0010.374<0.001
CT3.073.572.570.2220.1912.572.932.20.2370.5130.1690.1180.1360.941
EE90.0 a34.3 b33.0 b9.41<0.00196.6 a38.6 b42.3 b9.38<0.0016.5<0.001<0.0010.112
NDF418 c472 a446 b8.240.001409 c486 a455 b11.5<0.0016.87<0.0010.3410.11
ADF253 b313 a298 a9.42<0.001238 b326 a313 a13.9<0.0018.16<0.0010.3120.03
ADL173 b208 a211 a6.23<0.001160 b215 a224 a10.1<0.0015.76<0.0010.2640.002
IVOMD698 a476 c512 b34.5<0.001704 a484 c522 b34<0.00123.5<0.0010.1080.936
ME10.2 a6.69 c7.29 b0.54<0.00110.3 a6.90 c7.40 b0.539<0.0010.371<0.0010.0250.829
Myrtus communis
DM554 a531 b437 c18.1<0.001533 a515 a420 b17.7<0.00112.4<0.001<0.0010.851
OM9529399483.790.4289629539593.310.5712.810.2450.0450.959
CP89.7 a72.3 b82.7 ab3.030.0375.7 b84.7 ab90.7 a2.570.0221.940.0610.4150.002
CT96.0 b128 a115 a4.880.00288.0 b116 a110 a4.470.0023.37<0.0010.0080.562
EE42.6 a41.0 a24.0 b3.09<0.00146.3 a48.0 a27.6 b3.37<0.0012.29<0.0010.0050.543
NDF379 a362 ab336 b7.370.0273693723466.050.174.640.0060.6420.462
ADF2182282424.650.067205 b242 a252 a7.760.0034.41<0.0010.4750.087
ADL1029394.32.010.16993.7 b108 a106 a2.550.0151.740.5710.0190.003
IVOMD493 b550 a485 b10.6<0.001500 b556 a504 b9.21<0.0016.94<0.0010.0240.374
ME7.24 b8.24 a7.16 b0.178<0.0017.41 b8.02 a7.28 b0.12<0.0010.104<0.0010.6730.047
Phillyrea media
DM523 c612 a570 b13.1<0.001507 c598 a555 b13.5<0.0019.31<0.0010.0040.974
OM9609709622.740.3389749789752.520.782.30.30.010.843
CP109 a86.3 b81.0 b4.4<0.0019697.3921.480.3512.28<0.0010.124<0.001
CT2.732.673.170.280.7842.42.22.770.2410.6850.1860.5530.3440.991
EE23.625.6281.220.40128.628.631.61.470.6921.040.3370.0760.918
NDF399 b435 a424 ab6.790.047387 b448 a429 a9.980.0075.860.0030.7650.379
ADF272 a259 ab250 b4.070.043258 b272 a255 b2.920.0052.440.0060.6510.011
ADL171 a124 b119 b8.42<0.001159 a136 b134 b4.560.0084.69<0.0010.1260.007
IVOMD515 a429 b413 b16.90.002523 a435 b425 b16.70.00311.6<0.0010.3960.967
ME7.29 a5.96 b5.82 b0.2470.0017.46 a5.76 b5.90 b0.285<0.0010.183<0.0010.8890.504
Pistacia lentiscus
DM547 c622 a590 b11.1<0.001530 c607 a578 b11.5<0.0017.98<0.0010.0060.882
OM959 a927 b954 a5.250.001973 a941 b964 a5.210.0053.91<0.0010.0010.79
CP93.0 b91.7 b106 a2.410.00378.0 b105 a113 a5.45<0.0012.9<0.0010.32<0.001
CT1751911853.640.187161 b177 a172 ab2.850.0332.790.0120.0030.997
EE27.3 a23.6 b23.3 b0.7410.016343027.31.520.2141.070.0270.0020.716
NDF448 b483 a422 c8.98<0.001437 b493 a426 b10.4<0.0016.68<0.0010.5910.012
ADF284 a248 b270 ab5.950.0142682632844.330.1093.610.0060.3630.037
ADL118 b165 a168 a8.50.002109 b178 a186 a12.57<0.0017.42<0.0010.1320.069
IVOMD505 a443 c471 b9.28<0.001508 a453 c483 b8.18<0.0016.09<0.0010.0220.453
ME7.216.66.740.1210.0697.37 a6.42 b6.83 ab0.1590.0180.0970.0010.8760.567
Rubus ulmifolius
DM371 b409 a410 a6.69<0.001356 b394 a406 a8.430.0085.42<0.0010.0330.601
OM908 b924 ab939 a5.260.025922 b936 ab947 a4.330.0283.590.0010.020.862
CP12511913940.101110 b132 ab152 a6.610.0043.770.0010.4040.036
CT1361381193.770.0521161211092.310.1222.870.007<0.0010.419
EE18.61821.30.9270.3412421.6271.350.3060.9910.1120.0090.856
NDF3653693614.780.8343523803726.480.2113.920.3030.730.381
ADF1992082013.070.485186 b221 a207 a5.590.0053.110.0030.5480.065
ADL75.370.362.32.750.14567.3 b83.0 a76.7 ab2.80.0392.050.1660.0550.017
IVOMD405 b443 a444 a7.190.008413 b452 a457 a7.750.0125.27<0.0010.0950.936
ME5.55 b6.62 a6.24 ab0.1770.0015.71 b6.46 a6.43 a0.1470.0270.1120.0010.6370.511
DM, dry matter, OM, organic matter; CP, crude protein; CT, condensed tannins; EE, ether extract; NDF, neutral detergent fiber; ADF, acid detergent fiber; ADL, lignin; IVOMD, in vitro organic matter digestibility; ME, metabolizable energy; S, season; Y, year; SEM, standard error of the means. Within a row, values with different letters are significantly different (p < 0.05).
Table 2. Chemical composition (g/kg DM), IVOMD (g/kg), and ME (MJ/kg DM) of trees species (n = 4) and herbaceous browsed by goats in Southern Mediterranean forest rangeland of Northern Morocco during two contrasting years.
Table 2. Chemical composition (g/kg DM), IVOMD (g/kg), and ME (MJ/kg DM) of trees species (n = 4) and herbaceous browsed by goats in Southern Mediterranean forest rangeland of Northern Morocco during two contrasting years.
Item2016 (Dry Year)2017 (Wet Year)SEMp-Value (2016–2017)
SpringSummerAutumnSEMp-ValueSpringSummerAutumnSEMp-ValueSYY × S
Olea europaea
DM461 a437 b426 b5.570.003444 a423 b414 b4.760.0023.95<0.001<0.0010.801
OM954 a912 b907 b8.10.004970 a920 b918 b8.670.0015.94<0.0010.0170.753
CP76.379.3822.860.77366.7 b94.0 a89.3 a4.920.0192.810.0240.3590.101
CT4.20 a2.17 b3.20 ab0.320.0043.87 a1.77 b2.63 ab0.3270.0020.228<0.0010.0510.888
EE94.0 b123 a79.6 c6.39<0.00199.0 b131 a85.0 c7.02<0.0014.67<0.0010.0050.722
NDF415 b449 a442 a5.720.005404 b459 a450 a8.8<0.0015.1<0.0010.5120.065
ADF314 a258 b265 b9.740.0083022792765.290.0675.44<0.0010.3060.14
ADL1591511612.510.282147 b164 a174 a4.410.0082.530.0110.1780.012
IVOMD4995175174.280.105505 b528 a530 a4.840.0433.350.0050.0620.886
ME6.85 c8.12 a7.59 b0.193<0.0016.96 b7.97 a7.75 a0.161<0.0010.122<0.0010.6490.305
Quercus canariensis
DM564 c690 a634 b18.3<0.001548 c678 a620 b18.8<.000112.8<0.0010.0010.859
OM9399619634.910.0549509679744.380.0543.410.0030.0580.871
CP104 a63.7 c72.3 b6.23<0.00190.7 a77.3 b79.0 b2.220.0013.22<0.0010.095<0.001
CT2026.717.32.150.1941416.312.71.090.4381.450.0810.0080.554
EE18.2 b24.6 a24.0 a1.180.01521.727.3281.30.0750.9470.0020.0140.897
NDF488 c550 a525 b9.25<0.001480 c560 a535 b11.9<0.0017.33<0.0010.1570.036
ADF322 c372 b394 a10.7<0.001317 c382 b404 a13<0.0018.19<0.0010.010.003
ADL114 c157 b176 a9.44<0.001103 c168 b189 a13.1<0.0017.85<0.0010.2550.025
IVOMD602 a406 c446 b30<0.001607 a414 c454 b29.3<0.00120.4<0.0010.0070.694
ME8.69 a6.00 b6.30 b0.427<0.0018.82 a5.81 c6.42 b0.462<0.0010.305<0.0010.7750.159
Quercus ilex
DM571 b612 a601 a6.25<0.001551 b596 a588 a7.14<0.0015.02<0.001<0.0010.509
OM9439539573.340.2519559609683.360.3222.590.0860.0480.926
CP114 a70.3 b71.3 b7.60.00199.7 a83.0 ab78.0 b4.030.0394.18<0.0010.6870.029
CT26.3 b60.0 a55.0 a5.31<0.00122.7 b48.3 a45.7 a4.17<0.0013.42<0.001<0.0010.097
EE17.719.1190.830.80319.622.623.61.10.350.7860.3020.0340.736
NDF568 a534 b506 b9.820.004553 a539 ab512 b7.580.0456.02<0.0010.8890.399
ADF3523223335.770.0713423343453.910.5593.440.0630.4160.256
ADL1701631713.180.582162 b175 ab192 a5.160.0243.130.0390.0830.071
IVOMD508 a410 c459 b14.4<0.001513 a424 c468 b13.1<0.0019.49<0.0010.0490.673
ME7.16 a5.99 c6.59 b0.172<0.0017.38 a5.87 c6.70 b0.223<0.0010.137<0.0010.3170.146
Quercus suber
DM587 b650 a604 b9.7<0.001573 b639 a589 b10.2<0.0017.03<0.0010.0040.932
OM968 a957 ab947 b3.80.049789699633.50.2122.940.010.0090.829
CP8578.388.32.360.22975.391953.830.0572.220.0560.3740.056
CT1191321242.930.1961101181162.70.482.290.110.020.796
EE252627.31.120.75228.629.330.31.090.860.8610.6530.0810.988
NDF579 a502 b485 b14.7<0.001565 a511 b490 b11.50.0049.07<0.0010.9820.164
ADF377 a348 b311 c10.10.001367 a367 a321 b8.220.0036.36<0.0010.2230.097
ADL168 a134 b133 b6.120.0021611461492.980.0693.41<0.0010.0610.036
IVOMD543 a406 c506 b20.4<0.001550 a421 c513 b19.2<0.00113.6<0.0010.0030.412
ME8.05 a6.00 c7.34 b0.296<0.0018.19 a5.84 c7.51 b0.353<0.0010.223<0.0010.6050.085
Herbaceous
DM463 c631 a516 b25<0.001446 c616 a495 b25.6<0.00117.5<0.0010.0190.923
OM916 a870 b855 b9.80.002931 a883 b872 b9.50.0016.87<0.0010.0130.958
CP156 a78.3 b65.7 b14.2<0.001142 a91.0 b76.7 b10.1<0.0018.45<0.0010.2910.006
CT2.42 b4.17 a2.97 b0.3230.0482.23.532.430.2870.1160.2170.0060.1760.865
EE19.92223.30.8230.26222.626.326.31.120.3490.7890.110.0270.872
NDF51756849715.40.14750858050716.20.08710.80.0140.8270.865
ADF339 b363 a269 c14.70.001326 b379 a283 c14.60.00110.1<0.0010.4260.216
ADL71.475.764.32.760.26567.0 b89.0 a79.0 ab3.950.0432.530.0310.0560.106
IVOMD804 a651 c705 b22.7<0.001807 a658 c719 b22<0.00115.3<0.0010.2230.771
ME12.0 a9.25 c10.7 b0.405<0.00112.1 a9.57 c10.8 b0.364<0.0010.265<0.0010.0970.334
DM, dry matter, OM, organic matter; CP, crude protein; CT, condensed tannins; EE, ether extract; NDF, neutral detergent fiber; ADF, acid detergent fiber; ADL, lignin; IVOMD, in vitro organic matter digestibility; ME, metabolizable energy; S, season; Y, year; SEM, standard error of the means. Within a row, values with different letters are significantly different (p < 0.05).
Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Share and Cite

MDPI and ACS Style

Chebli, Y.; El Otmani, S.; Chentouf, M.; Hornick, J.-L.; Cabaraux, J.-F. Temporal Variations in Chemical Composition, In Vitro Digestibility, and Metabolizable Energy of Plant Species Browsed by Goats in Southern Mediterranean Forest Rangeland. Animals 2021, 11, 1441. https://doi.org/10.3390/ani11051441

AMA Style

Chebli Y, El Otmani S, Chentouf M, Hornick J-L, Cabaraux J-F. Temporal Variations in Chemical Composition, In Vitro Digestibility, and Metabolizable Energy of Plant Species Browsed by Goats in Southern Mediterranean Forest Rangeland. Animals. 2021; 11(5):1441. https://doi.org/10.3390/ani11051441

Chicago/Turabian Style

Chebli, Youssef, Samira El Otmani, Mouad Chentouf, Jean-Luc Hornick, and Jean-François Cabaraux. 2021. "Temporal Variations in Chemical Composition, In Vitro Digestibility, and Metabolizable Energy of Plant Species Browsed by Goats in Southern Mediterranean Forest Rangeland" Animals 11, no. 5: 1441. https://doi.org/10.3390/ani11051441

APA Style

Chebli, Y., El Otmani, S., Chentouf, M., Hornick, J. -L., & Cabaraux, J. -F. (2021). Temporal Variations in Chemical Composition, In Vitro Digestibility, and Metabolizable Energy of Plant Species Browsed by Goats in Southern Mediterranean Forest Rangeland. Animals, 11(5), 1441. https://doi.org/10.3390/ani11051441

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop