Shrubby and arboreal species diet preferences of creole goat in extensive husbandry during dry season at different altitude levels of dry forest in southern Ecuador

E L Aguirre-Riofrio^1*, V T Ullaguari-Ramirez¹, V Alvarado-Jaramillo², Z Aguirre-Mendoza² and H Viñan-Díaz¹

¹ Agricultural Faculty, Veterinary Medicine and Zootechnics School, National University of Loja, Loja, 110110, Ecuador
^* Correspondence to: edgar.aguirre@unl.edu.ec
² Herbarium “Reinaldo Espinosa”, National University of Loja

Abstract

The seasonal dry forest ecosystem occupies a third of the surface of the province of Loja in southern Ecuador. For most of the year, this environment has shortages of water and pastures, but the animal that has adapted and survived is the Chusca lojana goat. The objective of this study was to identify plant species that contribute natural feeding in dry season of these extensively husbandry goats at the different altitudinal levels of this ecosystem. Three herds of Chusca goats were chosen at each level, geographically separated by 5 km each. In the identification of the floristic composition at each level, 5 plots of 200 m² were used and in each of them 5 subplots of 8 m² were distributed, and the species within each subplot were counted. Regarding consumption preferences, these were carried out at each level and herd by direct observation with binoculars twice a day for 3 days. At the first altitudinal level the predominance of coverage in 86% is of 3 species: 2 shrubby Ipomoea carnea and Croton wagneri and one arboreal Handroanthus chrysanthus; at the second level, 4 species occupy 92.8%: the shrubby Croton wagneri and arboreals Ficus cuatrecasana, Acacia macracantha and Handroanthus chrysanthus; with 97% of coverage the third level is occupied by 2 shrubby Croton wagneri and Lantana cámara and 2 arboreals Acacia macracantha and Ceiba trischistandra. During this time, at all levels, the main food consumed by the goats is Croton wagneri, Handroanthus chrysanthus and Acacia macracantha. In conclusion, this ecosystem is characterized by the predominance, at all altitudinal levels, of shrubs where Croton wagneri and Ipomoea carnea abound, which have limitations in their consumption, but this goat has overcome them to survive. By processing the forage biomass of these plants and supplementing the diet in dry season, this would allow improvement in the productivity of this animal.

Keywords: browsing, borrachera, dry forest, floristic composition, moshquera, natural feeding

Introduction

In the Loja province located in the south of Ecuador (4°03′S 79°39′W), there is the seasonal dry forest ecosystem, which is characterized by extreme climatic conditions, with an annual rainfall of 400-600 mm, in a period of 3-4 months, generally from February to April, and with an average annual temperature of 24.9^ºC. The largest area of this ecosystem is in the range 0-1000 m.a.s.l, which includes the lowlands, low western foothills of the andes mountain range, located on hilly and abrupt terrains. The soils on which the dry forests develop are clayey and occasionally stony and s andy, forming quagmires in the rainy season and large cracks in the dry season (HerbarioLOJA 2001).

According to Aguirre (2012), dry forests are fragile and pressured ecosystems, where the human population lives and develops its productive activities. Standing out in this is the traditional use of the forest in the open-field grazing of goats, since it is this animal (due to its rusticity, resistance, and easy handling) that is best adapted to this environment, constituting an important source of economic income and livelihood for the inhabitants of this area.

The feeding behavior of goats is very selective (and opportunistic unlike other ruminants) when the vegetation is abundant, and very general when there is any shortage (Sánchez-Rodríguez et al 1993). These animals within the dry forest are surrounded by a great variety of vegetation, where trees and shrubs play an important role in the availability of food, especially in the dry season, and where browsing is the main source of food in arid and semi-arid regions of the world (Foroughbakhch et al 2013). Therefore, it is important to recognize both native and introduced plant species that the goat uses in its diet, in order to improve the conservation and sustainable management of this ecosystem and consequently the productivity of these animals within the extensive management that is carried out.

In the open field in times of food scarcity, the goat assumes a browsing feeding behavior, where the shrubs and trees present in the ecosystem of the dry forest constitute its main source of food. Considering that this ecosystem in the Province of Loja occupies different altitudinal levels, and therefore different types of vegetation in each one of them, the objective of the present investigation was to identify the plant species that contribute to the natural diet of the goat, when managed extensively in the dry season at the different altitudinal levels of the seasonally dry forest ecosystem in this province.

Materials and methods

Location of the study area

The study was carried out at three altitudinal levels of the seasonal dry forest ecosystem of the province of Loja. To determine the location of the study sectors, a map was prepared using the free QGIS^® software that determines the characteristics of altitude and coordinates. First level (0 to 400 m.a.s.l) in Limones and Pampa Blanca sectors in the Zapotillo Canton; Second level (400 to 900 m.a.s.l) in Algodona and Laguar sectors in the Macará Canton and Third level (900 to 1200 m.a.s.l.) in El Limon and Zapotepamba sectors in the Paltas Canton (Map 1). These study sites were selected due to the presence of Chusca goat herds that are managed extensively and the study was realized in the dry season of september to december 2022, with temperature between 28.5ºC in first level, 26ºC second level to 20.5ºC in the third level and an average daily sunhours of 8.3 and 40 mm of monthly rainfall for this time.

Characterization of the goat from the seasonally dry forest of southern Ecuador

The “Chusca lojana” goat, after the genetic characterization carried out by Aguirre et al (2020), it is considered a rustic local creole breed perfectly adapted to the arid and semi-arid environment of the dry forest, is an animal with a polychromatic coat (brown, black and white), ellipometric, long-lined and fleshless with a tendency more towards milk than meat and marked sexual dimorphism in terms of size, fronto-nasal profile, presence of beard in the males (Aguirre et al 2021).

Map 1. Sectors of the seasonal dry forest ecosystem in the province of Loja-Ecuador, where the research was carried out

Observational units

In this study, a herd of 60 adults Chuscas goats was considered an observational unit, and the herd selection criteria were: 1) more than 70 % of the goats belonged to this racial group, 2) herds were managed in an extensive environment, and 3) they were geographically separated at a distance of 5 km. Three goat herds were considered for each study zone or altitudinal level.

Recognition of the floristic and structural composition of the species in the study area

The zones were delimited by means of GPS and maps created in QGIS^®. For recognition of the species, five permanent plots of 10 x 20 m (200 m²) were used on each floor, delimited with wooden stakes and five subplots of 2x4 m (8 m²) were distributed in each one of them, and where the species found within each subplot were counted

Taxonomic classification of species

For the identification and classification of the botanical species found in the plots, samples were collected and taken for processing to the “Reinaldo Espinoza” Herbarium of the National University of Loja; The Web pages The Plant List and The World Flora Online, which are specialized scientific Web sites, were also used.

Preferences of the plant species for goats

Following a methodology used by Altmann (1974), Franco et al (2005) and Foroughbakhch et al (2013), the observation of which plant species are consumed by goats at each altitudinal level was carried out in each herd over a period of 3 days, twice a day, using sighting binoculars and choosing 30 goats at random according to the availability of observation, making the first observation from leaving the pen until 12:00 (noon) and the second from 2:00 p.m. to 5:00 p.m. when they returned to their pens, the animals did not receive any special handling or nutritional supplementation during this time.

The information on the type of vegetation that the animal consumed, the name of the plant, and the part of it that was consumed (leaf, flower, fruit, stem, bud), were recorded on a field record prepared for this purpose. The preference regarding the consumption of the plants was measured based on the number of animals that consumed them and was categorized as follows: high level, when the number of goats that consumed a certain plant exceeded 10 animals; medium level: when the number of animals was in a range between 6 and 9; and low level: when the animals that consumed a certain plant did not exceed 5 in number.

Ordering and tabulation of information

The collected field information was ordered by altitudinal floors based on the inventory of existing plants and the dietary preferences of the goats in each of these study ecosystems.

Results

Floristic and structural composition of the species in the study area

Table 1 shows the 22 floristic species found in the dry season at the 3 altitudinal levels of the seasonally dry forest and consumed by the population of Chusca goats.

At the first altitudinal level of the dry forest, the presence of 10 plant species between arboreal and shrubby was recorded, where Ipomoea carnea predominates (known as borrachera) with 46.6%; followed by Croton wagneri (known as Moshquera) with 26.6%. These species, being shrubby, cause an abundant density in this ecosystem that makes movement difficult. The tree species Handroanthus billbergii (guayacán), with 12.8%, Caesalpinia paipai (charan) and Piscidia carthagenensi (barbasco) with 4.6% and 4.1%, respectively, continue to be prevalent at this level, and these 5 plant species occupy 94.6% of the floristic composition of this floor. With a low percentage of prevalence (5.4%), 50% of the remaining species are found, such as Cordia lutea (overal), Coccoloba ruiziana (añalque), Geoffroea spinosa (almond tree), Neltuma juliflora (carob tree) and Loxopterygium huasango (hualtaco).

At the second altitudinal level, the shrubby presence of a large amount of Croton wagneri is observed in 64.3%, followed in order of prevalence by arboreal species such as Ficus cuatrecasana (higueron) in 11%, Acacia macracantha (faique) with 9.5%, and Handroanthus billbergii with 8.0%, showing the latter as a deciduous tree and with its leaves covering the ground surface, and these four species cover 92.8% of the surface at this altitudinal level. In addition to this, another eight species were found in this area that together occupied 7.2% of the floristic composition, such as Neltuma juliflora, Pithecellobium excelsum (chaquiro), Ziziphus thyrsiflora (ebano), Guazuma lmifolia (guazimo), Pisonia aculeata (pego pego), Celtis iguanaea (uva de pava), Caesalpinia paipai and Bougainvillea peruviana (papelillo).

In the floristic composition at the third level, more than 50% is made up of the Croton wagneri shrub (55.9%) and the presence of a large number of trees and small shoots of Acacia macracantha (26.3%), followed by the Lantana camara shrub with 10.9%, another tree species identified is Ceiba trischistandra (ceibo) with 3.9%. It is common to observe these large trees at this level, and these four plant species constitute 97% of the floristic composition of this ecosystem. Other plants found in a lower percentage were the shrub Bauhinia aculeata (pata de vaca) and the Tecoma stans (lame) and Coccoloba ruiziana (añalque) trees with 1.3%, 1.0%, and 0.7%, respectively.

Feeding preferences of goats towards plant species present in dry season

It is highlighting the varied topography of this ecosystem that influences the daily movement of the goat, so in the first and second level the terrain is semiplane in these places the goats move around 3 km daily while in third level through irregular terrain the move is around 2 km daily in search of food, which not finding it at ground level, and it is the arboreal and shrubby species provide what they seek, thus they developed a browsing behavior during this season. Table 2 shows the dietary preferences of goats at the different altitudinal levels of the seasonally dry forest ecosystem in the Province of Loja.

At all the altitudinal levels of the dry forest (Table 3), it was observed that the goat has a high preference in the consumption of Croton wagneri  and this probably due to its great distribution in this ecosystem and the ease of consumption, since they are shrubs and form numerous shoots on the ground. Likewise, another appreciated species that can be found up to 900 m.a.s.l. is Handroanthus billbergii, which in the dry season drops its leaves on the surface of the soil, and in this way they are easy to consume, just like the flowers during their flowering that occurs at the end of the dry season with the beginning of the first rains. The goat has a high preference for Acacia macracantha due to its availability from 400 m.a.s.l., and there buds, leaves and fruits are eagerly consumed by the goat

Table 3. Main plants consumed by the Chusca goat during the dry season at the different altitudinal levels of the dry forest of the Province of Loja

It is noteworthy that at each of the altitudinal levels studied there are specificities, so at the first altitudinal level, the goat has a high consumption preference for Caesalpinia paipai, consuming the leaves, flowers, and fruits. With a medium level of consumption, there are three species, which the Chusca goat, due to its adaptation to this environment, only consumes what is strictly necessary so as not to cause harm, since high consumption causes problems in the animal. These are: Cordia lutea, Ipomoea carnea, consumed by goats in small quantities, especially the dry leaves and buds, and Piscidia carthagenensis. Finally, with a low level of consumption are the leaves, flowers, and fruits of Geoffroea spinosa and Coccoloba ruiziana, highlighting that the low consumption of these plants is due to their low number within this ecosystem.

At the second altitudinal level, there are the trees Neltuma juliflora and Ficus cuatrecasana, for which the goat has a high preference, because both their leaves and fruits are arranged hanging from their branches or they fall to the ground. For species such as Pithecellobium excelsum and Caesalpinia paipai, their flowers, fruits, and leaves that fall to the ground are consumed with a medium level of preference, because these species are not very abundant in this ecosystem. For other species known in the environment, such as Pisonia aculeata, Bougainvillea peruviana and Celtis iguanaea, they are in sparse numbers at this level, consequently their level of consumption is low, as the goat takes advantage of everything available at that moment (leaves, flowers, or fruits).

At the third level, there is a low consumption of certain species such as Ceiba trischistandra, whose use is limited because it is a high-altitude arboreal species. However, when its fruits and leaves have fallen they are consumed, as well as its buds. Further, the shrubs Bauhinia aculeata and Lantana camara, as well as the arboreal Tecoma stans, for its green and dry leaves and flowers, are eaten by the goat.

Discussion

Floristic composition of the dry forest of the province of Loja

The floristic composition in the seasonally dry forest ecosystem in the province of Loja, which is located within the so-called “Tumbesino Endemism Region”, which covers the northwest zone of Peru and southwestern Ecuador, varies according to the altitudinal levels, which is in agreement with what was stated by Cueva et al. (2019), in that as the altitude increases, biodiversity increases. This is also corroborated in the work carried out by Lozano (2002); Leal and Linares (2005); Aguirre and Delgado (2005); Aguirre et al (2006, 2014 and Aguirre Z. (2012).

At the first altitudinal level, the predominance was of the shrubby species Ipomoea carnea, followed by Croton wagneri, and this regard Lozano (2002) and Aguirre et al (2006, 2014), in their studies carried out on the types of forests in southern Ecuador, state that the vegetation found in the lowest part of Zapotillo in a range between 280 and 350 m.a.s.l. is xerophytic, thorny, and stunted with the presence of columnar cacti, legumes, and species with latex of the genus Croton and Ipomoea. These details agree with the present study, where these two shrubs represent 73.2% of the species at this altitudinal level. The Handroanthus billbergii, Cordia lutea, and Piscidia carthagenensis trees were also found to a lesser degree, coinciding with those found by Aguirre and Delgado (2005) in the Cerro Negro-Cazaderos sector, Zapotillo. Caesalpinia glabrata, Netulma juliflora and Loxopterygium huasango are also found at this level, which are trees that have been recorded by Leal and Linares (2005) in the dry forests of northwestern Peru. A Fabaceae such as Geoffroea spinosa and a Polygonaceae Coccoloba ruiziana were also found at these sites, which are characterized by the fact that they do not completely lose their leaves during the dry season (Aguirre and Delgado 2005), constituting permanent sources of food for the goat.

At the second altitudinal level, Croton wagneri and the tree species Ficus cuatrecasana, Acacia macracantha and Handroantus billbergii predominate. This floristic composition coincides with the species described by Lozano (2002), who states that between 400 and 900 m.a.s.l, the vegetation supports long periods of low water (8 months), and it is at the beginning of the rains where the physiognomy changes and various legumes flourish. Other trees such as Neltuma juliflora, Pisonia aculeata, Celtis iguanacea, Zizipus thyrsiflora, Guazuma ulmifolia and Caesalpinia paipai are also found at this level in low numbers, as are the shrubby Pithecellobium excelsum and Boungainvillea peruviana. These same species were described by Aguirre et al 2006, in their study of the vegetation of the deciduous and semi-deciduous forests of the Tumbesina region. All of this inventory of plants found at the second altitudinal level, agrees with what was stated by Aguirre Z (2012), in his description of the plant species of the pluviseasonal dry forest.

According to what was stated by Lozano (2002), above 1000 m.a.s.l, the plant formation is montane dry scrub (ms-M) and lower montane semideciduous forest (bsd-MB), where the prevailing vegetation is low scrub of no more than 3 meters high. Being a mixture of plants armed with thorns with other species that have latex such as Croton, this description agrees with the findings of the present research, where four species predominated at this level, two shrubby Croton wagneri and Lantana camara, the latter having thorns, the arboreal Ceiba trichist andra, and Acacia macracantha, it is noteworthy that this floor is characterized by a rather irregular geography with slopes that reach 60%. Likewise, it is necessary to consider the low presence of Ceiba trichist andra (3,9%) at this level, and possibly this is due to what was stated by Aguirre et al (2015), that this typical dry forest tree has little regeneration, and this is due to the fact that goats and cattle that graze in the forest consume its regrowth and seedlings, thus preventing its growth. Finally, to corroborate the findings of this study, Sánchez et al (2006) also mention these two types of trees as the predominant at these altitudes.

Food preferences of the Chusca goat in the dry forest of the province of Loja

The goat, due to the extensive management that it receives and in ecosystems with little food availability, has become an opportunistic mixed-feeding herbivore that can diversify its diet according to the availability of the environment (Genin & Pijoan 1993; Perevolotsky et al 1998; Franco et al 2005). Likewise, adult goats have more facility, skill, and scope in selecting species than young goats. This experience is acquired with age and knowledge of the space that they cover daily, thereby improving feeding capacity and thus surviving in a given environment. (Launchbaugh and Howery 2005). In this environment, this has caused capriculturists to employ a management practice of keeping the kids in the pens to prevent them from being preyed on, and to help them with feeding until they grow up and engage in their own search for food.

According to Bailey et al (2001), Odo et al (2001) and Mellado et al (2004a), feeding habits, consumption preferences, and the botanical composition of the diet are influenced by the genetic factor, since breeds developed in a certain region, as is the case of the Chusca lojana in the dry forest of the province of Loja, have superior skills in the use of the aforementioned environment (they are more browsing) compared to other external racial groups. So, also populations of goats that feed on less preferred but abundant plants have a greater impact on long-term survival and reproductive success than other goats. In this same context, Franco et al (2005) state that goats raised in a free grazing transhumance system show a greater preference for the consumption of bushes and fallows, and in summer and fall the bushes contribute 69% of the plants selected for food, reducing this consumption by 34% in the rainy season. These Feeding strategies also help biological control of endoparasites by consuming plants that have higher tannin levels (Min et al 2003) and all this allows us to underst and and explain the survival of the Chusca goat in this environment.

The palatability or preference for forages is a multifactorial phenomenon, given not only by the presence of secondary metabolites with toxic potential, which are detected by the animals, but also by the ethological characteristics, animal category, biomass availability, season of year, arrangement of the branches, presence of physical defenses, roughness and pubescence of the leaves (Arnold 1981; Aerts et al 1999). It is important to consider what was stated by García (2004), that ruminants, according to the degree of adaptation to a certain environment, are provided with physiological defenses at the level of saliva and specialized ruminal bacteria as mechanisms to counteract the presence of metabolites such as phenols, cyanogenic, tannins, saponins, and alkaloids present in certain plants, such as Croton and Ipomea found in the present study.

Regarding Croton wagneri, it must be mentioned what was stated by Foroughbakhch et al (2013), that the least preferred plants by the goat in the semi-arid region of northeastern Mexico in the rainy season are Croton spp, among others, but that in the most critical seasons of the year they become part of the diet before the scarcity of fodder. A similar attitude occurs in the dry forest of Loja, where this plant is the most abundant shrub and is found in all altitudinal levels of this ecosystem, with the Chusca goat having a high consumption preference in the dry season, an ethological attitude that is explained by Franco et al (2005), stating that there is a high correlation (r² = 0.703) between the presence of the plant and its consumption. Teran et al (2020) in this regard indicate that this plant is endemic to Ecuador and grows at between 1000 and 3012 m.a.s.l. In the present study it is found even at below 400 m.a.s.l. Its leaves have a bitter and astringent taste with a high protein content (15.63±0.04%) and fiber (19.53%), fat (4.92%), ash (10.5%), total carbohydrates (44.59%), Omega 3 and 6 fatty acids (1.14% and 0.66 %, respectively). According to Cevallos et al (2016) and Teran-Portelles et al (2019), the plant is not toxic and can be used since no changes are observed in the health of the animals, and they are tolerant to the extracts of its different parts.

Regarding Ipomea carnea, it is a shrub that is found at the first altitudinal level of this ecosystem, and has a medium consumption preference for the goat in the dry season. According to Armien et al (2007), Sahayaraj and Ravi (2008), and Hussain et al (2022), it is a good source of flavonoids, terpenes, tannins, saponins, phenolic compounds, and alkaloids such as Swainsonine and Calistegine, which cause dysfunctions in the central nervous system, the former being more toxic to animals (Ikeda et al 2003; Armien et al 2007). According to those authors the animals do not suffer and they overcome the intoxication if the consumption of the plant is discontinuous. In this context, Rios et al (2008), state that when daily administering an amount of 50 g/kg LW of leaves, stems and flowers, the animals presented the first symptoms only 39 days after starting their consumption. An interesting fact regarding this plant is that the concentration of Swainsonin and Calistegin in the fresh leaves is 0.003% and 0.005%, respectively, while in the seeds (fruits) these concentrations are 10 times greater (Haraguchi et al 2003).

In this ecosystem and during the dry period in all the altitudinal floors studied, between 67 and 75% of the predominant vegetation cover is shrubby plants and the remaining percentage is arboreal, there being no herbaceous plants at this time (Table 1), therefore the diet of the Chusca goat, like the goat from the northeast of Mexico (Foroughbakhch et al 2013) and those of Oaxaca Mexico (Franco et al 2005), is based on the regrowth, leaves, and fruits of shrubby and arboreal plants that grow in such places. In this sense, Mellado et al (2004b) state that in the dry season the consumption of shrubby vegetation is similar for both adult and juvenile goats. Likewise, the body condition affects this consumption, since lower condition animals such as Chuscas goats consume more shrubs and are less selective than goats with a better condition.

Conclusions

The seasonal dry forest ecosystem of the Province of Loja is characterized by a predominance of shrubby and arboreal vegetation throughout its entire length, where Croton wagneri and Ipomoea carnea are abundant and have limitations in their consumption, due to the presence of certain metabolites that they contain. Hence, the breed of Chusca lojana goats that inhabits this environment, and that in the dry season becomes a browser, uses them to survive, which is why taking advantage of its nutritional ethology, to process the forage biomass of these plants in the rainy season and supplement the diet of the goats in dry season, would allow to improve the productivity of this population.

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