Concepts of Spiritual Nourishment in the Andes And Europe: Rosaries in Cross-Cultural Contexts

This article explores visual imagery connected with the Christian rosary and its connotations of spiritual nourishment in European art and Andean ritual practices. Early eighteenth-century religious art inspired by a vision received by Isidoro of Seville, a Capuchin friar, presented the Virgin Mary as the Divine Shepherdess with her flock of Christians, depicted as sheep with roses in their mouths. This imagery resonates in the flowering ceremony practised by Aymara pastoralists in Isluga, Chile. The floreo serves as a rite of passage for herd animals, its intention being to enhance their fertility by making them 'flower'. However the congruence between the visual codes produced by Europeans and Andeans conveys related but different social and moral messages. Whereas sheep can represent human beings in Christian art, they cannot stand in for human beings in Isluga, since Isluga people recognize that both human and herd animals require spiritual nourishment.     

Herders' Perceptions, Practice, and Problems of Night Grazing in the Sahel: Case Studies from Niger

A survey was conducted from February to June of 1997 among livestock herders in two villages of Niger, Kodey and Toukounous, on their perceptions, practice, and problems of night grazing. Cattle and sheep were the species that were taken out for night grazing by the herders. Small herd size and labor constraints were mentioned as the principal reasons for not practicing night grazing. Major benefits of night grazing included good body condition, herd growth, increased milk production, prevention of diseases, and reduction in herd mortality. Insecurity, difficulty in staying awake at night, labor constraints, and damage to crops by animals were given as problems of night grazing. According to the herders, grazing time (duration) during the day and night was shorter in the wet season than in the dry season. In the wet season animals were herded (followed and closely supervised by herders), whereas in the dry season, animals were mostly left to range freely in both villages. In general, children herded the animals during the day, while adults were responsible for night-time herding. Herders' perceptions on night grazing as regards animal production parameters such as weight development, water consumption, fecal output and feeding behavior are consistent with available experimental results. Therefore, future technical research needs to recognize the constraints faced by herders and determine how to overcome them to improve technical and economic efficiency.     

Livestock Acquisitions Dynamics in Nomadic Pastoralist Herd Demography: A Case Study Among Ngisonyoka Herders of South Turkana, Kenya

Despite the attention given to social relations in the pastoral literature, the role of livestock acquisitions—additions of livestock to herds through bridewealth, exchanges, gifts, payments, and begging (requests)—in herd build up has usually been assumed to be relatively minor compared to births and relevant mostly when the need for rebuilding arises after major losses. This study is based on an unusual set of data—the reproductive histories of the female cattle, camels, and goats and sheep of 13 Ngisonyoka Turkana nomadic herders in northwestern Kenya, collected in 1987. The article reports on the means by which mothers were added to the herd and how these changed through time. The results suggest that for this population in the late 1970s and 1980s, acquisitions were not merely relevant when disaster struck, but instead were a continuously important component of herd management. The results demonstrate the crucial role of social networks in the survival of Ngisonyoka pastoralists in their non-equilibrial ecosystem. Social exchanges, such as bridewealth, provide a resource security well suited to the challenges of coping with such unpredictable environments. Researchers and policymakers are urged to make efforts to support such indigenous networks if they want nomadic pastoralists to continue their effective use of arid, marginal lands.

Cattle and pastoralism: Survival and production in arid lands

Traditional subsistence pastoralists in East Africa tend to keep large herds, milk cattle in preference to eating them, and subject them to long foraging treks. Such practices are widely considered ill-suited to arid lands and are believed to arise because cattle are raised more for social prestige than food production. Whether this is true can only be judged by considering the responses of cattle to arid zones and, given the herder's goals and options, his management practices. In considering these factors, we show that indigenous East African cattle demonstrate energy-sparing capabilities during drought. Pastoralists can therefore herd cattle at great distances from water at little more cost than animals on the normal maintenance diet and watered more frequently. The physiological response of cattle to drought, the ecological constraints imposed by livestock and wildlife competition, and the energetic efficiency of mixed milk and meat pastoralism explain why herders traditionally select their characteristic management practices.     

Wealth-dependent and interdependent strategies in the Saami reindeer husbandry,Norway

It has been argued that decisions in relation to choosing strategies to a large degree depend on an organism's state. For nomadic pastoralists, wealth is an important state variable since it has been argued that differences in observed behaviours reflect alternative strategies dependent on varying socioeconomic circumstances. From a game theoretical point of view, however, strategies are also interdependent, i.e., the choice of a strategy cannot be made wisely without considering what other actors are doing since the outcome of a given strategy is dependent not only on individual state but also on the strategies of others. This study investigated to what degree slaughter strategies in the Saami reindeer husbandry are both state dependent and interdependent. The main findings in this study were that (a) the probability, (b) the amount, and (c) the type of animal slaughtered to a large degree were influenced by both individual herders’ herd size and the number of animals slaughtered by neighbouring herders. Moreover, this study also found that kinship represents a coordinating principle since the degree of genealogical relatedness had a positive effect on the slaughtering strategies adopted by herders.     

Population status of Andean Condors in central and southern Bolivia

Andean Condors (Vultur gryphus) are a Near Threatened species that was formerly distributed along the entire length of the Andes from western Venezuela to Tierra del Fuego. Populations have been severely reduced north of Peru, but several thousand Andean Condors still exist in the southern portion of their range in Argentina and Chile. Little is known, however, about the size of the Andean Condor population in the central part of their range in Peru and Bolivia. From June to September 2012, we used feeding stations to attract Andean Condors and estimate the size and structure of the population in the eastern Andes of central and southern Bolivia. We estimated a minimum population of 253 condors, an adult male‐to‐female ratio of 1:0.6, an immature male‐to‐female ratio of 1:0.9, and an adult‐to‐immature ratio of 1:1.1. At our five survey areas, estimated abundance ranged from 15 to 100 condors per area. Males outnumbered females in three areas and the opposite was true in two areas. Our estimated adult‐to‐immature ratio, overall and in each area, suggests that the populations could be reproducing at a high rate. As previously observed in other Andean Condor populations, skewed sex ratios could be associated with differences between sexes and age classes in habitat selection. Although our results suggest that Bolivian populations of Andean Condors are still reasonably large, population monitoring is urgently needed, including use of feeding stations throughout the entire Bolivian range of the species and intensive searches for roosting and nesting sites.     

Diversification patterns in the CES clade (Brassicaceae tribes Cremolobeae,Eudemeae, Schizopetaleae) in Andean South America

Dated molecular phylogenetic trees show that the Andean uplift had a major impact on South American biodiversity. For many Andean groups, accelerated diversification (radiation) has been documented. However, not all Andean lineages appear to have diversified following the model of rapid radiation, particularly in the central and southern Andes. Here, we investigated the diversification patterns for the largest South American‐endemic lineage of Brassicaceae, composed of tribes Cremolobeae, Eudemeae and Schizopetaleae (CES clade). Species of this group inhabit nearly all Andean biomes and adjacent areas including the Atacama–Sechura desert, the Chilean Matorral and the Patagonian Steppe. First, we studied diversification times and historical biogeography of the CES clade. Second, we analysed diversification rates through time, lineages and associated life forms. Results demonstrate that early diversification of the CES clade occurred in the early to mid‐Miocene (c. 12–19 Mya) and involved the central Andes, the southern Andes and the Patagonian Steppe, and the Atacama–Sechura desert. The Chilean Matorral and northern Andes were colonized subsequently in the early Pliocene (4–5 Mya). Diversification of the CES clade was recovered as a gradual process without any evidence for rate shifts or rapid radiation, in contrast to many other Andean groups analysed so far. Diversification time/rates and biogeographical patterns obtained for the CES clade are discussed and compared with patterns and conclusions reported for other Andean plant lineages.     

Nomadization in Rajasthan, India: Migration, Institutions, and Economy

Despite a global trend toward settlement, the incidence of pastoral nomadism is on the rise in the Marwar region of Rajasthan, India. Typical explanations for this change use models of population pressure; increasing herds and decreasing pasture are held to blame. This explanation, however intuitive, is unsatisfactory. Instead, changing institutional and economic patterns are creating new contexts for strategic movement. Bottlenecks in the yearly resource calendar, caused by the disintegration of obligatory social relationships, force migrations during periods of scarcity. Changes in the volume and pattern of the meat and wool markets have also created opportunities for migrating pastoralists. Producers increase their access to markets and the reproductive rate of their herd through long, annual, migration. While nomadism is a general adaptation to changes in the socioeconomic conditions of the region, differential resource endowments account for the range of strategies; wealthy herders have opportunities not enjoyed by more marginal producers.     

Evolutionary islands in the Andes: persistence and isolation explain high endemism in Andean dry tropical forests

Aim The tropical Andes are a world biodiversity hotspot. With diverse biomes and dramatic, geologically recent mountain uplift, they offer a system to study the relative contributions of geological and biome history to species richness. There are preliminary indications that historical species assembly in the Andes has been influenced by physiographical heterogeneity and that distinct biomes have evolved in relative isolation despite physical proximity. Here we test this ‘Andean biotic separation hypothesis’ by focusing on the low‐elevation, seasonally dry tropical forest (SDTF) biome to determine whether patterns of plant diversification within the SDTF differ from those in mid‐ and high‐elevation biomes. Location Tropical Andes, South America. Methods Densely sampled time‐calibrated phylogenies for five legume genera (Amicia, Coursetia, Cyathostegia, Mimosa and Poissonia) containing species endemic to the Andean SDTF biome were used to investigate divergence times and levels of geographical structure. Geographical structure was measured using isolation‐by‐distance methods. Meta‐analysis of time‐calibrated phylogenies of Andean plant groups was used to compare the pattern and tempo of endemic species diversification between the major Andean biomes. Results Long‐term persistence of SDTF in the Andes is suggested by old stem ages (5–27 Ma) of endemic genera/clades within genera, and deep divergences coupled with strong geographical structure among and within species. Comparison of species diversification patterns among different biomes shows that the relatively old, geographically confined pattern of species diversification in SDTF contrasts with the high‐elevation grasslands that show rapid and recent radiations driven by ecological opportunities. Main conclusions The SDTF biome has a long history in the Andes. We suggest that the diverse SDTF flora has been assembled gradually over the past c. 19 Ma from lineages exhibiting strong phylogenetic niche conservatism. These patterns suggest that Andean SDTFs have formed stable and strongly isolated ‘islands’ despite the upheavals of Andean uplift. Indeed, the Andean SDTFs may represent some of the most isolated and evolutionarily persistent continental plant communities, similar in many respects to floras of remote oceanic islands.     

Phytogeographical relations of the Andean dry valleys of Bolivia

Aim The objective of this study is to examine the phytogeographical affinities of the Andean dry valleys of Bolivia in order to contribute to a better understanding of the Andean dry flora's distribution, origin and diversity. Particular emphasis is given to the analysis of the floristic connections of this flora with more austral parts of South America. Location The dry valleys of Bolivia are located in the Andes of the southern half of the country, at elevations between 1300 and 3200 m. Methods An extensive floristic list compiled by the author to evaluate plant diversity in these Andean regions was used as the base for this study. To accomplish this, all recorded genera and species were assigned, respectively, to 11 and 12 phytogeographical elements established previously by the author. Two phytogeographical spectra were thus obtained and analysed. Results At the genus level, the Andean dry valleys of Bolivia are clearly dominated by genera that have widespread distributions (cosmoplitan and subtropical genera). Many of these reached the Andes from the lowland region of the Chaco. At species level, Andean elements constitute more than 60% of the species total, most of which are restricted to the central‐southern Andes. This suggests that Chaco‐related and Andean genera had considerable levels of speciation in these valleys. Many genera and more than half the species have their northernmost distribution in the dry valleys of Bolivia, thereby underlining strong relationships with central‐southern South America (mainly Argentina, Paraguay and southern Brazil). The data supports the belief of the existence, in central‐southern Peru, of a floristic disjunction in dry to arid environments that separates a tropical dry flora north of this limit from a dry subtropical/warm temperate flora south of it. Main conclusions The Andean dry valleys of Bolivia are diverse plant communities with high levels of endemism (c. 18% of the species). The species of this region are more related to those present in central‐southern South America than to the flora of northern South America that ranges southwards to Peru. Many of the species have restricted distributions in the dry Andes of Bolivia and Argentina, and many genera of these dry valleys have their northernmost distribution in Bolivia/southern Peru, too. The data point to high levels of speciation also in the central Andes.     

Pastoralist Decision-Making on the Tibetan Plateau

Despite a growing body of research about rangeland degradation and the effects of policies implemented to address it on the Tibetan Plateau, little in-depth research has been conducted on how pastoralists make decisions. Based on qualitative research in Gouli Township, Qinghai province, China, we analyze the context in which Tibetan herders make decisions, and their decisions about livestock and pastures. We refute three fundamental assumptions upon which current policy is premised: that pastoralists aim to increase livestock numbers without limit; that, blindly following tradition, they do not actively manage livestock and rangelands; and that they lack environmental knowledge. We demonstrate that pastoralists carefully assess limits to livestock holdings based on land and labor availability; that they increasingly manage their livestock and rangelands through contracting; and that herding knowledge is a form of embodied practical skill. We further discuss points of convergence and contradiction between herders’ observations and results of a vegetation analysis.     

Patterns of diversification in the high Andean Ponderacris grasshoppers (Orthoptera: Acrididae: Melanoplinae)

The Andes, the world's longest mountain chain, harbours great taxonomic and ecological diversity. Despite their young age, the tropical Andes are highly diverse due to recent geological uplift. Speciation either followed the orogeny closely or occurred after the Andean uplift, as a result of subsequent climatic changes. Different scenarios have been proposed to explain the diversification of high Andean taxa. The Melanoplinae grasshopper Ponderacris Ronderos & Cigliano is endemic to the eastern slopes of the Andes of Peru and Bolivia, mostly distributed between 1000 and 4000 m above sea level. Diversification in several montane habitats of Bolivia and Peru allows tests via cladistic analysis of distinct possible geographic modes of speciation. Eight species are recognized, with three described here as new with revised diagnostic morphological characters provided: Ponderacris carlcarbonelli sp.n., P. chulumaniensis sp.n. and P. amboroensis sp.n. Cladistic analyses of 15 species (8 ingroup and 7 outgroup) and 38 morphological characters, under equal and implied weighting, confirm the monophyly of Ponderacris. Characters from the external morphology and colour pattern provided less phylogenetic information than did the male abdominal terminalia and phallic complex. Species distributed in the Peruvian Andes constituted a monophyletic group, whereas those from the Bolivian Andes formed a basal paraphyletic grade. Dispersal–vicariance analysis resulted in one ancestral distribution reconstruction indicating that the most recent common ancestor was distributed in the Lower Montane Yungas of Bolivia. Eleven dispersal and one vicariant events are postulated, with a South‐to‐North speciation pattern coincident with progressive Andean uplift. Vicariance could relate to fragmentation of montane forest during the dry intervals of the late Cenozoic. From the Bolivian area, ancestral Peruvian Ponderacris may have dispersed northward, coinciding with the rise of the Andes. Ten of 11 dispersal events occurred at terminal taxa and are likely to be recent. However, diversification of Ponderacris cannot be explained solely by the South‐to‐North speciation hypothesis, but may also include both vicariance and dispersal across barriers influenced by Pleistocene climatic cycles.     

Reconsidering the Role of Absentee Herd Owners: A View from Mongolia

Since the privatization of livestock in 1992, rates of absentee ownership of livestock have increased sharply in Mongolia. Unlike other documented instances of absentee herding in pastoral societies, absentee herd ownership has few detrimental ecological or social impacts in Mongolia. Rather, the relationship between absentee herd owners and herders may be viewed as a revitalized institution, with links to customary patterns of urban-rural exchange, emerging to meet the needs of both herders and town-dwellers during the transition from a socialist planned economy to a free market economy. Absentee herding in Mongolia differs from absentee and contract herding accounts from Africa and the Middle East in its continuing emphasis on subsistence rather than speculative investment and accumulation. Other important distinctions include: (1) absentee owners and herders are usually kin or friends; (2) herders tend their own private herds in addition to absenteeowned animals;(3)few ethnic, caste, or class differences existbetween herders and absentee herd owners; and (4) herders from all wealth strata tend absentee-owned animals. Policies to restrict or regulate absentee livestock ownership must be carefully considered in the Mongolian context, making clear distinctions between informal, mutually beneficial subsistence-driven arrangements among kin and friends, and more formal investment-driven contracts between businesses or investors and herders.     

Estimating the size of the Andean Condor population in the Apolobamba Mountains of Bolivia

ABSTRACT.   Population estimates of Andean Condors ( Vultur gryphus ) are urgently needed across their range to more accurately assess their conservation status and design appropriate conservation measures. From July to September 2005, six feeding stations in the Apolobamba mountain range of Bolivia were monitored for 3 days each and all visiting condors were digitally photographed and video-taped. Using distinctive marks, the size and shape of crests, and plumage patterns, we were able to individually recognize 23 adult male Andean Condors, or 30% of all observed condors. By extrapolation, we estimated a minimum population size for Apolobamba during the sampling period of 78 condors. Considering that the current estimate of the total population of Andean Condors in Colombia is 180 individuals, our results illustrate the importance of Apolobamba for Andean Condors and the potential importance of the Bolivian Andes for the conservation of this species. The methods we used to attract, observe, photograph, videotape, and recognize adult males may be useful for other populations of Andean Condors, as well as for other species of threatened vultures.     

Environmental Stochasticity and Long-Term Livestock Viability—Herd-Accumulation as a Risk Reducing Strategy

This study tests the hypothesis that herd accumulation can be a risk reducing strategy aimed at increasing security in an unpredictable environment. Saami reindeer husbandry in Norway is characterized by environmental unpredictability and occasionally harsh winters can have dramatic negative effects on reindeer population densities. While herd accumulation has been found to be an adaptive risk reducing strategy in stochastic environments (i.e., individually rational), the accumulation of large herds may also result in collectively negative density dependent effects, which may negatively affect individual herders (i.e., collectively irrational). We found that individual husbandry units’ strategies, such as accumulating reindeer, have a larger effect on individual husbandry units’ herd size than a negative density-dependent effect.     

Quinua and Relatives (Chenopodium sect.Chenopodium subsect.Celluloid)

Traditionally viewed as an Andean grain crop,Chenopodium quinoa Willd. includes domesticated populations that are not Andean, and Andean populations that are not domesticated. Comparative analysis of leaf morphology and allozyme frequencies have demonstrated that Andean populations, both domesticated(quinua) and free-living(ajara), represent an exceptionally homogeneous unit that is well differentiated from allied domesticates of coastal Chile(quingua) and freeliving populations of the Argentine lowlands(C. hircinum). This pattern of relationships indicates that Andean populations represent a monophyletic crop/weed system that has possibly developed through cyclic differentiation (natural vs. human selection) and introgressive hybridization. Relative levels of variation suggest that this complex originated in the southern Andes, possibly from wild types allied withC. hircinum, with subsequent dispersal north to Colombia and south to the Chilean coast. Coastal populations were apparently isolated from post-dispersal differentiation and homogenization that occurred in the Andes. Other data point toward a center of origin in the northern Andes with secondary centers of genetic diversity subsequently developing in the southern Andes and the plains of Argentina. Comparative linkage of South American taxa, all tetraploid, with North American tetraploids of the subsection will eventually clarify this problem. While the possibility of a direct phyletic connection betweenC. quinoa and the Mexican domesticate(C. berlandieri subsp. nuttalliae,) cannot be excluded, available evidence indicates that the latter represents an autonomous lineage that is associated with the basal tetraploid, C. b. subsp.berlandieri, through var.sinuatum, whereas South American taxa show possible affinities to either var. zschackei or var.berlandieri. An extinct domesticate of eastern North America,C. b. subsp.jonesianum, represents either another instance of independent domestication, possibly from subsp. b. var.zschackei, or a northeastern outlier of subsp.nuttalliae.     

The role of cross-border transhumance in influencing resident herders’ cattle husbandry practices and use of genetic resources

For centuries, the humid West African coastal country of Benin attracts cattle herders from neighboring Sahelian countries such as Niger, Burkina Faso and Nigeria. Each year, several thousands of cattle are trekked over hundreds of kilometers to Benin following established and non-established transhumance corridors. This cross-border mobility has been shown to play an important role in ensuring the productivity of the transhumant herds through adequate late dry season access to pastoral resources. Yet, its effects on the traditional agro-pastoral production systems that are encountered along the routes in Benin have never been explicitly investigated. Therefore, we collected socio-economic household data, herd characteristics and management data from 104 resident herders and 38 transhumant herders in 2 vegetation zones of Benin. To determine whether or not the proximity to transhumance corridors affects sedentary production systems, characteristics of cattle herds and farmers’ management practices were compared between villages close to (within a 25 km buffer) and far from (outside a 25 km buffer) transhumance corridors within and between vegetation zones using non-parametric statistical tests. Existing relationships between resident and transhumant herders were also identified and characterized. Subsequently, herd characteristics and management practices were compared between resident herders having relationships with transhumant herders and those without. Herd sizes of resident herders living close to transhumance corridors were larger (P < 0.01) than those of their counterparts living far away. Also, proximity to transhumance corridors had positive effects on herd management practices. The relationships between resident and transhumant herders were governed by a variety of interests including encampment/manuring contracts, exploitation of grazing lands and watering points, trading and bartering of cattle. This exchange of cattle is an important driver of change in the breed composition of local herds and represents an opportunity for resident herders to enhance their herds’ productivity through crossbreeding. However, the mere replacement or indiscriminate crossbreeding of local cattle breeds with those kept by transhumant herders threatens the sustainability of the traditional resident herding systems by increasing the risk of genetic erosion and loss of valuable adaptive traits in indigenous animal genetic resources.     

Stepwise colonization of the Andes by Ruddy Ducks and the evolution of novel β‐globin variants

Andean uplift played a key role in Neotropical bird diversification, yet past dispersal and genetic adaptation to high‐altitude environments remain little understood. Here we use multilocus population genetics to study population history and historical demographic processes in the ruddy duck (Oxyura jamaicensis), a stiff‐tailed diving duck comprising three subspecies distributed from Canada to Tierra del Fuego and inhabiting wetlands from sea level to 4500 m in the Andes. We sequenced the mitochondrial DNA, four autosomal introns and three haemoglobin genes (α^A, α^D, β^A) and used isolation‐with‐migration (IM) models to study gene flow between North America and South America, and between the tropical and southern Andes. Our analyses indicated that ruddy ducks dispersed first from North America to the tropical Andes, then from the tropical Andes to the southern Andes. While no nonsynonymous substitutions were found in either α globin gene, three amino acid substitutions were observed in the β^A globin. Based on phylogenetic reconstruction and power analysis, the first β^A substitution, found in all Andean individuals, was acquired when ruddy ducks dispersed from low altitude in North America to high altitude in the tropical Andes, whereas the two additional substitutions occurred more recently, when ruddy ducks dispersed from high altitude in the tropical Andes to low altitude in the southern Andes. This stepwise colonization pattern accompanied by polarized β^A globin amino acid replacements suggest that ruddy ducks first acclimatized or adapted to the Andean highlands and then again to the lowlands. In addition, ruddy ducks colonized the Andean highlands via a less common route as compared to other waterbird species that colonized the Andes northwards from the southern cone of South America.     

Nutritional Consequences of Wealth Differentials in East African Pastoralists: The Case of the Datoga of Northern Tanzania

Among African pastoralists evidence that wealth is associated with other measures of household success is scant and the significance of wealth differentials for family welfare remains poorly understood. Predictions that wealth in livestock is associated with increased adequacy of household food supply and with improved child nutrition are tested with longitudinal data on herd size, household composition, food supply, and anthropometric status collected in 1992 for a sample of traditional Datoga households (n = 20) living in the Eyasi basin in northern Tanzania. Although a majority subsisted below poverty cutoffs estimated for east African pastoral populations, there was large variation among households in measures of wealth and resource availability. Individuals in relatively wealthy households did not appear to benefit either in terms of adequacy of household food supply or in terms of average growth performance of young children. Theoretical and methodological challenges in the study of the relationship between wealth and health in nomadic pastoralists are discussed.     

Coping with Uncertainty and Variability: The Influence of Protected Areas on Pastoral Herding Strategies in East Africa

A large number of East African pastoralists reside around protected areas (PAs). Over the last few decades pastoralists have been affected by the loss of grazing lands and increasing climatic variability. Many pastoralists who reside around PAs have resorted to grazing inside PAs to counter environmental variability. However, there is little information on how PAs influence the herding strategies of pastoralists. This case study from southern Kenya employs a spatially and temporally explicit mixed-methods approach to understand and evaluate the herding strategies of pastoralists around a PA. The results find that pastoralists access PAs on a regular basis, regardless of seasonality or herd size. Movement into PAs was partly driven by the loss of grazing land to conservancies. PAs affected pastoral herding by presenting differential opportunity costs to disparate groups. However, households with large herd sizes utilized the most flexible strategies to counter environmental variability and uncertainty.