Use and management of wild fauna by people of the Tehuacán-Cuicatlán Valley and surrounding areas, Mexico

The Tehuacán-Cuicatlán Valley (TCV) is an area of about 10,000 km², located in south-eastern Puebla, north-western Oaxaca and the limits of Veracruz [34, 35] (Fig. 1). A high environmental heterogeneity can be found in the area, with warm, semi-warm and temperate climates and average annual precipitation ranging from 400 to 700 mm [42]. The region is part of the Papaloapan River basin and comprises several valleys and mountain chains with elevations from 595 to 2950 m above the sea level [43]. The surrounding mountains, belonging to the physiographic regions known as “Sierras Orientales” and “Sierras Centrales de Oaxaca”, harbour a gradient from thorn-scrub, tropical deciduous forest, oak forest, pine-oak forest in leeward slopes, and cloud forest and tropical rain forest in windward slopes [35]. The orographic shadow caused by the Sierra Madre Oriental prevents humidity from the Gulf of México and causes the arid condition of the TCV. The TCV is one of the five floristic provinces belonging to the Mexican Xerophytic Region in the Neotropical region [44]. At least 29 plant associations belonging to six general vegetation types have been recognized in this area, which stands out from other arid zones in North America for its high floristic diversity [34, 43] and its high species richness of mammals and birds [45].

The use and management of regional biodiversity have been part of the subsistence strategies of the human groups inhabiting the TCV for approximately, the last 14,000 years [46]. Wild plants and animals ranging from big vertebrates (i.e. antelopes, horses, deer) to insects have been part of the human diet since prehistoric times [47, 48]. Archaeological records suggest that they were obtained through hunting, trapping and gathering [47]. The management of agroforestry systems, a current practice in the area [49, 50] is also millenary in the region [46]. The human groups currently inhabiting the Valley (i.e Ixcatec, Nahua, Cuicatec, Mixtec, Popolocan, Chocholtec, Chinantec, Mazatec and Mestizo) use over 2,000 of the more than 3,000 vascular plant species occurring in the TCV, and at least 610 of them are subject to management practices [4, 25]. However, information regarding the current use and management of fauna by human groups in the area is not systematized. In 1998, a Biosphere Reserve of about 500,000 ha was decreed by the Mexican Federal Government Ministries SEMARNAT and CONANP [51] within the TCV. Within this Biosphere Reserve, an area of 145,255 ha was included in 2018 as a Mixed Site in the World Heritage List by the UNESCO because of its cultural and biological significance [52].

In our research, we included documented contemporary records of fauna occurring in any of the 107 municipalities (1436 localities) of the TCV and surrounding areas (Fig. 1a; the list of localities and municipalities is provided in the Additional file 1). We also included evidence from the archaeological records analysed by Kent V. Flannery, Eric O. Callen and Richard S. MacNeish [46‐48].

We conducted documental research between January 2018 and January 2019 for the available scientific literature about the occurrence, use and management of wild fauna in the studied region. Firstly, we consulted key documents (published between 1967 and 2018 according to the taxa) [47, 48, 51, 53‐71]. Afterwards, we consulted the digital repositories of EBSCO, SCOPUS, Web of Knowledge and Google Scholar for subsequent records; in addition, those of specialized journals (Therya, Revista Mexicana de Biodiversidad, Acta Zoológica Mexicana, Etnobiología, Huitzil, Journal of Insects for Food and Feed, Journal of Ethnobiology and Ethnomedicine, Journal of Ethnobiology, Ethnobiology letters, Ethnobiology and Conservation); and those from dissertations of three Mexican universities (UNAM, UAM, BUAP). Our search included 70 keywords in English and Spanish (Additional file 2). In order to collect the information, we considered the following criteria: (a) geographic: we included those records of the occurrence, use or management of fauna that felt within the study area (see Fig. 1 and text in the “Study area” section), (b) conceptual: for management, we took Zeder’s approach and excluded works about zootechnical intensive management (see “Background” section), (c) time-line: all included records are subsequent to the research by Flannery, Callen and MacNeish 1967, (d) publication criteria: theses, dissertations, peer-reviewed journals, published abstracts and, in the case of fauna occurrence records, we included 7 databases hosted by CONABIO and GBIF). These criteria made possible to include practically all found works except for two sources that felt out of the geographical area and were instead considered in the references as other areas of Mesoamerica. We included records about the presence of fauna from 38 documentary sources (21 scientific reports published in peer-reviewed journals, seven public databases, four lists, three book chapters, two dissertations, one book) and registers about the use or management of this fauna from 15 documentary sources (five scientific reports published in peer-reviewed journals, four dissertations, three book chapters, one book, one scientific dissemination article, one conference abstract). We also included primary information generated through the inventory of records of our own observations of living individuals, hunting evidence, and commercialization and interchange in the regional markets, as well as mentions in semi-structured interviews conducted with 30 Cuicatec households [72], 30 people in a mestizo locality with Popolocan ancestors in Zapotitlan, Puebla [73], 16 Ixcatec people in 2016, and with 16 people in three Nahua localities in Coyomeapan, Puebla (2018-2019). Additionally, we went through the checklist of species present in TCV with no local reports of use and checked their uses reported in other regions of Mesoamerica. This search based on the set of animal species whose presence has been reported so far within the TCV (references cited in Table 1). Specifically, if ethnozoological information for an animal species occurring in the study area but no information reported for the TCV region, we searched for at least one kind of use of the animal outside the TCV. This was done in order to analyse the supposition that in the TCV region, there would be more species under any use or management categories than those that have been documented so far. We included 38 documental sources (25 scientific reports published in peer-reviewed articles, ten book chapters, one dissertation, one book, one scientific dissemination article). To avoid repeated information associated with synonymy, the nomenclature was checked with the Integrated Taxonomic Information System 2019 and 2000 [144].

Vertebrates in the TCV were represented by 652 species, nearly 11.50% of them reported as being used by people of the area. However, when including the uses reported for other regions of Mesoamerica, we found that 35.73% of the vertebrate species occurring in the TCV are potentially used as resources. Out of the 765 species of insects that have been reported in the TCV, only 4.18% was reported to be used in the region, but about 6.79% of them are used in other regions of Mesoamerica (Table 1).

Overall, the animals used in the TCV were found distributed in the following use categories: 70 edible species, 30 ornamental, 23 medicinal, 13 weather indicators; six are used as amulets, five as ceremonial or for rituals, four as animal companion and tools, respectively, three are melliferous, two are used for crop protection and one for recreational purposes (Table 2). The number of different uses per species ranges from 1 to 5 (all species mean = 1.54 ± 0.08 SE, n = 109; birds mean = 1.36 ± 0.09 SE, n = 36; mammals mean = 2.34 ± 0.22 SE, n = 26; reptiles mean = 1.14 ± 0.14 SE, n = 7; insects mean = 1.28 ± 0.12 SE, n = 38). The species with more uses reported are the naturalized bee Apis mellifera and the temazate deer Mazama temama Kerr, 1792 (Fig. 2). Bee’s larvae are edible, honey is extracted, wax is used as glue, to elaborate candles with ceremonial motives and to make art pieces. The meat of the temazate deer is edible, the male reproductive structures are used to facilitate childbirth, heads and legs are prepared through taxidermic techniques to be used as ornaments and their skulls are exposed as a protection against “mal aire” (cultural illness or the agent that causes it).

We found evidence for treatments of at least twelve health issues, which involve animals (Table 3). Childbirth facilitation involves the use of 26.08% of the medicinal species, mainly mammals. Cancer treatment included 21.73% of the species, mainly snakes and birds. Treatments for skin problems and epilepsy both involved 17.39% of the species. Mammals, snakes and immature stages of Lepidoptera are used in the treatment of skin problems while mammals, snakes and birds are used for alleviating epileptic seizures. Trochilidae (hummingbirds) is the family used for more medical treatments. According to our ethnographic records in Coyomeapan, inhabitants of the region perceive that prior to the arrival of the vaccines in the 1970s, the use of birds and mammals to treat illnesses was much more common. An example of this is the larger former use of species of Trochilidae and Geomyidae Bonaparte, 1845 in remedies for whooping cough, especially for children.

Each species can be used by 1 to 4 human groups (mean = 1.65 ± 0.09 SE, n = 93); 36 species are used by the Mestizo people, 32 by the Cuicatec and the Nahua, respectively, 23 by the Ixcatec, 19 by the Mazatec and 13 by the Popolocan people. The animals used by more cultural groups of the region are the mammals Nasua narica Linnaeus, 1766, Pecari tajacu Linnaeus, 1758, Sciurus spp. and Sylvilagus spp. (Fig. 3).

The animals used in any of the categories documented have been reported in at least eight vegetation types along the TCV (Table 4, Fig. 4). This information should be considered as potential systems from which fauna is extracted, rather than the actual systems from which it is extracted.

Animals in the TCV region are obtained through extractive practices. We recorded the following general management categories:

The number of management categories per species ranges from 1 to 4 (all animals mean = 1.51 ± 0.08 SE, n = 71; birds mean = 1.76 ± 0.16 SE, n = 26; mammals mean = 1.61 ± 0.16 SE, n = 21; reptiles mean = 1 ± 0 SE, n = 6; insects mean = 1.27 ± 0.13 SE, n = 18) (Table 2, Fig. 7).

The ordination of species along with the number of uses, the number of management categories and the number of cultural groups suggested correlation among these three variables but this was only significant between the number of uses and cultural group (r_s = 0.586, n = 71, p value < 0.001). The first principal component explained 59.84% of the variance and the second one 26.69%. Most of the variation along the first principal component is given by the differences in the number of uses and the number of cultural groups that use the species. Variation in the number of management categories was explained by the first and the second principal components (Fig. 8, Table 7). In the right half of the biplot are the species with more types of uses, more types of management and which are used by more cultural groups. The species related to more types of management practices such as doves, the edible Lepidoptera larvae A. armida, the temazate deer (M. temama), cottontail rabbits (Sylvilagus spp.) and P. tacaju are concentrated in the upper right quadrant of the biplot.

Early settlers in the region integrated vertebrates and insects to their diet since before 12,000 B.P. Although the dates have been subject to discussion, the history of animal use in the TCV proposed by MacNeish, Flannery and Callen remains useful to characterise the general chronology of events. At least 20 of the species reported by these authors are still currently used in the area (Table 8).

According to the chronology reported, there is evidence of hunting and edible use of an American horse (Equus sp.) Linnaeus, 1758 and an antelope (Antilocapridae, probably Antilocapra americana (Ord, 1815)) about 12,000 B.P. along with small prey such as jackrabbits (Lepus sp. Linnaeus, 1758), coyotes (C. latrans), skunks (Spilogale sp. Gray, 1865, Conepatus sp. Gray, 1837), foxes (Urocyon sp. Baird, 1857), ring-tailed cats (Bassariscus astutus (Lichtenstein, 1830)), squirrels and turtles. Later, horses and antelope were extirpated from the area. From 10,000 B.P., the white-tailed deer (O. virginianus) appears in the archaeological record as hunted and consumed by humans, along with cottontails, lizards, skunks and other small prey. According to MacNeish, Flannery and Callen, small preys were probably obtained by trapping. Ants and immature stages of Lepidoptera were found in coprolites dated between 8500–7000 B.P. and 7000–5500 B.P., respectively, in the Coxcatlán area. Between 7000 and 4300 B.P., hunting of the white-tailed deer seems to have intensified, and there was also hunting of P. tajacu and trapping of small games. Around 5000 B.P., the domestic dog (C. lupus familiaris) was introduced into the area. Between 4300 and 3500 B.P., remains of dogs with evidences of human consumption increased. Between 3000 and 1800 B.P., gophers increased in the archaeological record. Then, around 1500 B.P., the first turkey (M. gallopavo) appeared in the area.