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Orb-web spider Argiope (Araneidae) as indigenous arrow poison of G/ui and G//ana San hunters in the Kalahari [1]

['Tharina L. Bird', 'General Entomology', 'Ditsong National Museum Of Natural History', 'Pretoria', 'South Africa', 'Department Of Zoology', 'Entomology', 'University Of Pretoria', 'Hatfield', 'Department Of Biological Sciences']

Date: 2023-01 

Hunting has been crucial in early human evolution. Some San (Bushmen) of southern Africa still practice their indigenous hunting. The use of poisons is one remarkable aspect of their bow-and-arrow hunting but the sources, taxonomic identifications of species used, and recipes, are not well documented. This study reports on fieldwork to investigate recent indigenous hunting practices of G/ui and G//ana San communities in the Central Kalahari Game Reserve (CKGR), Botswana. Here we discuss their use of spider poison. The hunters use the contents of the opisthosoma (‘abdomen’) of a spider as sole ingredient of the arrow poison and discard the prosoma that contains the venom-glands. Using taxonomic keys, we identified the spider as the garden orb-web spider Argiope australis (Walckenaer 1805) (Araneidae). The hunters’ choice of this species is remarkable given the scientific perception that A. australis is of little medical importance. The species choice raises questions about how the spider fluids could kill game, particularly when the prosoma, which contains the venom glands, is not used. Possibilities include trauma, as a source of pathogens, or abdomen-containing toxins. Based on characteristics of Argiope Audouin 1826, we hypothesize that the choice of this species for arrow poisons might have evolved from the recognition of aposematic signalling or spiritual symbolism. Indigenous knowledge (IK) is an important source for advances in biotechnology but is in decline worldwide. The study contributes to the documentation of the San people, and their ancient IK, which is threatened by marginalization, political pressures, and climate change.

Funding: This study was funded by the National Geographic Society, USA ( https://www.nationalgeographic.com/ ) (PI CSC) under Grant # NGS-181R-18 to CSC, RK, Casper Nyamukondiwa, megan Biesele, and TLB. TLB acknowledge, with thanks, funding provided by Botswana International University of Science and Technology (BIUST) ( https://www.biust.ac.bw/ ) for the symposium and contribution to expedition logistics. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Introduction

Indigenous knowledge (IK) and traditional ecological knowledge (TEK) (or traditional knowledge (TK); Native Science) are highly specific to location, environment, and season [1–4]. Indigenous hunting is one aspect of IK that that has evolved over the course of human history. An important event in early human cognitive evolution is the shift from hunting with spears to hunting with bow-and-arrow [5]. Stone points on traditional arrows were replaced with bone tips, possibly some 60,000 years ago [4,6,7]. Poisoned arrows have also been part of ancient human hunting kits. The use and evolution of such poisons in indigenous hunting kits remain little understood, and are difficult to investigate because organic matter on archaeological artefacts is hard to detect in archaeological sites and degraded residues on artefacts, and there is not necessarily continuity between prehistoric and contemporary use of resources [8]. In southern Africa, the use of poison in hunting kits has now been dated to ~24,000 years bp [9]. Knowledge about contemporary indigenous hunting and contemporary hunting poisons might provide insight into the evolving cognitive processes involved in selecting materials and ingredients.

An important clue to the evolution of hunting poisons might be the extent to, and the manner in which, venomous versus poisonous ingredients are included in hunting poisons. ‘Poison’ is often used loosely to encompass all kinds of toxins. There is, however, an important distinction between ‘venom’ and ‘poison.’ Following the definitions of Nelsen et al. [10], ‘venom’ refers to toxins that are delivered through a wound made through, for example, a fang (e.g., snakes, spiders, centipedes) or a sting (e.g., scorpions, bees), whereas ‘poison’ refers to toxins that enter the body through, for example, inhalation or ingestion (e.g., ingestion of milkweed latex, Asclepias L. spp. (Asclepiadaceae) is poisonous to humans and many other species; monarch butterflies, Danaus Kluk 1802 spp. (Lepidoptera: Nymphalidae) feed on the milkweed as caterpillars, accumulate milkweed toxins, and thereby become similarly poisonous to avoid predation). Traditional hunters use the word ‘poison, as in ‘arrow poison’, colloquially (which we will adhere to here, consistent with previous literature), but by scientific technical definitions, it is a venom because it enters the body by penetration.

By these definitions, spiders are categorized as venomous, not poisonous, due to the injection of toxins through their fangs. Spiders have been documented as hunting poisons in different cultures globally. The earliest report was by von Siebold [11: 201) who recounted observations that Hayashi Shihei made of the Ainu on Hokkaidõ Island, Japan, in the 18th century. Spider poisons have also been reported from the Americas, Asia, and Africa (e.g., [5,12–20]). Indigenous Americans used macerated black widow spiders (Latrodectus Walckenaer 1805 spp., Theridiidae), sometimes with their eggs [21,22]. In Asia, the North Pacific Ainu tribes reportedly crushed spiders with poisonous aconite (monkshood) root or surku poisons (Aconitum L., Ranunculaceae) [17,23–25]. In southern Africa, the earliest mention of spiders incorporated in arrow poison is by Farini [26]. Similar reports emerged over the years [14,15,27–33], but Farini [26] appears to remain the only first-hand witness of poison preparation by the San when spiders are included in the poison recipe.

These mostly Western anecdotes of Native Science cultural practices have not been ground-truthed with first-hand interviews, scholarly approaches to biodiversity observation, and collections of specimen vouchers to verify taxonomic identifications. The type of spider and manner of preparation are rarely placed in context with the particular communities. The situation is similar to that highlighted by Chaboo et al. [34: 12] in their investigation of the use of beetles as arrow poisons amongst San (Bushmen):

Today, the San’s bow-and-arrow hunting and attendant tracking knowledge have a mythical status, but the facts of the poison sources and preparations are largely unclear. Several factors contribute misconceptions, outright errors, and ambiguous information about San arrow poisons. First, the use of the term ‘Bushmen’ for diverse San [communities] obscures apparent geographic variation in poison sources, recipes, and preparations. Second, insect taxonomists have rarely been involved in specimen identifications. Third, chemists analyzed specimens with presumed taxonomic identifications and left no specimen vouchers to confirm the species involved.

The San are considered the first indigenous people of southern Africa and have historically lived as hunter-gatherers (e.g., [35]). They use poisons from both plant and animal sources [5,14,16,20,34,36–39]. In addition to Chrysomelidae leaf beetles [5,34, and citations therein], reported animal sources of San arrow poisons are snakes, geckos, centipedes, scorpions, and spiders [14–16,27,28,31]. San indigenous hunting techniques have included snares [37,40], nets and pitfalls (or game pits) [15,27,31,41], traps ([40]; spring-traps [37]), hooks (= springhare hook [1,31,40]), spears, and bows and arrows [42]. Tools used by contemporary indigenous San hunters often show design modifications using alternative contemporary raw materials. For example, the use of wire instead of sinew or plant fibre for snares [31], plastic pipes for quivers instead of wood [8], and the replacement of bone or later stone tips with glass [27] or metal [27,40,43]. Metal can be cut from tins [31], fence wire [8,37,39), steel from vehicles [37], or metal nails hammered to triangular arrow points [29,34]. Information of change in arrow poisons is scant (see also [20]).

Due to years of exploitation and encroachment into their territories, San groups have lost much of their ancestral land in the Kalahari [44,45]. Today, there is unlikely any San in southern Africa who depends entirely on hunting-gathering as a way of subsistence [46,47]. However, whether hunting with bows and arrows persists in these areas remain largely unknown. Similarly, information regarding the identity and diversity of arrow poison ingredients, and preparation methods thereof all remain scant.

Traditional hunting with plant and beetle poisons by Hai//om and Ju’hoansi San communities in Namibia was investigated previously [5,34]. To answer these same questions, we visited G/ui and G//ana San communities in the Central Kalahari Game Reserve (CKGR), Botswana (Fig 1). We set out to investigate particularly whether indigenous G/ui and G//ana San are still hunting with bow-and-arrow, whether they use arrow poisons, and the identity of the poisons they use. We interviewed hunter-informants on their tools and poisons, and we collected voucher specimens. Here we discuss only our findings relating the spiders as ingredients in their arrow poisons, specifically whether spiders are used, which species, and how. We discovered that spiders are used in arrow poisons, but, surprisingly, not the venom of the spider. Thus we consider here the possible mechanisms through which this spider-based poison could facilitate a successful hunt. We also explore the cognitive processes that may have influenced the selection of this species for poisons. Given the erosion in hunting IK [48], and the marginalisation of hunting and gathering, our study is significant in documenting the remnants of critical San knowledge on the use of spiders as sources of arrow poisons, methods of preparation, application process, and use.

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