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[Jeannine Lander]

Copyright2023 Bird et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Funding: This study was funded by the National Geographic Society, USA ( ) (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) ( ) 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.

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):

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.

Places visited and indicated in text. Spider symbols indicate communities where the garden orb-web spider, Argiope australis (Walckenaer 1805) (Araneidae), is thought to be used as arrow poison. Map generated in QGIS 3.24.1 by T.L. Bird using personal datasets and open source shapefiles.

Research into IK is highly sensitive due to a long history of colonization, slavery, abuse, and marginalization of indigenous communities and because of multiple legal cases about uses, patents, and profits from such knowledge [49]. One particular case of the San people has involved lengthy legal battles over the indigenous hunger-suppressant plant, Hoodia Sweet ex Decne. (Apocynaceae) [49]. Given this backdrop, we took great care that our research approach was unambiguous, respectful, and legal. In addition to the legal requirements of the government of Botswana, the San communities also have their own regulations over data sharing (interviews, photographs, films, and data). Thus, our research met several layers of legal permits, ethical considerations, and community permissions at both the national and at each local community level.

We visited three G/ui and G//ana San communities (villages) in the CKGR, Botswana (Fig 1; Tables 1 and S1). These communities were identified based on size and presence of people with past hunting experience. Smith Moeti (SM) conducted two solo follow-up expeditions to the same communities in December 2019 and 2021 with follow-up questions and observations in different seasons. In February 2022, during a third solo visit to one of the communities, he observed a hunt.

Our multidisciplinary team consisted of 12 members from the social and biological sciences, and from several institutions: 1) the Botswana International University of Science and Technology (BIUST), Palaype, Botswana, 2) the Nebraska State Museum, USA, 3) the University of New Mexico, USA, and 4) the Kalahari Peoples Fund (KPF, an NGO), Botswana. Co-author SM is of San heritage and a speaker of mother tongue languages G/ui and G//ana. During the group expedition, SM, originally of Metsiamonong, served as a translator and guide. Other members include TLB, a spider systematist; CSC, a beetle systematist; and RKH and MCK, who are anthropologists. Although no botanist accompanied the team, plant samples were collected and identified by the National Herbarium of the Botswana National Museum & Monuments (hereafter Botswana National Museum), Gaborone, Botswana.

The central Kalahari is a subset of the Kalahari (the latter is often termed Kalahari Desert, e.g., [58]). The Kalahari is an ill-defined semi-arid area that forms the core of the Kalahari Basin [58]. The vegetation is classified as arid savanna-type vegetation [59] (Fig 4A). Except for the Delta areas, the Kalahari has no permanent surface water. Temporary pans, which are shallow depressions of a few to several km2 that hold rainwater after rains [58] are widespread [59,60,61]. Oscillations in wet and dry, along with loss of surface water, affected vegetation, wildlife, and humans [62].

Two legally-recognized conservation areas, the Central Kalahari Game Reserve (CKGR) and the Kutse Game Reserve, occupy most of the central Kalahari. Our fieldwork was restricted to the CKGR (Fig 1). Indigenous peoples, as well as ecologists and anthropologists, recognize subtle ecological variation with specific vegetation, wildlife, and geomorphology [37,52]. The Okwa and Deception valleys are two noteworthy topographical features within the CKGR [37]. Silberbauer [37] recognized three distinct vegetation regions within the CKGR: (a) dune woodlands in the north, (b) central grassland plains between the Okwa and Deception valleys, and (c) savanna thornveld in the south. The wildlife population is typical Kalahari species. Apart from mammals, much of the wildlife is largely undocumented: some checklists for Botswana are available for plants (Bryophytes, Pteridophytes and Angiosperms [65]), mammals [66], birds [67], reptiles [68,69], and some arthropods (Araneae [70]; Lepidoptera [71]; Odonata [72]). In the past, the central Kalahari has seen large-scale migrations of wildebeest toward the Boteti River and to Lake Xau [73].

In 1961 the Central Kalahari Game Reserve (CKGR) was created in Botswana for the purpose of protecting the land and what is on it, and to secure the living on, and utilization thereof, by the San that lived on the land [85]. Today there are approximately 350 San settled in five communities scattered in the CKGR, namely Gope, Gugamma, Metsiamonong, Molapo, and Mothomelo (Fig 1; Tables 3 and S1).

The San that live in the CKGR receive some government support. However, anecdotal evidence suggests that food distribution is minimal and can be irregular. Individual access to income- and age-based government aid may also be hampered when they lose their identity cards (e.g., in fires) given the logistical complexities to get replacements (data from our community interviews). The Baboalongwe Bakgalagadi practice sedentary agropastoralism [86], but communities employ gathering and some hunting strategies to supplement food. Small animals such as duiker Sylvicapra grimmia (Linnaeus 1758), steenbok Raphicerus campestris Thunberg 1811, and springbok Antidorcas marsupialis (Zimmermann 1780) are hunted through trapping and snares while larger animals such as hartebeest Alcelaphus caama (. Geoffroy Saint-Hilaire 1803) and kudu Tragelaphus strepsiceros (Pallas 1766) are hunted by bow-and-arrow or spear [40; W. Barnard, pers. comm.] (Table 4).

We followed the World Spider Catalog [88] for current scientific names of spiders. For plants we followed IPNI [90], and Wilson and Reeder [91] for mammals. Spiders were identified using the relevant keys to species, namely Bjrn [92] for Argiope australis (Walckenaer 1805); De Wet and Dippenaar-Schoeman [93] and Gallon [94] for Ceratogyrus darlingi Pocock 1897. Spiders and beetles (Chrysomelidae) collected as voucher specimens were deposited in the Ditsong National Museum of Natural History, Pretoria, South Africa. Plants collected were deposited in the National Herbarium of the Botswana National Museum, Gaborone, Botswana (S2 Appendix).

Through our fieldwork and interviews we 1) established that traditional hunting with bows and arrows persist today in the San communities of the Kalahari, Botswana; 2) found that Chrysomelidae beetles are being used as poisons; and 3) discovered that spiders (Arachnida: Araneae) are also used as poisons. No other venomous animals (scorpions, snakes) are used as poisons in these communities. We discuss these findings below and explore more deeply the discovery of spiders as hunting poisons.

This pouch holds small equipment and small game (e.g., rabbit). Our hunter informants indicated a preference for leather made from gemsbok Oryx gazelle (Linnaeus 1758) as they consider other leathers too thick to pull to shape.

Based on his solo interviews with hunters, SM concluded that spider-poisoned arrows tend to be used for smaller antelope (duiker, steenbok), and beetle-poisoned arrows for larger animals. Arrows for hunting rabbits are not poisoned. Our informants indicated that they may sometimes use spider poisons to hunt large mammals such as eland Taurotragus oryx (Pallas 1766), giraffe Giraffa camelopardalis Linnaeus 1758, hartebeest, impala Aepyceros melampus (Lichtenstein 1812), leopard Panthera pardus (Linnaeus 1758), and lion Panthera leo (Linnaeus 1758).

The informants that we interviewed reported that they use beetles as the active ingredients in arrows. However, due to the seasonality of our expedition, we were unable to collect specimens to verify the identity. During follow-up visits, SM vouchered some adult beetles that the San informants indicated as the kind used in their poisons. These adult beetles are identified by CSC as Diamphidia Gerstaecker 1855 (Coleoptera: Chrysomelidae). No juveniles were collected so we cannot associate lifestages to specify the species used for poisons, nor detail further their use in the CKGR. Our fieldwork only confirmed that beetle poison is still used, similar to that of Hai//om San in Namibia (see [5,34]).

Our field notes and observations of spider arrow poison application by the G/ui and G//ana San in the Kalahari in Botswana provide evidence that A. australis is applied as an active ingredient in their arrow poison, but that the active component is not the venom since only the abdomen (which does not contain the venom glands) is used. Here, we highlight the implications of our observations, we consider possible mechanisms of how an Argiope-based poison (contents from abdomen, e.g., haemolymph, tissues, and digestive fluids) could lead to a kill, and we propose hypotheses of cognitive processes involved in selecting an Argiope sp. as an arrow poison ingredient and in using the abdomen rather than the prosoma thereof. These hypotheses need to be scientifically investigated. We conclude with a note on the loss of San IK.

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