diamphidia nigroornata or bushman arrow-poison beetle , is an african leaf beetle species in the genus diamphidia .

the larvae and pupae of diamphidia produce a toxin used by san people as an arrow poison .

the finnish explorer hendrik jacob wikar , who travelled in southern africa in 1773 – 1779 , described the larvae as " poisonous worms " .

hans schinz was the first scientist to document the process by which the san people extract and use the poison .

the adults and larvae of diamphidia nigroornata feed on commiphora angolensis .

diamphidia nigroornata

the aduwts and warvae of diamphidia nigroornata feed on commiphora angowensis ( engwer ) , whereas diamphidia vittatipennis eat african myrrh .
the bushman arrow toxin , diamphidia toxin : isolation from pupae of diamphidia nigro - ornata .
the bushman arrow toxin , diamphidia toxin : isolation from pupae of diamphidia nigro - ornata .
the bushman arrow poison toxin , diamphidia toxin : isolation from pupae of diamphidia nigro - ornata .
preliminary studies on the chemical properties of the toxic principle from diamphidia nigroornata larvae , a source of bushman arrow poison .
preliminary studies on the chemical properties of the toxic principal from diamphidia nigroornata larvae , a source of bushman arrow poison .
the bushman arrow toxin , diamphidia toxin : isolation from pupae of diamphidia nigro - ornata . - pubmed - ncbi
( 1984 ) . the bushman arrow poison toxin , diamphidia toxin : isolation from pupae of diamphidia nigro - ornata .
preliminary studies on the chemical properties of the toxic principle from diamphidia nigroornata larvae , a source of bushman arrow poison . - pubmed - ncbi
( 1982 ) . preliminary studies on the chemical properties of the toxic principal from diamphidia nigroornata larvae , a source of bushman arrow poison .
minden pictures stock photos - flea beetle ( diamphidia nigroornata ) larvae used to poison arrows are thought after by kalahari or kung desert bushman . . .
minden pictures stock photos - flea beetle ( diamphidia nigroornata ) feeding on myrrh tree ( commiphora sp ) larvae produce a toxin not found in adults w . . .
based on our specimen collections of larval , pupal and adult stages ( we did not collect egg stages ) with the ju’hoansi san at nyae nyae , we determine those beetles as diamphidia nigroornata . bryant ( 1942 ) indicates several species names as synonyms of diamphidia nigroornata . other researchers have photographed poison beetles used in the nyae nyae region and which are a different species from our sample . thus , several diamphidia species are used for poison .
diamphidia toxin , the bushman’s arrow poison : possible mechanism of prey - killing .
diamphidia or bushman arrow - poison beetwe , is an african genus of weaf beetwe . diamphidia in de chrysomewidae famiwy , de same famiwy as de coworado potato beetwe .
effect of diamphidia toxin , a bushman arrow poison , on ionic permeability in nucleated cells .
diamphidia toxin , the bushmen ' s arrow poison : possible mechanism of prey - killing .
a drop of haemolymph of the diamphidia vittatipennis larva is squeezed from the larva onto the arrow shaft .
arrow - poison beetles of the ju | ’hoansi , tswumke conservancy , namibia ( photos : cs chaboo ) . 11 typical collecting for beetle cocoons at base of a commiphora shrub in the drip line 12 ostrich egg - shell full of cocoons of diamphidia nigroornata ab . locusta 13 parasitoid carabid larva ( left ) and diamphidia larva ( right ) extracted from collected cocoons 14 cleaned 4 th instar diamphidia larva extracted from cocoon 15 adult diamphidia beetle in cocoon 16 squeezing the contents of leaf beetle larvae onto giraffe bone to prepare arrow poison 17 typical hunting implements , quiver , bow , fire - sticks , and arrows .
woollard , j . m . r . , fuhrman , f . a . & mosher , h . s . 1984 . the bushman arrow toxin , diamphidia toxin : isolation from pupae of diamphidia nigro - ornata . toxicon 22 : 937 - 946 .
larvae are parasitised by a species of carabidae beetle called lebistina holubi . the lebistina larvae by attaching themselves to full - grown diamphidia vittatipennis larvae , get into their cocoons where they feed on blood and soft parts of the diamphidia larvae , eventually killing them .
preliminary studies on the chemical properties of the toxic principle from diamphidia nigro - ornata larvae , a source of bushman arrow poison .
the tsila , botswana . the tsila ( ~ 500 people ) are found in the central kalahari game reserve , in the northern kweneng district and the eastern central district . vierich ( pers . comm . 2014 ) observed their use of arrow poison derived from diamphidia nigroornata and an unidentified plant .
arrow - poison beetles of the san people and their host plants ( photos : cs chaboo , or indicated if otherwise ) . 2 diamphidia nigroornata ståhl ( = diamphidia simplex péringuey , = diamphidia locusta fairmaire ) , namibia ( chrysomelidae ) 3 polyclada sp . ( chrysomelidae ) 4 blepharida sp . , kenya ( photo : c smith , usnm ) 5 lebistina sp . ( carabidae ) 6 diamphidia femoralis ( above ) and its predator - parasitoid enemy , lebistina ( below ) , on commiphora plant in south africa ( photo : k ober ) 7 lebistina sanguinea ( boheman ) adult beetle on a commiphora plant in south africa ( photo : e . grobbelaar , sanc , arc - ppri ) .
woowward jm , fuhrman fa , mosher hs ( 1984 ) .
the bushman arrow toxin , diamphidia toxin : isowation from pupae of
this beetle in different stages of its live cycle can be found on and in the near vicinity of its host plant the commiphora sp . in particular the dza or also known as sand corkwood ( c . angolensis ) , which is a very important tree for the san for various reasons , and host plant for diamphidia nigroornata and the hairy corkwood or also known as ' haarige kanniedood ' in afrikaans , a host to the diamphidia simplex and diamphidia vittatipennis . these cocoons can be found near the roots of their particular host plants at between 20 cm and 1 meter deep .
diamphidia is parasitised by a carabid lebistina beetwe , de warva of which attaches itsewf to a mature diamphidia warva , cwinging to it untiw de diamphidia has formed its cocoon , encwosing bof host and parasite , and den feeding on its host ' s soft tissue . the lebistina warvae are more toxic dan deir hosts and are preferred by san hunters for arrow poison , uh - hah - hah - hah . [ 1 ]
two main species of diamphidia are used namely diamphidia nigroornata ( host plant commiphora angolensis ) and diamphidia vittatipennis ( host plant commiphora africana ) . in addition , larvae of the chrysomelid beetle polyclada flexuosa ( host plant sclerocarya caffra ) can also be used as a toxin . each of these three species is also parasitised by carabid beetles in the genus lebistina . the lebistina larvae , by attaching themselves to full - grown diamphidia and polyclada larvae , remain with them when they make their cocoon . inside the cocoon , they feed on blood and soft parts of the chrysomeld larvae , eventually killing them . the lebistina larvae are also toxic , in fact the san hunters evidently find them more toxic than the chrysomelid larvae and favour these parasitic larvae for their arrows .
fecal ecology in leaf beetles : novel records in the african arrow - poison beetles , diamphidia gerstaecker and polyclada chevrolat ( chrysomelidae : galerucinae ) .
steyn dg . a bushman arrow poison ( diamphidia nigro - ornata stal . ) . s afr med j . 1957 ; feb 931 : 119 - 20
new records of diamphidia gerstaecker 1855 and polyclada chevrolat 1840 in the republic of mozambique ( chrysomelidae : galerucinae s . l . ) . the coleopterists bulletin .
aduwt femawes of diamphidia femorawis gerstaecker and diamphidia nigroornata ( ståw ) way deir eggs on de stems of commiphora species and coat de eggs wif deir faeces which hardens into a protective covering . as de warvaw instars devewop , de pewwets of deir own faeces remain attached to deir backs and posteriors . [ 3 ] the finaw instar sheds dis faecaw coat when entering de soiw to pupate . the same behaviour is found in bwepharida , a fwea beetwe and powycwada , de african leaf beetwe . [ 3 ] the diamphidia warvae burrow down for a depf of up to 1 metre in de sand under de food pwant , where dey may wie dormant for severaw years before going drough a very rapid pupaw phase .
digging up cocoons of diamphidia beetles in the vicinity of their host plant commiphora sp . the cocoons are usually buried 0 . 5 - 1 m below ground surface .
steyn dg . a bushman arrow poison ( diamphidia nigro - ornata stal . ) . s afr med j 1957 ; 31 ( 6 ) : 119 - 20 urltoken accessed july 9 , 2018 .
steyn dg . ( 1957 ) . a bushman arrow poison ( diamphidia nigro - ornata stal . ) . south african medical journal = suid - afrikaanse tydskrif vir geneeskunde , 31 , pp . 119 - 20 .
csc observed that the ‘beetle paste’ of diamphidia nigroornata larva is applied with a twig to the dried sinew that fastens the arrowhead to the wooden shaft ; the hunter never touches the poison mixture . the arrows are then propped up against a log or hung up to air dry , and stored in a quiver made of the bark of the root of acacia luedertizii engl . , false umbrella thorn ( fabaceae ) ( leffers 2003 ) . finally , the hunter cleans his hands with loose sand . different san groups squeeze diamphidia beetle tissue directly onto the shaft of the arrow ( see photographs in campbell 1968a and chaboo et al . 2007 ) . hunters indicated that cocoons or prepared poison arrows may be traded with hunters in other communities .
thomas ( 1959 : 94–97 ) described the arrow poison preparation of people she terms the gwikwe ( = g | ui ) . the poison was derived from grubs , extracted from underground cocoons ; she drew attention to the similarity of the life cycle with that of diamphidia simplex ( = diamphidia nigroornata ) , as used by the ju | ’hoansi . she described two different colored pupae in these cocoons , which the gwikwe regarded as male ( a small , yellow , black - headed pupa with poison only in the legs ) and female ( a larger orange pupa with poison throughout the body ) . she may have been observing different stages of maturity of the 4 th instar larva of the chrysomelid and the lebistina parasitoid . this poison was applied directly to the arrow shaft .
chaboo cs , grobbewaar e , larsen a , fecaw ecowogy in leaf beetwes : novew records in de african arrow - poison beetwes , diamphidia gerstaecker and powycwada chevrowat ( chrysomewidae : gawerucinae ) ( 2007 ) the coweopterists buwwetin : vow . 61 , no . 2 pp . 297–309
steyn dg :
a bushman arrow poison ( diamphidia nigro - ornata stal . ) .
south african medical journal = suid - afrikaanse tydskrif vir geneeskunde , vol . 31 , no . 6 , 1957 , pp . 119 - 20 , urltoken accessed july 9 , 2018 .
the various groups of hunters have used a variety of poisons . some use plant - derived substances or toxins derived from poisonous snakes . however , in the northern kalahari , the most commonly used poisonous substance for arrows is that derived from the larva and pupae of chrysomelid beetles in the genus diamphidia .
three feet . three feet of red sand and clay the hunter - gatherer bushmen of the kalahari desert dig through to find the poison which they use to fall their prey . do they search for the venom of snakes , spiders , or scorpions ? no , they dig for the larvae of a lowly leaf beetle . beetles of the genus diamphidia lay …
the warvae and pupae of diamphidia produce a toxin used by bushmen as an arrow poison . [ 1 ] [ 2 ] the finnish expworer hendrik jacob wikar , who travewwed in soudern africa in 1773–1779 , described de warvae as
poisonous worms
. hans schinz was de first scientist to document de process by which de bushmen extract and use de poison , uh - hah - hah - hah .
it is interesting that the toxin from these beetle larvae / pupae ( called diamphidia toxin ) in non - toxic to mammals by ingestion and is only toxic after it enters the blood stream . this is puzzling because it then raises the question as to how this toxin provides protection to the beetle larva from predators and parasitoids . perhaps this non - toxicity by ingestion does not apply to non - mammals .
in blepharida , diamphidia , podontia , and polyclada , larvae retain their feces directly on the dorsum . this coating acts as a deterrent to attacking enemies such as ants ( vencl and morton 1998 , 1999 ) . the fecal coat may also serve to moderate body temperature or to reduce water loss but the functions have not been tested . fecal retention and the dorsally - positioned anus represent complex characters supporting the monophyly of the blepharida - group ( paterson 1943 ) .
ty - jour t1 - a bushman arrow poison ( diamphidia nigro - ornata stal . ) . a1 - steyn , d g , py - 1957 / 2 / 9 / pubmed py - 1957 / 2 / 9 / medline py - 1957 / 2 / 9 / entrez kw - poisons sp - 119 ep - 20 jf - south african medical journal = suid - afrikaanse tydskrif vir geneeskunde jo - s . afr . med . j . vl - 31 is - 6 sn - 0038 - 2469 ur - urltoken er -
san ( ' bushman ' ) hunters use lightly constructed bows and arrows in their hunting which means that the injury inflicted by the arrow is usually insufficient to subdue large prey unless it is supplemented by a poison that has been applied to the arrow . a variety of poisons have been used by the various groups of hunters . some use plant - derived substances or toxins derived from poisonous snakes . however , in the northern kalahari , the most commonly used poisonous substance for arrows is that derived from the larva and pupae of chrysomelid beetles in the genus diamphidia .
the ǂ x’ao - ǁ’ aen ( = makaukau , auen ) , botswana and namibia . the ǂx’ao - ǁ’aen san ( ~ 7 , 000 people ) are sometimes called makaukau or auen ( schapera 1930 : 33 ) . they are found in the northern and southern parts of ghanzi district ( groot laagte , kuke , d’kar , hanahai ) , botswana and into the omaheke region , eastern namibia ( otjinene , skoonheld , donkerbos , gobabis ) . the ǂx’ao - ǁ’aen use diamphidia beetles for arrow poison ( maria sapignoli , job morris , pers . comms . , 2012 , 2013 , 2014 ) .
diamphotoxin , de poisonous compound in diamphidia , is highwy wabiwe , and has a wow mowecuwar weight . it is bound to a protein dat protects it from deactivation , uh - hah - hah - hah . [ 2 ] [ 4 ] it causes an increased permeabiwity of ceww membranes , which , whiwe not affecting normaw ionic fwow between cewws , awwows easy access to aww smaww ions , dereby fatawwy disrupting normaw cewwuwar ionic wevews . [ 5 ] awdough it has no neurotoxic effect , it produces a wedaw haemowytic effect , and may reduce haemogwobin wevews by as much as 75 % , weading to haemogwobinuria . [ 6 ]
habitus of adults of blepharida - groupgenera , size < 2 cm long . 1 . asiophrida marmorea ( wiedemann ) ( photo by c . - w . shin ) . 2 . blepharida rhois ( forster ) ( photo by c . - w . shin ) . 3 . blepharida vittata baly ( photo by c . - w . shin ) . 4 . crimissa cruralis stål ( photo by m . tavares ) . 5 . diamphidia femoralis gerstaecker ( photo by c . s . chaboo ) . 6 . ophrida spectabilis ( baly ) ( photo by c . - w . shin ) . 7 . podontia affinis ( gröndal ) ( photo by c . - w . shin ) . 8 . podontia lutea ( olivier ) ( photo by c . - f . lee ) . 9 . podontia rufocastanea baly ( photo by c . - w . shin ) . 10 . polyclada flexuosa baly ( photo by c . s . chaboo ) .
each cocoon usually contains a larva in an arrested stage of development prior to pupation . larvae can stay in this dormant stage for several years .
a san hunter squeezing the haemolymph of a beetle larva on to an arrow . up to 10 larvae may be applied to one arrow . the poison is applied to the foreshaft behind the point and not to the point itself , evidently to avoid accidental poisoning to the hunter from being scratched by the arrow point . the poison is a protein and becomes less potent with time but generally can retain its toxicity for up to a year .
squeezing the contents of larvae / pupae directly on to the arrows ( see above photos ) . this might be followed by drying the arrows over a fire .
making a mixture of beetle larvae / pupae plus plant juice plus spit and then applying it to the arrows . the plant juice acts as an adhesive .
drying the larvae / pupae in the sun , grinding them into powder and then mixing the powder with plant juices . this mixture is then applied to the arrows .
after shooting an animal with an arrow , the hunters then follow its spoor until the poison subdues it which can be within a few hours but which can take 4 - 5 days in the case of large prey such as giraffe . a wounded animal can evidently travel 40 - 70 miles ( 64 - 112 km ) before slowing down and finally falling down dead . noli ( 1993 ) comments on the fact that this system of hunting animals is very time consuming and in fact would be much more efficient if more powerful bows with more injuring arrows were used . there are various reasons why they might not have adopted more powerful bows including the possibility that hunting is more than just about survival but also about a culture and sport with rules and status .
koch , c . 1958 . preliminary notes on the coleopterological aspect of the arrow poison of the bushmen . pamphlet of the south african biological society 20 : 49 - 54 .
noli , h . d . 1993 . a technical investigation into the material evidence for archery in the archaeological and ethnographical record in southern africa . unpublished ph . d . thesis , university of cape town , 252 pages .
shaw , e . m . , woolley , p . l . & rae , f . a . 1963 . bushman arrow poisons . cimbebasia 7 ( 2 ) : 2 - 41 .
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some species are used by san ( ' bushmen ' ) as sources of poison for their arrows .
adult . feed on the host plant by day and if disturbed they quickly drop to the ground or fly off .
eggs are elongate , ovate with an orange tint and are laid on the branches of commiphora plants ( right ) in batches of up to 15 eggs each , closely attached to one another . after laying a batch of eggs , the female covers it with greenish excrement , using her back legs to build up the deposit . on drying , this excrement becomes dark - brown in colour and becomes well camouflaged against the branch to which it is attached .
larvae pass through three instars ( i . e . stages : each instar is separated by moulting of the cuticle ) . they feed on the leaves of the host plant .
upon completion of their development on the host plant , the larvae burrow down 0 . 5 - 1 m into the sandy soil in close proximity to the host plant bush . they then makes a cocoon round themselves from sand , and secretions from the anal region . the larvae remain curled up in the cocoon for up to several years before finally moulting into pupae . this variable period of arrested development explains why san hunters are able to find cocoons throughout the year , for using as poison on their arrows .
[ all photographs by hg robertson , iziko © . photographed in ' bushmanland ' , ne namibia , in february 1996 ]
read about how san hunters use the cocoons as a source of poison for their arrows .
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beetle and plant arrow poisons of the ju | ' hoan and hai | | om san peoples of namibia ( insecta , coleoptera , chrysomelidae ; plantae , anacardiaceae , apocynaceae , burseraceae ) .
arrow poisons in south asia . part 1 . arrow poisons in ancient india .
[ legal evidence by adi poison and the arrow poison of natives of dahomey ] .
caroline s . chaboo , 1 megan biesele , 2 robert k . hitchcock , 3 and andrea weeks 4
1 division of entomology , biodiversity institute and department of ecology and evolutionary biology , 1501 crestline drive , suite 140 , university of kansas , lawrence , ks , 66045 , u . s . a .
2 kalahari peoples fund , 4811 - b shoalwood , austin , tx , 78756 , u . s . a .
3 department of anthropology , university of new mexico , albuquerque , nm , 87131 , u . s . a .
4 ted r . bradley herbarium and department of biology , george mason university , fairfax , va , 22030 , u . s . a .
copyright caroline s . chaboo , megan biesele , robert k . hitchcock , andrea weeks
this is an open access article distributed under the terms of the creative commons attribution license ( cc by 4 . 0 ) , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
this paper concerns the arrow poisons used by the southern african san . the term san ( also known as khoisan , basarwa , or “bushmen” ) covers many indigenous groups using so - called ‘click languages’ in southern africa (
; see our materials and methods below for further notes on san nations ) . in africa , the origin of projectile points currently is dated to ~ 64 , 000–60 , 000 years ago , with the oldest from sibudu cave , south africa (
) . poison arrows are discussed in ancient mythologies ( e . g . , odysseus’ hellebore - poisoned darts in the iliad ,
) and in ancient religious texts ( e . g . , the rig veda from india ( 1100–1700 bce ) , see
map showing contemporary distribution of major san groups in southern africa ( prepared by marieka brouwer burg ) .
the most commonly used poisons across cultures are extracts from single plants or mixtures of plants ( neuwinger 1996 , 1998 ) . south american indians in the orinoco basin use curare , extracted from plants ( loganiaceae : strychnos l . ; menispermaceae : chondrodendron ruiz et pavón , sciadotenia miers ( wintersteiner and dutcher 1943 ) . poison dart frogs ( anura : dendrobatidae ) are used by the indigenous chocó in colombia ( myers et al . 1978 ) ; the frogs secure poison from their insect , mite and millipede prey ( clark et al . 2005 ; daly et al . 2005 ) .
) . hunting with poison arrows continues in southern africa today among a number of different san groups ( fig .
) . some san groups still practice limited hunting and gathering , depending on their living conditions , population densities , technology , and the legal context in which they are operating . san use several hunting techniques : pursuit hunting on foot with bows and arrows , spears , or clubs ; running down game animals and then dispatching them ( “persistence hunting” ; see
, 2007 ) ; hunting from ambush , sometimes with bows and arrows or spears ; mounted hunting , usually on horses but sometimes on donkeys , driving animals toward waiting hunters ; and hunting with dogs .
provides a comprehensive review of archery in southern africa . hunting with guns is rare among most san . in general , the san hunt small game with traps and snares and use poison arrows on large game—antelope , buffalo , cheetah , eland , elephant , gazelle , giraffe , impala , lion , puku , springbok , warthog , wildebeest , and zebra (
( baly ) , used for bushmen arrow poisons ; he noted geographic variation in poison formulae and that only the kalahari bushmen used the beetles . john
: 378 ) reported , “there are four kinds of poison—a root which is rarely used , two grubs , and the pod of a tree . one of the two kinds of grubs is the larva of an unidentified beetle that lives in a bush ; the other is the larva of
complex…lastly , still a fourth insect—small and hairy… . ” this third beetle has been identified as
) whose larva is a parasitoid , killing off the host chrysomelid larva as it develops .
indicated the involvement of multiple insect species belonging to various genera and families ; he wrote that “results of all previous knowledge . . . . . will have to be revised” ( p 53 ) . during the last 200 years , various sources of poison have been implicated ( table
summary of southern african san groups using poisons on hunting arrows and the source of the poison .
de la harpe et al . 1983 ; woollard et al . 1984 ; woollard 1986 ; kann 1989
koch 1958 ; heinz 1966 , 1973 ; steyn 1971 ; heinz and maguire 1974 ; main 1987 ; woollard et al . 1984 ; woollard 1986
plants : eythrophylaeum guineense g . don ( caesalpiniaceae ) , palisota barteri hook ( commelinaceae ) combretum sp . ( combretaceae )
marshall thomas 1959 ; gakelebone , pers . comm . ; sesana pers . comm .
campbell 1968a , b ; nonaka 1996 ; gakelebone , pers . comm . ; sesana pers . comm .
böhm 1897 ; marshall 1958a , 1976 ; lee 1979 ; mebs et al . 1982 ; de la harpe et al . 1983 ; green 1998 ; chaboo 2011 ; robbins et al . 2012
bleek 1928 ; steyn 1971 ; guenther 1986 ; alan barnard , mathias guenther , maria sapignoli , pers . comm .
tobias 1957 ; alan barnard , mathias guenther , maria sapignoli , pers . comm .
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 unclear . several factors contribute misconceptions , outright errors , and ambiguous information about san arrow poisons . first , the use of the term “bushmen” for diverse san tribes 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 .
in this paper , we synthesize the anthropological , entomological , and chemical literature about san arrow poisons . based on our fieldwork , we report arrow poison sources , their preparation and use , bow and arrow construction , and poison antidotes for the ju | ’hoan san in north east namibia and the hai | | om san at etosha national park , namibia . these two ethnic groups represent the largest san groupings in namibia (
) . we also summarize what is known about beetle poison use in seven other san groups—the g | ui , g | | ana , g | | olo , kua , naro , tsila , and x’ao - ǁ’aen . we note arrow poison by one non - san group—valley bisa in zambia (
) . our paper supplements ethno - entomological documentation of the g | ui and g | | ana in and around the central kalahari game reserve ( see fig .
we synthesize literature from anthropology , botany , chemistry , and entomology to develop a better picture of the arrow poisons used by the san in southern africa . we present novel data and images ( figs
: 213–219 for hai | | om ) . individual data is indicated by author initials [ i . e . , caroline s . chaboo ( csc ) , andrea weeks ( aw ) , megan biesele ( mb ) , and rk hitchcock ( rkh ) ] . additional collecting of
) , was done by csc and a colleague , e . grobbelaar ( agricultural research council , plant protection research institute , biosystematics division ) , in south africa .
8 sclerocarya birrea ( anacardiaceae ) , south africa ( photos : cs chaboo ) 9 commiphora africana ( burseraceae ) , south africa , with sanseviera ( dracaenaceae ) at base ( photo : e grobbelaar ) 10 adenium bohemianum ( apocynaceae ) used as arrow - poison by hai | | om around etosha national park , namibia .
symbols in san languages . the san languages reported on here are characterized by sounds that are called clicks . this article is concerned primarily with the ju | ’hoansi and the hai | | om ( see lee 2013 ) and the un - sponsored site , written in the sand , 2015 ) . they have four clicks in their languages ; the symbols for these clicks used throughout this text are as follows :
/ / ( | | ) lateral click ( as in g | | ana ) .
san words for aspects of the poisons and their preparation are available in linguistic dictionaries ( see tanaka 1978 ; dickens 1994 ; visser 2003 ) .
data collection . in addition to the ju | ’hoansi , our literature survey revealed beetle poison use by seven other san groups—g | ui , g | | ana , g | | olo , kua , naro , tsila , and ǂx’ao - | | ’aen . we summarize these data below . insect inventories were conducted by csc in expeditions to botswana , namibia , and south africa ( 2005–2007 ) , including a field study for three weeks with hai | | om and ju | ’hoan communities in namibia in october 2007 . hitchcock worked among the ju | ’hoansi in 1987–2015 and hai | | om in 2011–2012 . biesele has extensive experience among the ju | ’hoansi from 1970 to the present .
throughout this manuscript , we refer to the kalahari . the kalahari desert is a basin - shaped plateau extending over northern south africa , from eastern namibia , across most of botswana to southwest zambia , and southern angola (
) . our data is drawn from our experiences with different communities in different parts of this vast region and our map ( fig .
) illustrates contemporary distributions of major san groups that have been studied in southern africa . in this paper we focus mainly on the ju | ’hoansi and the hai | | om . the former are found in both namibia ( tsumkwe district ) and botswana ( north west district , ngamiland ) , and the latter are found in odangwa , cunene , and otjozondjupa regions . some groups are not shown because of the history of forced removals of people from their ancestral lands and the small population sizes . an accurate socio - linguistic map of historic and contemporary distributions of southern african peoples is fraught with problems but will be developed as our research expands .
the environmental settings of nyae nyae ( namibia ) and tsodilo - dobe - / xai / xai ( botswana ) is kalahari sandveld and consists of tree - bush savanna and pans where water collects in the rainy season ( yellen and lee 1976 ; marshall 1976 : 82–91 ; lee 1979 : 87–115 ; thomas and shaw 2010 ) . the climate of the area is semi - arid , with a 4–6 month summer rainy season and moderate to cool winters with no rainfall ( yellen and lee 1976 ; thomas and shaw 2010 ) . rainfall at tsumkwe averages 450 mm per annum ( range 219 . 8–627 . 8 mm ; namibia meteorological service data for 1984–2010 ) . one of the unique features of this region is that there are between 15 and 25 permanent waterholes in the molapos ( lowland areas ) that lie between east - west trending sand dunes . some of these waterholes are pans that are fed either by springs or by rainfall ( marshall 1976 : 64 ; yellen and lee 1976 : 36 ) . as yellen ( 1977 : 21–22 , 30–31 ) notes , the pattern in the distribution of dunes and molapos or mekgacha ( low depressions between dunes ) is important , as it provides different resource areas within relatively short distances that can be exploited by resident san populations .
interviews with hai | | om , namibia . since 1954 the hai | | om have not been permitted to hunt in etosha national park , their original designated homeland . subsequent displacement , settlement and shifting away from traditional ways have the consequence that hunters are rare and difficult to locate . seven senior former hunters were interviewed within etosha national park , or on private farms south of etosha , between 18–22 october 2007 : jan tsumeb , daniel apia , katison khomob , willem dauxab , fritz khamuάb , abram geesep , and jakes kamaxάb . interviews lasted several hours and involved a seated conversation and walks to collect specimens . hunting nowadays is illegal ; most informants said they were unaware of any active traditional hunters , but recalled their own hunting days or that of their fathers and uncles . three elderly male hai | | om informants discussed their own hunting in their younger days , and pointed out traditional plants within the park . no poisons or arrows were prepared during the hai | | om interviews ( this would have been illegal ) .
interviews with ju | ’hoansi , namibia . the ju | ’hoan communities together form a politically self - regulated body in the nyae nyae conservancy . the region is part of the savanna biome , and is ecologically arid with brush or grass , almost no trees , and ranges from flat to gently hilly ( mendelsohn et al . 2009 ) . temperatures can be below freezing during winter and over 100°f in summer . the ju | ’hoansi live in small grass huts or rondavels ( round adobe constructions with thatched roofs ) . communities comprise 10–100 people living semi - permanently in some 36 villages around or near boreholes used for water . they are scattered throughout the area within one or more days of walking , the most common form of travel regionally . today , men are still hunters where allowed and women are still the primary gatherers but more commonly both men and women carry out subsistence farming of maize , melons , beans , pumpkins , and other crops and some care for domestic animals and poultry .
the three - person field team , led by csc , traveled to tsumkwe ( 19°35’34 . 99 s , 20°30’07 . 99 e ) in 2007 , and then spent two weeks travelling to 10 villages scattered in the nyae nyae conservancy . in each village , current and former hunters were interviewed , with questions presented by csc in english and translated by team members ( in german , afrikaans , and oshiwambo ) or by native san speakers ( three cases ) . hunters are traditionally male ; we did not encounter or hear of female hunters . within 30 minutes of our arrival , most community members had surrounded our group , inserting comments from time to time , the senior women in particular correcting or debating details .
twelve hunters were interviewed in nyae nyae conservancy : leon ‡oma tsamkao , tsumkwe lodge ; trechie ( ‡nlundi village , aha hills , 9–10 oct 2007 ) ; andreas ( n | ama village , 11 oct 2007 ) ; xushe sao ( n | ama village , 11 oct 2007 ) ; david sao iui ( n | ama village , 11 oct 2007 ) ; / aice n ! aucu ( xamsa village , 11 oct 2007 ) ; | aice ‡oma ( tambuti village , 11 . oct . 2007 ) ; g | aq’o ‡oma ( tambuti village , 11 . oct . 2007 ) ; n ! aici kaqece ( makuri village , 12 . oct . 2007 ) ; kaqece ikaece ( makuri village , 12 . oct . 2007 ) ; il’ao n‡ao ( tsumkwe lodge , 14 . oct . 2007 ) ; and n ! aici ‡oma ( tsumkwe lodge , 14 . oct . 2007 ) . interviews were conducted over one or two days . each involved an initial interview , followed by a walk in the desert with the informants to look for the host plants and beetles ; digging up beetle larvae ( fig .
) ; and complete the interview . each poison and arrow preparation session produced 10–20 arrows [ > 150 arrows observed in total ] . some reports indicate the ju | ’hoan word for the poison beetle is “nga” or “n’gwa” (
author mb has worked among the ju | ’hoansi since 1970 , and served as director of the nyae nyae development foundation of namibia from 1988–1992 . she and rkh collected data about hunting and the use of poison from 46 people in 1987 , 1992 , 1995 , 2001 , and 2011–2014 .
interviews with ju | ’hoansi , bostwana . rkh collected data from 56 interviews in dobe , / xai / xai , tsodilo in the years 1976 , 1978–1982 , 1985 , 1988 , 1992 , 1995 , 1997 , 1999–2000 , 2005 , and 2011–2013 . hunters’ names are omitted here to protect their identities .
interviews with g | ui , g | | ana , kua , and tsila , bostwana . rkh has conducted > 140 interviews in about 15 visits ( 1976–2013 ) with these communities from the central and eastern kalahari , many specifically on hunting . these data were collected in the central kalahari game reserve or in the resettlement sites outside the reserve .
taxonomic identifications . due to the diversity of scientific names for plants and beetles used herein , our generic names are abbreviated to avoid confusion ( aca . = acacia ; ade . = adenium ; aco . = acocanthera ; bob . = bobgunnia ; comm . = commiphora ; euph . = euphorbia ; sans . = sansevieria ; scl . = sclerocarya ) .
. they confirmed the plant from photographs in our regional plant guides or by pointing out planted specimens in nearby gardens , including labeled plants in the educational garden of the research unit , etosha national park . several members of the
csc obtained beetle specimens in several ways : receiving gifts of cocoons from informants from their stored supplies , collecting them with informants on bush walks , and purchasing containers from hunters’ stored supplies ( e . g . , fig .
) . more than 5 , 000 cocoons were accumulated by csc during fieldwork in namibia ; she also conducted nightly light - trapping to sample flying adults . five hundred cocoons were dissected in the field to determine the beetle species and their life stages .
beetle species were identified by examination of types and as part of a taxonomic review of specimens from museum collections in france , germany , south africa , and the usa . the chrysomelid beetle species in the namibian ju | ’hoan arrow - poison case is
dissected cocoons revealed mostly larvae , some pupae , and some adults of leaf beetles . about 5 % of the coccoons held only mature parasitoid carabid larvae (
beetle vouchers are deposited in the collections of the national museum of namibia ( nmwn , windhoek ) , the american museum of natural history ( amnh , nyc ) , agricultural research council , plant protection research institute , south african national collection of insects ( arc - ppri , sanc , south africa ) , and the university of kansas entomology collection ( semc , ks , usa ) . john irish , national botanical research institute , namibia , confirmed the identity of the plants .
our cross - disciplinary synthesis of historial literature , reports of anthropologists , and our own collective fieldwork in southern africa indicates that beetle arrow poison is used by seven san groups—the g | ui , g | | ana , g | | olo , naro , kua , and tsila in botswana , and the hai | | om in namibia . two san groups , the shua and tshwa from the north - eastern kalahari of botswana and zimbabwe , do not use arrow poisons . one san group , the hai | | om , uses a plant poison . the valley bisa in zambia is a non - san group that uses beetle arrow poison . our research focused primarily on two large san groups , hai | | om and ju | ’hoansi , in namibia but we assemble data for other smaller communities .
: 22–36 ) used the generic term kua to refer to g | ui , g | | ana , g | | olo , tsila , and all of the groups in the central kalahari region ( our fig .
map does not indicate a location of the g | | olo as we have not found maps with this information ) . we follow the usage of
who restrict the term kua to the people of the eastern and southeast parts of the central kalahari . since these diverse san nations are so poorly known , we briefly summarize the location , contemporary status , and knowledge of arrow poisons below .
) . they live nowadays in the central kalahari game reserve , botswana . both groups have been the subject of forced evictions after the botswana government removed them from the central kalahari in 1997 and in 2002 (
) . some do still hunt with the aid of bows and poisoned arrows , although this has decreased significantly due to governmental restrictions .
) for their arrow poison , but it is not clear how she identified the beetle species .
in the central kalahari , according to our g | ui and g | | ana informants , the beetle that is used for arrow poison is polyclada flexuosa ( baly ) . the larvae and adults feed on the leaves of marula trees ( sclerocarya birrea ) ( jumanda gakelebone , roy sesana , pers . comms . 2011–2013 ) ; pupation is likely to be in the soil around the host as in other species we have studied .
described kua poison preparation , with 8–12 grubs squashed together with saliva on a mortar , then smeared unto the binding behind the arrowhead . the arrow was then air - dried .
host , but specified the rubbing of larval tissue directly onto the arrow shaft , which was then heated to dry . observations of kua in the eastern - kalahari in 1975–1976 , by hitchcock and ebert , revealed that they made poison from beetles and from ‘spider’s nests’ ( hitchcock and ebert , field data ) ; they assumed that the beetle identification was
. vierich ( pers . comm . 2014 ) indicates that alternative plants were used when beetles were unavailable , however she did not confirm the identity of these plants .
in the east - central kalahari , arrow poisons were used until the late 20 th century . james chapman , who visited nkawane in this area in 1852 , notes that the bushmen there used bows , arrows and spears ( chapman 1971 ) . he noted that the bushmen killed elephants with spears , but he does not say whether or not poisons were used in elephant hunting . based on observations made in 1850 in the eastern kalahari , schulz and hammar ( 1897 : 84 ) indicated that bushmen used arrow poison “obtained from the juice of poisonous herbs and roots” and poisonous snakes .
the tshwa and shua , botswana and zimbabwe . ethno - historic evidence suggests that poisoning of arrows is uncommon among these san groups , living in the northeastern kalahari . hodson ( 1912 : 227 ) made the following observation “the bushmen in this district are called mashuakue , their headman being kotama . . . they do not use the bow and poison arrow so common with bushmen in the far kalahari , but carry long assegaais [ light spear ] , with which they stalk game . some have rifles and are good shots at a close distance . ” detailed fieldwork with tshwa and shua ( about 4 , 000 people ) , beginning in 1975 , revealed that spears were the most common weapon used , along with clubs and guns of various kinds ( crowell and hitchcock 1978 ; hitchcock and bleed 1997 ; hitchcock et al . 2014 ) . tshwa who were forcibly moved to the east - central kalahari in the 1950’s by bamangwato cattle owners , picked up the use of bows and poisoned arrows from the neighboring kua ( hitchcock 1978 ) .
the valley bisa , zambia . marks reported on his 1973 observations of valley bisa hunts in zambia ( 1977 , pers . comm . ) . hunters stalked up to large game before letting an arrow fly , a distance that was further than that for shotguns and even rifles . their tactic was to stay slightly beyond the species ‘flight distance’ ( distance before the prey flees ) , force the herd into a smaller space , and then arc the arrow to fall within the anticipated space . they did aim to get closer to some individual mammals ( e . g . , warthog ) . the arrows only had to lance the animal to get poison into the blood stream , but the wounded prey normally did not move very far away . marks found that hunters used two different types of arrows for mammals and for birds , both poisoned with an extract made of pods and roots from an acokanthera sp . ( apocynaceae ; identified by biologists at university of zambia ; marks 1977 ) .
the ju | ’hoansi in namibia . the ju | ’hoansi represent one of the earliest - diverging lineages of modern humans ( knight et al . 2003 ; tishkoff et al . 2009 ; mitchell 2010a ; pickrell et al . 2012 ; schlebush et al . 2012 ) and are therefore some of the best - documented peoples on the planet as they have been studied intensively and over a long period , since the early 1950s ( marshall and ritchie 1984 ; marshall 2003 ; marshall 1976 : lee and devore 1968 ; lee 1968 , 1978 , 2013 ) . although they may not be the only , or the best model , of human’s hunting and gathering past , in some ways their history is very enlightening . most ju | ’hoansi retain their language , culture , and many of their traditions till today . they have been able to secure some of their land and resources through advocacy , working with the namibian government and non - governmental organizations ( biesele and hitchcock 2011 ; lee 2013 ) . the ju | ’hoansi are the only indigenous people in africa who still have the right to hunt for subsistence purposes using their traditional weapons . some ju | ’hoansi continue to hunt with poisoned arrows , and they attempt to teach their young about finding poison materials , preparing the poison , and putting it on arrows . the ju | ’hoansi are , therefore , well - placed for the investigation of the use of arrow poisons .
hunts can last several days depending on the animal’s size and the slow paralysis by the poison . the tracks and spoor of the fleeing animal helps the hunter decide to immediately start stalking or return to the community to gather materials ( e . g . , water , food ) and other men to help with the hunt . animal tracks and dung inform the hunter about the size ( size of footprint ) , species ( nature of print ) , age ( depth of foot print ) , wound ( one side of foot prints heavier than other size ) , and travelling direction of the prey ( campbell 1968a ; liebenberg 2001 ; lee 2003 ) .
- group . blepharidines typically have large colorful adults ( ~ 1 cm ) ; eggs are deposited in clusters on host plants , coated with fecal material ; larvae that feed on host plant leaves , retain a fecal coat and eventually descend to the ground where they form a sandy underground cocoon ; here they remain in a suspended larval stage , ready for pupation . the cycle from egg to adult can span 2–4 years ; the underground phase is prolonged as a probable adaptation to unpredictable rainfall . blepharidines tend to show generic - level specialization on certain plant families—
arrow poison of the ju | ’hoansi , nyae nyae , namibia . locating host plants .
shrubs are located , the hunter initiates a new hole at the leaf - drop ( = drip line ) margin of the shrub ; in some cases , ditches of previous digs ( past years ) were still apparent and our informant jumped into the 1m deep ditch and extended the ditch to encircle the plant . we observed some ditches forming a complete moat around plants . ju | ’hoan traditionally use a wooden digging stick ( e . g . , of the widespread kalahari christmas tree ,
: 5 ) reported that ju | ’hoan hunters “take measures to protect marula trees by building fire breaks around them during the dry season” and that the indigenous word for the larva is “ ! oan / aqro” .
) , but csc found none of these trees in the nyae nyae conservancy .
) . when sufficient cocoons have been collected into an ostrich egg shell ( fig .
what follows is our typical observation compiled after 12 interviews with hunters who each made their poison with our observation . first , he arranges his tools , stabilizing an old giraffe or kudu knuckle bone with the concave surface facing upwards in the sand in front of him and placing the beetle cocoons nearby . a small fire is lit ; traditionally a fire stick was used , usually made of
) , but nowdays a cigarette lighter is used . he breaks open a cocoon and taps out the single larva ; non - larval forms ( adults , pupae ) are discarded . the larva is then rolled between his fingers , loosening the inner tissues from the integument . using a stick as a pestle , he rubs hard against the skin to loosen tissue , then extracts it to mix on the bone mortar ( fig .
) ; about 10 larvae are used per arrow . he chews the bark of
) to produce saliva which is mixed with the larval tissue and hemolymph . published accounts of similarly chewed extracts list the plants used as :
) is heated over the fire , cooled , and added to the poison mixture . an unidentified toxic bean ( j .
) appear as ingredients in some poison recipes . ju | ’hoan informants at | xai | xai in botswana told hitchcock that they use the juice of
literature sources reported that saliva made by chewing the bark of dicerocaryum eriocarpum ( decne . ) abels ( devil’s thorn ) ( pedaliaceae ) or the leaf of sansevieria aethiopica thunb . ( asparagaceae ) , are used to moisten the poison if it dries out ( leffers 2003 : 87 ) . the efficacy of the poison has been reported to last from three months ( green 1998 ) to two years ( clark 1975 ) . lee ( 1979 : 137 ) reported that the high initial potency declines over time , and is essentially harmless after a year . others report it to decline seasonally ( hitchcock et al . 2015 ) .


cotana bisecta is a moth in the eupterotidae family .

it was described by rothschild in 1917 .

it is found in new guinea .

the wingspan is about 52 mm .

the forewings are dark brown , somewhat suffused with chestnut-rufous and with a median black band , a postmedian indistinct blackish band beyond which is a lunate buffish band on the outside of which is a line of black dots joined by a chain of indistinct blackish lunules .

the hindwings are orange-yellow with an antemedian transverse line of sooty black and a median indistinct blackish cloud band followed by two lunulate bands .

cotana bisecta

cotana neurina turner , 1922 ( eupterotidae : eupterotinae ) , female - qld , 9 km ene of mt tozer , 5 . jul . 1986 , j . c . cardale leg . ( anic ) .
cotana neurina turner , 1922 ( eupterotidae : eupterotinae ) , male - qld , iron range , 29 . may . 1978 , m . s . & b . j . moulds leg . ( anic ) .
cotana neurina turner , 1922 ( eupterotidae : eupterotinae ) , male - qld , bamaga , 30 . mar . 1964 , i . f . b . common m . s . upton leg . ( anic ) .
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lunulata ( bethune - baker , 1904 ) borealis rothschild , 1932 ab . unicolor rothschild , 1932 [ infraspecific ] montium rothschild , 1932 occidentalis rothschild , 1917 satisbona rothschild , 1917
pallidipascia rothschild , 1917 pallidifascia ( sic sensu auct . ) postpallida ( rothschild , 1917 )
rubrescens walker , 1865 ssp . kapaura rothschild , 1917 [ male ] ssp . oetakwensis rothschild , 1917 [ male ]
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
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