1 Historical Introduction TOM PIEK Farmacologisch Laboratorium Universiteit van Amsterdam Amsterdam, The Netherlands Due to the powerful venoms secreted by bees, social wasps and ants these insects appear early in man's history and prehistory. Paleolithic paintings, discovered early in the twentieth century in Spain, show the early interest of humans in honey-bee products (Bodenheimer, 1928). Similar paintings of Mesolithic humans (Fig. 1) have been found in India (Gordon, 1936, 1958). The way those honey hunters nowadays approach bees' nests has been described by Clauss (1983). Skilled honey hunters in Botswana (Africa) approach nests of the aggressive Apis mellifera adansoni with a remarkable gentleness, only using mild smoke of buffalo or cow dung. Paintings made between 8500 and 2000 BC of bees attacking humans and animals have been described by Pager (1976) and Dams (1978). About 1500 BC the Ebers Papyrus deals with prescriptions to prevent wasps from stinging. According to von Buttel-Reepen (1921) the use of smoke to dislodge wasps and bees was already known to the ancient Egyptians. Moses reminded the Israelites of their enemies when he recalled, 'Then the Amorites, who lived in that hill country came out against you and chased you as bees do and beat you down [Deuteronomy 1:44]', and the Israelites were encouraged by the words, 'And I will send hornets before you, which shall drive out Hevite, Canaanite and Hittite from before you [Exodus 23:28]'. Greek sailors may have used bee skeps against Turkish enemies (Gautier, 1974). The properties of social wasp and bee venoms were obviously well known. However, there is a paucity of early records indicating a knowledge of the venoms produced by solitary wasps. A Chinese illustrated encyclopedia called the Erh-ya, in which Bodenheimer (1928) has dated the illustrations as originating between 500 1 VENOMS OF THE HYMENOPTERA Copyright © 1986 by Academic Press Inc. (London) Ltd. All rights of reproduction in any form reserved 2 Tom Piek Fig. 1 Prehistoric painting of humans collecting honey from bees, found in the Mahadea Hills, India. From Gordon (1958). and 400 BC, gives an interesting description of a wasp (kuo-ying-pu-lu) that paralysed a green 'worm' (ming'ling-sang-ch'ung) living in mulberry trees (Fig. 2). According to a German translation by Huelle (Bodenheimer, 1928) the 'worm' was stored by the wasp in a wood-nest. Seven days later the 'worm' was transformed into the son of the wasp. One might doubt whether the original Chinese character concerning the condition of the 'worm' collected by the wasp has been correctly translated into 'paralyse', but it could be argued that if the original author believed in a metamorphosis of the 'worm' into a wasp, he must have been convinced that the 'worm' was alive and not dead. The description in the Erh-ya is probably the most ancient report of paralysis of an insect by a solitary wasp. Social and solitary wasps and bees were also known to the ancient Greeks. The Greek name for the social wasps was Sphex or Sphecio (Moffet, 1634). Today these names are used for genera of solitary wasps in the family Sphecidae, division Aculeata (suborder Apocrita). Aristotle ( — 350 BC) called the solitary wasps Ichneumon and he described them as spider-killing wasps 1. Historical Introduction 3 Fig. 2 A solitary wasp, Kuo-ying-pu-lu, which paralyses an insect larva, Ming-ling-sang-ch'ung. From a facsimile reprint of the Erh-ya yin t'u, or Dictionary of Old Terms with Sung illustration, AD 276-324, in the Royal Library, The Hague. (probably Sceliphron spp.). The name Ichneumon is now used for a genus of the family Ichneumonidae, division Terebrantia (suborder Apocrita). Aristotle also observed a second group of solitary wasps which he called the Pseudosphecae or bastard wasps. This group is now known as the super family 4 Tom Piek Sphecidae or digger wasps. These wasps are described by Oviedus as big, yellow or white coloured wasps with black spots on their wings, which sting flies to death (Jonston, 1653). The wasps described by Oviedus were probably Bembix species. The name Vespae is a Latin name for social wasps, however, the spider- killing solitary wasps are called by Plinius (AD 23-79; Historia Naturalis) Vespae ichneumones. The Latin name for bee is Apis (Steier, 1912). The Indo-European languages, Brythonic and Celtic, give a name to bees which originally described their stinging ability: gwenynen, gwenenen or gwenaneum (Le Sage, 1974). Aristotle (Historia Animalia Lib. V. 21, see Klek and Armbruster, 1919) recognized that honey-bees possess a sting, that the individual he called the 4king' has a sting which it did not use, and that the drones are stingless. Aristotle knew that honey-bees are able to kill animals and recorded that even a horse could be killed (Pseudo-Aristotle IX, 40; see Klek and Armbruster, 1919). About two thousand years separate the original observations of the Erh- ya, and of Aristotle, from those of Butler (1609) and Swammerdam (1669, 1672-1673). In his Feminine Monarchie, Butler (1609) described the defensive mechanisms of honey-bees: Hir speere she is very loth to use, if by any other meanes she can shift hir enimy, as knowing how dangerous it is to hir selfe: for if shee chance therwith to strike any hard part, as the brest or shou'der, she is enforced to leave hir speere behinde her, and so she killeth and is killed with the same stroke. Yet when the Bees are very angry; as namely whe they are assalted with a multitude of robbers at once, they fal sodainly upo the with their poisoned Speeres. Their Speeres or stings they use chiefly against things of other sort, as men, beastes, and fowles, which have outwardly some offensive excremet, as haire or feathers. Hair and feathers cause the Bees to sting. Butler's text (1609) clearly shows that the venom apparatus of honey-bees is normally used against enemies, including vertebrates. This is probably the case for all bees, ants and social wasps. In his work on metamorphoses of insects Swammerdam (1669) described a number of wasps. He also described the sting of bees and wasps extensively (Fig. 3) (Biblia Naturae Sive Historia Insectorum, 1672-1673). In Germany Frisch (1720-1738) described about 300 insects, among which appear spider- and caterpillar-killing wasps. He gives no information on paralysis of the prey by these wasps. The work of Swammerdam was continued by Ferchauld (better known as Monsieur de Reaumur). He described the sting of bees, the life of social Fig. 3 Morphology of the intestine and the sting apparatus of a honey-bee worker (I—III) and the glandular part of the venom organ of a social wasp, probably Vespa spp. (IV). From Swammerdam (1672-1673), Biblia Naturae Sive Historia Insectorum, Tab. XVIII. AjJZ. JFy.T. 6 Tom Piek bees and wasps (Ferchauld, 1740) and the life of the solitary wasps (Ferchauld, 1742). The solitary wasps were divided into the ichneumon flies (des mouches ichneumons), which were probably identical to the Hymenoptera Terebrantia, and the ichneumon wasps (des guespes ichneumons), which were probably identical to the Hymenoptera Aculeata (see end of this chapter for subdivision of the Hymenoptera). He based his division on the ichneumon flies having an 'external sting' and the ichneumon wasps having an 'internal sting'. This distinction had already been recognized by Aristotle. The ichneumon wasps were also distinguished from the social wasps by the fact that members of the former group could not fold their wings. This distinction is no longer useful since the Eumenidae, which are also solitary, do fold their wings. One of the 'ichneumon' wasps described by Riaumur (1742) is probably Odynerus spinipes (cf. Dufour, 1839). He observed the nest of this wasp to be very well supplied with green 'worms', which were obviously close to their metamorphosis. Reaumur (1742) wrote that the 'worms' were stored as rings, and in a deep sleep: he believed that they were still alive. It could be that Reaumur did not realise that the 'worms' were paralysed. We will discuss these green 'worms' again in reviewing the work of the nineteenth century French entomologists. According to Lepeletier (1841), Reaumur (1742) also described an observation by Cossigni of a wasp Ampulex, which stings and paralyses a cockroach, Lepeletier's (1841) citation is, 'La kakerlac a perdu ses forces; eile est tombee en paralysie; eile ne peut resister ä Phymenoptere'. The original text by Reaumur is, 'La kakerlac a alors perdu ses forces, eile est hors d'etat de resister ä la guepe ichneumon'. Reaumur did not say that the prey was paralysed, although he possibly did intend this meaning. The change in behaviour, other than a paralysis, of the cockroach stung by Ampulex compressa is described in Chapter 5. In 1671 the Royal Society of London published a letter from an American, WiUoughby, 'containing some considerable Observations about that kind of Wasps, call'd Vespae Ichneumones; especially their several ways of Breeding and among them, that odd way of laying Eggs in the Bodies of Caterpillars'. Willoughby's letter did not mention paralysis of the prey and neither did Banister in a letter dated 1680, communicated to the Royal Society by James Petiver and published in the Philosophical Transactions of the Royal Society of London (1701). This letter was entitled 'Some Observations Concerning Insects in Virginia' and describes 'Dirt Wasps, making their Nests of Dirt and put into it 6 or 8 live Spiders, leaving them to brood upon their young, something like that of Aristotle in his Historia Animalia, Lib. I. Cap. 20' (see Klek and Armbruster, 1919).