Historical Introduction

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

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). Although Banister did not speak of paralysis, he did not follow Aristotle in saying that the spiders were killed, on the contrary he recognised them to be alive. A letter from Bartram about 'some very curious Wasps Nests made of Clay 1. Historical Introduction 7 in Pensilvania' appeared in the Philosophical Transactions of the Royal Society of London (1744). The spider-hunting wasp discussed in this letter was probably a Sceliphron species. Bartram (1744) wrote, 'But it is further wonderful to observe, that they (the wasps) only in some manner disable the Spiders, but not kill them; which is to answer two Purposes; first that they should not crawl away before the cell is finished and next that they may be preserved alive and fresh until the Egg hatches, which is soon'. Bartram (1749) also described another wasp (Great Black Wasp)* catching grasshoppers, 'But their peculiar Skill is to be admired in disabling them either by Bite or Sting, so as not to kill them; for then they would soon putrify, and be unfit for Nourishment', . . . Life sufficient is left to preserve them for the time the Maggot is to feed on them.' It is obvious that these American observers were convinced that the prey were alive and that Bartram, in particular, because he used the term 'disabling', must have recognised that the prey was paralysed. Leaving the American observers of the seventeenth and eighteenth centuries and returning to Europe, we encounter the work of De Geer (1771), who described a number of solitary wasps; among them was Sphex sabulosa, which immobilized and collected spiders (Taraignee devient immobile'). De Geer also implied that the spiders were insensible (Taraignee etoit fraiche et ne paroissoit avoir aucune blessure; cependant ell'etoit absolument sans mouvement et comme parfaitement engourdie'). Therefore, it is surprising that Dufour (1839), continuing the work of Reaumur and also describing 'green caterpillars rolled up and live, and collected by Odynerus spinipes\ did not realize that these larvae were paralysed. Dufour (1839) included these observations in a letter to the Annales des Sciences Naturelles, in reply to which one of the editors (Audouin, 1839) presented a description of his observations on two groups of the same green 'caterpillars', one group collected by the wasp and the other group collected in the field by Audouin. The latter larvae metamorphosed very soon, the adult insects being weevils. Thus Audouin discovered that the 'worms' of Reaumur and the 'caterpillars' of Dufour were in reality beetle larvae. The group of beetle larvae collected by the wasp did not show any sign of metamorphosis, confirming the conclusions of Reaumur (1742) and Dufour (1839). Because Audouin compared larvae collected by himself with larvae collected by the wasp, he was able to recognize the lethargic state of the latter group of insects. In the same letter Audouin (1839) described another wasp, Odynerus parietum, which attacked true caterpillars. These caterpillars also did not metamorphose. Their bodies could contract but they were not capable of

The drawings of the wasp and grasshopper leave little doubt that the Great Black Wasp represents Ammobia pennsylvanica (Frisch, 1938). 8 Tom Piek locomotion. The larvae collected from the wasps's nest in August 1835 were still alive in June 1836. Audouin (1839) was convinced that the wasps were responsible for what he called paralysis. Two years later Dufour (1841) answered Audouin's letter. Dufour argued that adult beetles collected by Cerceris bupresticida and bees collected by Palarusflavipes were not paralysed because these insects did not respond when manipulated at dissection. According to Dufour this indicated a complete lesion of the central nervous system and not a state of paralysis which is, according to him, characterized by an intact sensible system. Dufour (1841) agreed with Audouin (1839) in believing the wasp's venom to contain a preservative compound. However, Dufour was sure that the beetles and bees were dead. Lepeletier de Saint-Fargeau (1841) described Bembix rostrata stinging a fly, resulting in paralysis ('cette piqüre met ses victimes dans une espece de paralysie, qui n'est mortelle'). He also observed that Philanthus triangulum paralysed workers of honeybees (41 räsulte de la piqüre un engourdissement subit; mais l'abeille conserve la vie une quinzaine de jours'). Passerini (1841) described what he called paralysis in larvae of Oryctes nasicornis caused by a. sting of Scoliaflavifrons(. . . . 'paralizzate, ma ehe movevano lentamente i piedi e i segmenti del corpo'). It fell to Fabre (1855) to settle conclusively this conflict of views. He observed that the weevils attacked by Cerceris tuberculata, and also other prey of solitary wasps, were sometimes partly lethargic. Stimulation of such larvae often resulted in movements. Completely lethargic beetles, stimulated by electric currents, showed muscle contractions and movements. Fabre's conclusion was that prey of these solitary wasps were paralysed and not dead, 'Non en presence de parails faits, on ne peut invoquer Faction d'un antiseptique et croire ä une mort reelle; la vie est encore la, vie latente et passive'. Therefore, it seems to be a contradiction that Fabre (1879-1910), despite the above description by Lepeletier de Saint-Fargeau, believed that honey-bee workers stung by Philanthus triangulum were dead and not paralysed, which is not true (see Chapter 5). After Fabre's publications, a great number of entomologists were to described their findings on the paralysis of prey by solitary wasps. In his Souvenirs Entomologiques Fabre (1879-1910) presented the view that solitary wasps, at least the Aculeata, sting their victims in the central nervous system. He observed prey having a concentrated central nervous system (bupresticid beetles, weevils and some beetle larvae) to be stung once, and that prey with a more diffuse central nervous system were stung more than once. Among his examples were Sphex spp., which sting crickets three times, once in each of the three main nerve ganglia, and Ammophila spp., which 1. Historical Introduction 9 sting caterpillars in every segment containing a major nerve ganglion (Fabre, 1879-1910). Fabre's conclusion, that prey of most solitary wasps is paralysed and not killed, has been criticized by Peckham and Peckham (1898). They suggest that the prey of the solitary Aculeata are normally killed and that wasps that paralyse their prey should be 'in the novitiate state and not entitled to rank as masters in the art'. Current knowledge on the stinging behaviour of solitary wasps and on paralysis of prey (Chapters 4 and 5) allows criticism of the Peckhams' hypothesis and gives support to the overall picture presented by Fabre. We must, however, remember that both Fabre and the Peckhams were pioneers in the description and interpretation of the behaviour of solitary wasps. Their work is part of the excellent inheritance we derive from the nineteenth century entomological literature. Impressed by the powerful actions of the venoms of the social wasps, bees and ants, many entomologists have studied the anatomy of the stings of the insects. Aristotle described the sting of the bees as an internal organ. Swammerdam (1672-1673) was the first to give a detailed description of the chitinous and glandular parts of the venom apparatus of bees and wasps (Fig. 3). He described the sting as consisting of three parts, two of them, the lancets (called by Swammerdam the legs or crura) moving inside the sting sheath (Swammerdam's vagina). The different names used for the chitinous parts of the sting of bees and wasps by Swammerdam and by a number of other workers have been summarised in Table I. The confusion in the terminology of the different authors has been unravelled using their excellent drawings. The embedding technique, discovered by Swammerdam, made possible his extensive studies of the chitinous parts of the sting. These techniques did not preserve the glandular parts of the venom system very well and the studies of Swammerdam and Riaumur on the venom glands lacked the precision of their observations of the chitinous apparatus. Swammerdam described the venom of bees and social wasps as being stored inside a reservoir. In what he called the common wasp and in the hornet this reservoir is surrounded by muscle. He could not find this particular muscle around the reservoir of worker honey-bees (Biblia Naturae Sive Historia Insectorum, T. II, p. 457). Leydig (1859) confirmed that worker honey-bee's venom reservoir walls do not contain muscle fibres. That this is not entirely true has been shown by Bridges (see Chapter 2, Sections II and III). The venom gland and reservoir of Hymenoptera were believed to contain a strong acid (Wray, 1670; Berg, 1865; Carlet, 1890). Referring to the observations of Fisher, of Langham and of Tragus, Wray (1670), described the way in which flowers of chicory, and some other blue flowers, changes their colour to red when brought into contact with disturbed ants. Quoting Table I Comparison of the Names Used for Some of the Different Chitinous Parts of the Sting Apparatus of Bees and Wasps" bulb of sting sheath sting sheath sting bulbe du gorgeret gaines de l'aiguillon aiguillon ^ Crane, 1951 Stachelrinnen Kolben Stachelrinne (Scheide) Stachel bulb of stylet of sting stylet of sting lancets of sting Snodgrass, 1956 koker (vagina) benen (crura) Swammerdam, 1672-1673 la base des auguillons le fourreau des aiguillons* les aiguillons Reaumur, 1740 Rinnenkropf Stachelrinne Stechborste Sollmann, 1863 Schienenrinne Schieber des Stachels Fenger, 1863 Furche der Schienenrinne Leiste der Schienenrinne Stechborste Kraepelin, 1873 gorgeret la tige du gorgeret les stylets d'aiguillon Carlet, 1890

flTop: English, French and German equivalents given by Crane, and the English equivalents used by Snodgrass. Bottom: equivalent names used in history. öIn The Dictionary of Bee-Keeping Terms, with allied scientific names (Crane, 1951, ed.) the French name fourreau de laiguillon is translated as sting chamber or sting cavity (Stacheltasche). 1. Historical Introduction 11

Fisher's work, Wray (1670) wrote that when an ants' nest was stirred with a stick, a liquor was deposited on the stick. Although Fisher did not know 'to what sort of liquor this juyce refer to, it twinges the nose like newly distilled spirit of vitriol'. Wray (1670) was easily induced to think 'that this juyce of Pismires might be of the same nature with the oyl of vitriol'. Fisher found the acid to be very similar to vinegar. Wray found it strange 'that Nature should prepare and separate in the Body of these Insects without any sensible heat and that in a good quantity, considering the bulk of the animal, a liquor the same for kind with those Acid spirits which are by Art extracted out of some Minerals, not without great force of Fire'. The chemical identification of formic acid was performed by Manitius and Sperling (1689; see also Hagen, 1863; Horn and Schekling, 1928) and by Neumann (1730). Huber (1792, see Huber, 1814) described the ability of both ants with a sting and stingless ants to produce formic acid. For the recently discovered pathway of biosynthesis of formic acid see Chapter 9, Fig. 5. Bert (1865) found that the content of the venom reservoir of the carpenter bee Xylocopa violacea was acidic. According to Carlet (1890) the active principle in the venom of other bees and also of wasps should be formic acid. We now know that this is not true (Chapters 5-8). The early authors have only recognized one gland system in their observations of the honey-bee's venom apparatus. This gland consisted of a reservoir with two bifurcated glands (Swammerdam, 1672-1673) or a single tubular gland (Reaumur, 1740). A second gland system was later described in bees and wasps by Dufour (1834). Dufour's gland contains a weak alkaline solution (Carlet, 1884a) and has been commonly referred to as the alkaline gland. Laboulbene (1852) suggested that Hymenoptera with a barbed sting always killed their prey and that hymenopterans with a smooth sting paralysed the prey. Carlet (1884a, 1980) extended Laboulbene's hypothesis by suggesting that the alkaline gland should only be present in Hymenoptera which have a barbed sting (bees and social wasps) and that this gland would be absent in Hymenoptera with a smooth sting (solitary wasps). He explained the paralysis of the prey of solitary wasps by the action of the product of the acid gland and the lethal action of the venom of bees and social wasps by the combined action of the products of the acid and alkaline glands. This view was rejected by Bordas (1894a), who showed that the Hymenoptera he knew possessed at least two glands, the acid gland and the alkaline gland (or Dufour's gland). Bordas (1894b) has also described a third gland in the Terebrantia and in a number of Aculeata. From Bordas's drawings it seems that this third 'venom' gland is present in Sphecidae but not in Pompilidae (Psammocharidae), Vespidae or Apidae. The work of Swammerdam Table II Survey of the Most Important Groups (Suborders, Superfamilies and Families) of Hymenoptera Treated in This Book

SUBORDER [DIVISION] Superfamily Family

SYMPHYTA1 APOCRITA [TEREBRANTIA] Ichneumonoidea Ichneumonidae2 Braconidae3 Cynipoidea Cynipidae4 , Figitidae Chalcidoidea Torymidae,Chalcididae5, Eurytomidae, Pteromalidae Encyrtidae, Eupelmidae, Eulophidae, Elasmidae Proctotrupoidea Platygasteridae [ACULEATA] Bethyloidea Sclerogibbidae, Dryinidae, Bethylidae, Chrysididae Scolioidea Scoliidae6,Typhiidae, Mutillidae Pompiloidea Pompilidae7 Sphecoidea Sphecidae8 Vespoidea Eumenidae9 Vespidae10 Apoidea Colletidae Halictidae Andrenidae Melittidae Megachilidae Apidae11 Formicoidea Formicidae12 1. Historical Introduction 13

(1672-1673), Reaumur (1740), Dufour (1834), Bordas (1894b) and Carlet (1884b, 1885) provides a historical introduction to more recent studies on the morphology of the chitinous and glandular parts of the stinging apparatus of the Hymenoptera (Chapter 2). In the fourth edition of the Systema Naturae the order of the Hymenoptera was established by Linne (1744) under the name Gymnoptera (naked wings). According to Westwood (1840) the Hymenoptera are distinguished as an order from other insects by the number, the comparative size and the structure of the wings, the mandibulated mouth enclosing the labium ensheathed by the maxillae, the ovipositor, or the sting and the nature of their metamorphosis (Westwood, 1840). He suggested the following arrangement of the different groups within the order, founded chiefly upon the ideas of Latreille: section I: Terebrantia, with the subsections Phytiphaga and Entomophaga, and a section II: Aculeata with the subsections Praedones and Mellifera. In the modern literature the Hymenoptera are normally divided into the suborders Symphyta and Apocrita. The Symphyta (or phytophagous Hymenoptera) are scarcely dealt with in this book (Chapter 5). It has long been customary to separate the suborder Apocrita into two main divisions, the Aculeata and the Parasitica or Terebrantia (Imms, 1960). The name Terebrantia is to be preferred, since the use of the name Parasitica suggests that the other division, the Aculeata, does not include parasites*. In this volume only the Apocrita are extensively described, and as a matter of simplicity we divide the Apocrita into two major groups, the Terebrantia, having an ovipositor (terebra or drill) which is also used as a ductus venatus, and the Aculeata, which differ from the first group in having an 'ovipositor' (aculeus or sting) fully modified for injection venom into a host, prey or enemy, though in some primitive families, for example Drynidae and Chrysididae, the ovipositor has retained its egg-laying function (Imms, 1960). A survey of the most important (super)families of Hymenoptera is presented in Table II.

*A parasite is here defined as an organism living in or on another organism (its host) from which it obtains food (Abercrombie et al.y 1951). According to Evans and Eberhard (1970) a number of Aculeata destroy their host. They therefore prefered the name parasitoid, a word first used by Reuter (1913). Because of the doubtfulness of a number of cases we do not distinguish between parasites and parasitoids and only use the first name.

REFERENCES

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