Coprophagy insect brethren of the crowned pigs of Borneo

Cuenot, the French biologist, trying to demonstrate the finality of the things of this world, once wrote in a humorous vein, "Excretion (he meant egestion) is pleasant!"

Horatius, the Latin poet, in his Satires, wrote one day the following verses:

"Mentior at si quid, merdis caput inquiner albis

Corvorum ; atque in me veniant mictum atque cacatum...

(If I am lying in something, may my head be covered with the white shit of the crows, that come to me poo and pee).

This aspect of life has been touched in Latin or Greek more bluntly in writings of Suetonius, Petronius and Aristophane comedies. During that classical period a spade was called a spade.

This aspect of life, egestion, is generally considered a humiliating aspect, and we try to keep it in privacy, and do not talk about it. But as students of life, we should have no hesitation in discussing it boldly, here mainly in relation to insects.

Human and certain animal excreta provide food to Scarabeidae, the dung beetles. Those insects are however selective and pure gourmets. Some prefer human excreta, others monkey's shit, still others are found to live on bird droppings, or on antelope feces or elephant discharges. Under our climates, as under the tropics, the selection is strict and nearly as strict as is plant selection among the phytophagous insects. Certain species are specialized for feeding on semi-liquid excreta, like cow's dung, while others choose dry excreta as that from rabbit, marsupials, or horse. With the introduction of cattle, flies multiplied fast in Australia. It was the same in Hawaii with introduced mammals normally producing liquid feces, such as cattle. The problem was solved to some extent by Waterhouse in CSIRO in Canberra by introducing African and other tropical scarabs to feed on the liquid dung. This led to drastic reduction in the number of flies. However, PJ must say that there are still flies in the land of kangaroos, and they still are quite abundant. Somehow this introduction of dung beetles did not fully succeed. In New Guinea, the local scarabs (Onthophagus) are adapted naturally to feeding on horse excreta (Paulian, 1972), and they could never take to cow dung. In New Caledonia and Vuanatu (New Hebrides) also there were no big mammals in the past, except bats and a few rodents, and local scarabs fed normally, near Noumea, among others, on bird (cagoo) excreta. In Australia, New Guinea, New Zealand and New Caledonia no beetle has adapted to semi-liquid cow dung. An introduction of one Onthophagi-ni, two Oniticellini and one Sisyphini, all scarabs, has brought to New-Caledonia and Vuanatu cow dung feeders (Gutierrez et al, 1988). Three among the introduced species appear to be established on those islands. In Australia, there are dung beetles, which are adapted naturally to plant food as normal diet.

One day, in Ethiopia, in Ogaden valley, in a locality named Awash, one Ethiopian student of PJ observed big scarabeids, very busy around buffalo dung, feeding on it. He looked around and said: "It is their injera". Injera could be translated as bread. Injera in Ethiopia is the acid pancake made with teff flour. Teff is a peculiar Poaceae (Gramineae), Agrostis teff, with very small grains, cultivated in Ethiopia and in Eastern Sudan. It tastes a bit like sarracen (buckweed) flour. Yes, it was injera for those scarabs, who were eating it with delight. All we can say is that everyone has his own taste, even if that taste seems disgusting to us. This holds for the human species too. One man's joy is another man's sorrow (Jolivet, 1991).

It is a fact that nature has done well in adapting every species to search for and locate the source of suitable nourishment, and only exceptionally animals feed on their own excreta. An example of this is gorillas, which probably take to autocoprophagy to get vitamins. The smelly substance of excreta, the skatol (3-methylindol) is used, in small quantities, as a stabiliser of the flavour of vanilla in ice creams. Nature synthesizes skatol in Arum flowers (Zantedeschia aethiopica) to attract pollinating flies, and civet cats in Africa also have glands producing skatol. This chemical is widely used in perfume industry for giving long shelf life to odours. PJ remembers those civet cats in Ethiopia, which were expressing their hate when people were pressing their perfume glands to extract the precious molecules. The Abyssinian cheaters used to mix up sometimes child poo and that was why our perfumers in Paris used to pay the shipment only after analysis of the contents. After all, consistency and smell and even similar chemical formula, were present in the two, above mentioned, producers. Now an anonymous poem looks rather appropriate:

Why Your Offal Smell Awful.

Your body ejects poisons and things It won't need,

And these are the things on which You mustn'feed.

If your offal smelt tasty, like fresh Cherry-pie,

Then you might eat it, and then you Would die.

Your offal smells so bad that you kick

It off your plate

You then may survive and may

Procreate.

If ever an animal found its waste in Good taste,

Evolution has cured that condition Post-haste.

It would be necessary to ask gorillas why they behave that way. However, animals feed on other animal excreta, but practically never on their own.

Excreta are an excellent attraction not only for scarabs, but also for the splendid butterflies, often sensitive to smells, which to us seem nauseating. Around 50 years ago, one Belgian M.D., whom I want to keep anonymous, was the director of a hospital in Congo, and was also an ardent collector of butterflies. This medico-entomologist accumulated human excreta, with help of his boys, deposited over paw-paw leaves (Carica papaya), generally used in that country to tenderise the emaciated and leathery chickens living there. Those paw-paw leaves, so beautifully decorated with feces, were religiously deposited on the various shelves of the hospital refrigerator. Each morning the medico used to take one leaf, and place it on the garden lawn. Nearby a native, a green net in his hand, was trying to catch the beautiful Charaxes or other butterflies feeding on the decorated leaves.

The truth is that with modern traps, suspended on trees, one can catch very easily these Lepidoptera, attracting them with rotten bananas and also human urine. Such secret recipes are also known to entomologists as making a cocktail of excreta, urine, rum and rotten fruits. I think that the good Belgian doctor was trying to do his best in his circumstances.

The splendid Graphium weskei, a Papilionidae, of which the host-plant remains unknown, gets down to drink in the middle altitude (1500 m) water falls in New Guinea. It is equally well sensitive to traps with organic matter and to pure water. It normally drinks on humid sand along the shores. Beetles, other than the dung beetles, may be also attracted by feces. In Florida, an American colleague, Bob Woodruff, found one day a chrysomelid (Lamprosomatinae) Oomorphus florid-anus, feeding on rodent excreta. Normally that genus is an ardent feeder on Araliaceae plants. Is it an aberration? No, it is simply attraction towards a medium rich in water, salts and nitrogenous compounds (Jolivet and Verma, 2002).

Recently a good book has been written on this topic (Lewin, 1999). In this book not less than 22 paragraphs have been dedicated to insects. It is known that merdigerous, merdicolous, scatophagous, stercorarious, onthophagous flies and beetles are numerous. Merda in latin, skatos, stercos, onthos in Greek, all mean shit. These words show the richness of Latin and Greek languages. In this book, excretion is minutely studied and described, including its structure, its smell and its chemistry.

Lewin seems to assign partial production of greenhouse gases to the flatulence of cows. Termites also contribute gases in this process. In amber, when termites are caught, you can see the fossil traces, as tiny bubbles, of their palaeontological flatulence. He also mentions frequency of defecation among insects and vertebrates. He says that the Guatemala viper excretes once a month (practically it is the same for the sloth, but the ceremony involves also moths living in the sloth's fur) and that the rabbits do it once every three minutes. Termites feed on plant matter and fungi along with their intestinal fauna (Hypermastiginae). They produce gases abundantly. Their number compensates for their small body size. Americans have attributed yellow rain in Vietnam to the overabundant excreta of the bees.

It is certain that dung beetles help to clean our planet by feeding on animal excreta, while necrophores, like vultures and hyaenas also help by feeding on cadavers. Many books have been dedicated to the biology of the dung beetles (Hanski and Cambefort, 1991), and this significance of the scarabs has been well brought out by Lewin. Lewin is also a great oceanographer, and is married to a marine entomologist. He combines qualities of an eminent scatophile and a poet, celebrating plants and insects (Lewin, 2003).

— Fig. 42.1. Urinating posture of Allomyrina dichotoma (Linnaeus) (Scarabeidae, Dynastinae), in Japan (after Ohtani and Kuribayashi, 1985).

Human excreta are normally rich in salts, and the pigs in the Orient feed on them with relish. All tastes are in nature, but for the pig it is Lucullus cenat apud Lucullum. In South East Asia, thanks to pigs, everything is eaten and thus removed, and river banks are clean, unlike what we see in the Middle East. It is true that in Vietnam, for instance, the poor pigs have nothing else to eat, but for duckweed, which is collected dextrously by the women in ponds. One of PJ's former colleagues, Dr. Farinaud, eminent malarialogist with WHO, told us once that one day in Vietnam, as he was busy relieving himself in a rice field, he felt something humid on his rear. It was a pig, which was removing directly the product from the producer. In Borneo, the Dayaks live in houses, raised on poles. People, during night, relieve themselves through gaps between wooden planks in the floor. Below chickens and pigs fight for the manna dropped from the sky. We nicknamed them the crowned pigs of Borneo, as they were generally crowned with excreta, but they all seemed to be happy and to be fighting for the fodder. In Vietnam the piggery is often near toilets, and the pork is excellent, if you don't think of the nourishment source for the pigs. Sweet and sour pork is always good everywhere. Those pigs do not eat their own excreta. They don't live in close confinement, as their food collection requires perpetual movement. Pigs are teachable, and the dwarf ones, used as pets in the US, are as clean as cats and dogs.

Ethiopians, as also Jews or Arabs, do not eat pork meat. Pigs, except wild, are rare in the country. Hyaenas do cleaning during nights in the streets of Addis Ababa. In Brazil and in Guyana, there are the urubus, sort of big crows, which do the job. Nature has an army of nonpolluting cleaners, such as hyaenas, pigs, birds, insects, worms etc. Armadillos also contribute to the cleaning in America. Bacteria and fungi achieve the final destruction. Plastics, however, resist destruction by such natural cleaners.

During the Palaeozoic, recycling of dead bodies was done by the great proto-myriapods, the Arthropleura, two metres long monsters, which were feeding on lycopods and decaying matter. Scavengers were numerous at that time. During the Mesozoic, there were necrophagous dinosaurs, like Tyrannosaurus, and they were necessary to clean the enormous bodies of their herbivorous 'colleagues'. There were also very probably numerous coprophagous beetles, since the first Coleoptera appeared during the Permian, just before the dinosaurs. The dung beetles were elevated to the rank of sacred animals during Egyptian times, because they symbolically carry the world with their ball of dung.

A folklore runs as follows. During the colonial period, a French administrator flew into a towering rage on seeing pigs wandering on the streets of a village near the Chinese border. He complained to the chief of the village. The chief knew the French people and he agreed formally to send away or to destroy the pigs, but he did not do anything. During his second visit, the administrator saw again the wandering pigs in the streets. He got angry again. The chief again politely agreed to take necessary steps, but again did nothing. During his third visit, the administrator shot at the pigs, and the village chief thought that it was time to do something. With the help of a peasant he carried the pigs to Yunnan on the other side of the border. When the administrator came back for the fourth time, he found the village very dirty. He complained that to the chief, who said that without the pigs it was perfectly normal to see the village dirty. Then the administrator asked to bring back the pigs. The chief brought them back from Yunnan, and again the village was totally clean.

All animals excrete, even the smallest members of plankton in the sea, and almost all the insects. Urinating postures of some Dynastinae in Japan has been described by Ohtani and Kuribayashi (1985). Some, like big saturniid moths, don't have an opened intestine and don't excrete at the adult stage, but they do well as a caterpillar. Some chironomids and many other insects also don't eat or excrete as adults.

There are many books on excretion and we would recommend a classic, the Microscopic Coprology of Langeron and Rondeau du Noyer (1930). Coprology we find in the writings of many authors (Rabelais, Swift, Lord Byron, even Shakespeare). Sigmund Freud, who was victim of a chronic constipation, is one of the celebrities involved. Finally, all living things must excrete at certain moment of their life. I cannot say we must feel proud of it, but we are not ashamed of it. Ad augustaper angusta. In view of the universal process of excretion in the animal world the importance of coprophagous cleaners like the dung beetles should be appreciated.

References

Gutierrez, J., Macquee, A. and Brun, L. O. 1988. Essais d'introduction de quatre espèces de bousiers Scarabaeinae en Nouvelle-Calédonie et au Vuanatu. Acta Oecologica 9 (1): 39-53.

Ohtani, T. and Kuribayashi, S. 1985. Urinating posture of the Japanese horned beetle, Allomyrina dichotoma (Linnaeus). Kontyu, Tokyo 53: 247-248.

Hanski I. and Cambefort, Y (eds.). 1991. Dung Beetle Ecology. Princeton University Press, Princeton, N. J.

Jolivet, P. 1991. Curiosités Entomologiques. Chabaud publ., Paris.

Jolivet, P. and Verma, K.K. 2002. Biology of Leaf-Beetles. Intercept Publisher, Andover, U.K.

Langeron, M. and Rondeau du Noyer, M. 1930. Coprologie microscopique. Masson & Cie. Publs., Paris.

Lewin, R. A. 1999. Merde. Excursions into Scientific, Cultural and Socio-Historical Coprology. Aurum Press, London and Random House, uSA.

Lewin, R. A. 2003. Blue Green. A Collection of Poems by Ralph Lewin. Kluwer Academic publs., Dordrecht. Boston.

Paulian, R. 1972. Récoltes de M. P. Jolivet en Nouvelle-Guinée (Col. Scarabeidae Onthophagidae). Bull. Soc. Entom. Fr.: 215-217.

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