THEOPHRASTUS ON PI-ANT-ANIMAL INTERACTIONS

COSTAS A. TT{ANOS

Institute of General Botany, University of Athens,

Athens 15784,Greece

Theophrastus,the Ancient Greek philosopher,was born around 370 BC at

the town of Eresus,in Lesbos island and lived to the age of 85. During his
lifetime the City-State system of Classical Greece declined and was eventually
replaced by a more or less unified Greek State dominated by the northern
Greeks, the Macedonians, under the leadership of Alexander the Great.
Theophrastus,though not a major philosopher,proved to be an extraordinary
scientist. His two voluminous works IIEPI OYTON ETOPIA: (Historia
Plantarum, Enquiry into Plants) and IIEPI OYTON AITION (De Causis
Plantarum, Causesof Plants) are the first truly scientific botanical writings and
have led to his international recognition as the founder of the Scienceof Botany
(e.g. Morton 1981, Evenari 1984). These works were written in their final
version around 300 BC, exactly23 centuries ago, and their translation into
English was completed only very recently.
During a great part of his [fe, perhapsfor aslong as30 years,Theophrastus
was very closely associated to Aristotle, whom he succeededas the director of
the Lyceum of Athens. Aristotle, probably the major Ancient Greek philosopher,
is further credited nowadays as the founder of Biology in general and of
ZooIog in particular. It seems that Aristotle had not studied animals
systematically until 345 BC when he moved to Lesbos, together with
Theophrastus and probably after the latter's suggestion.This date may constitute
the landmark of the creation of Zoolory and Botany as a result of the two
friends' joint decision to pursue an organised study of the living world.
Therefore, a rough division of labour was mutually agreed and this may also
explain why both refrained from penetrating, in their writings, into each other's
specific field. Nevertheless, overlapping and casual references of interactions
between plants and animals were obviously inevitable.
Theophrastus, in particular, is repeatedly attempting comparisons and
analogies between plants and animals (the latter serving always for reference)
though only in basic or general issues; for instance in discussing general form
and basic functions such as nutrition and reproduction. In introducing these
subjects, Theophrastus writes: "We should not expect to find in plants a
completecorrespondence with animals".Among severalparticular casesone can

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 mention the similarity between the nature of seedsand that of eggs as well as
the analory of the leaf sheddinghabit in plants to sheddingof horns, feathers
and hair in animals.The principled comparisonsbetweenanimalsand plants in
both Aristotle's and Theophrastus's works reflect their conception of a single
natural kingdom of living organisms, a profoundly scientific view derived from
the naturalism of Ionian philosophersand in marked contrast to the idealism
of Plato.
Although Theophrastus was ernphatically teaching botany instead of
providing a local flora or a treatise on agriculture, the great economic
importance agriculture had already gained during his time is obviously reflected
to a certain degree in his works. Therefore, in numerouspassagesone finds
discussionsabout manuring, particularly concerningthe effect of the various
animal sourcesof manure to the growth of plants of variousimportant crops.In
addition, several referencesto animal pests and animal-causeddiseasesof
plants, particularly crops, are made. Considerable attention is devoted to seeds
that are consumedby the larvae of beetles(thought to be producedby the seed
itsel|, a most commoncasein legumeseedswhich are infestedby the larvae of
beetles (today identified as bruchid beetles).As an obviousresult of the great
importance of galls as a tannin source, Theophrastus deals with them in detail
and describes ten different types of galls produced by oaks (Ouercus spp.), as
well as additional ones occurring in terebinth (Pistacia terebinthus) and elm
(IJlmus glabra). Although the general belief of the tirne was that these were
formations of the host plant, Theophrastushad noticed the presenceof insects
within certain galls, as for examplein the terebinth one and in the transparent,
watery gall of the leaf rib of oak.
In discussingherbivory,Theophrastusmakesa strongpoint that no general
rule can be reached.Some parts of the plant may be edible and other inedible;
also some animals seem to prefer the tender parts while other prefer the dry
ones.IIe further statesthat it is most usual that leavesare not edible while the
fruits of the same plant can be consumedby both humans and animals. Less
usual is the case of lime (Tilia europaea) with edible leavesbut inedible fruits.
Another interesting observation is that animals find legumes a pleasure to
digest. Theophrastus also mentions several examples of specific plants that
produce toxic compounds and which may cause poisoning or death to the
animals that might consume the particular plant parts. Examples furnished
include black hellebore (Helleborus cl'cloph]'11us),fatal to horses and cattle, the
deadlyroot of Aconitum sp.which is not touchedby sheepor other animalsand
Euonvmus spp. leaf and fruiq both fatal to sheepand especiallyto goats.An
interesting exampleis the highly poisonousroot of Thapsiaearganicawhich was
never touched by the indigenous cattle of Attica, where this plant was
particularly abundant.Imported cattle, on the other hand,might feed on it and
perish of diarrhoea. The Cretan dittany (Qfiga4lq dictamnus) is an example
 of healing action. The plant is describedby Theophrastusasbeing rare and with
very narrow distribution attributed to the fact that the goats are fond of it and
graze it down. Theophrastus adds also, somewhat sceptically though, the story
of the arrow (cited by Aristotle as well) according to which a wounded goat
seeks to eat dittany and as a result the arrow drops off. This impressive story
inspired the Flemish engraver Dapper, who illustrated it in 1703 (Baumann,
1982).A final note concernsthe insectrepellantaction of certain plants or plant
products such as olive oil, the pungent smell of marjoram (Qftganum spp.) and
the likes, as well as both speciesof Inula (i.e. eraveolensand viscosa).
In the field of pollination one hasto admit that no great contribution to our
knowledge could be expectedduring Theophrastusera. The real nature of
flowers as sexualorganseludedTheophrastusbut this is no surpriseat all since
plant sex was suggested only two millennia later, in 1672, by the English
physician Thomas Millington while the first experimental proof was furnished
subsequently,in L694, by Rudolf Jacob Camerer, a German professor of
medicine at Tubingen (Bristow 1980).Nwertheless,Theophrastuswas able to
distinguish the two sexes in certain dioecious plants on the basis of their
fruiting abfity, the female being of course the fruit bearing type. The caseof the
date palm (Phqgail dactvlifera) has been well known since at least 1500 BC, as
illustrated in the bas-reliefsof Nimrud in Mesopotamia.Theophrastusproceeds
further to the point of comparing the dusting of the female flowers with the
male inflorescence to what was observedwith fish, when the male sprinkles his
milt on the eggs as they are laid. Another most interesting case is the
caprification of the fig fig tree (Ficus carica) being a close companion of man
for many millennia in Eastern Mediterranean. In certain cultivated fig varieties
caprification is the necessaryprocedureto promote proper maturation of the
syconium, the complex fruit of fig. Wild figs were hung on the cultivated fig tree
or wild trees were planted on eminencesnear the fig orchard to ensure the
insectsan easyflight down wind. For it is the insectsthat after having grown
inside the developing seeds of the wild figs will pierce the pericarp tissues,
sometimesleaving a leg or a wing behind, to seek another fig to get in and
spend the rest of their life (having carried the pollen from the first to the latter
while laying at the same time their eggs).These insects are the "psenes"of
Theophrastus,the now called fig wasps,Agaontidae (the particular one being
Blastophaga p!g!gc). Theophrastus also describes another qpe of waspswhich
never get out of the fig and are sluggishlike drones. These are actually the
wingless male waspsthat fertilise the females before the latter venture their trip
to their secondfig-host.
In the field of plant dispersal, numerous accounts are furnished by
Theophrastus.The cormlets of the corn-flag (Gladiolus segetum)are found in
moles' runs for this animal likes them and collects them, this habit leading
unintentionallv to the dispersal of the plant. A similar case is the cachine of
acorns by jays and other birds, resulting again in the accidentaldispersal of
oaks. An example of epizoochory is the fruit of Galium aoarine which sticks on
the clothes;endozoochoryis representedby ivy fruits reported to be consumed
by birds. The fruits (legumes) of Colutea arborescensare described as being
eaten by sheepand their seedsare found germinating promptly from within the
droppings.A final fine exampleof zoochory is the case of the mistletoe. It is
also a masterly treatise by Theophrastuswho having identified the two species
occurringin Greece(Loranthuseuropaeus and Viscumalbum),asksquestions
and provides answersconcerning the peculiar habit of these plants to grow only
on a host tree (that may belong to a number of speciesthough).He concludes
that it is the birds that consumethe mistletoeberries; the seedspassunharmed
through the digestive track of the bird and are able to establish their new
seedlingsonly when the droppingshappento fall on a host plant.
A final point to be born in mind is that althoughTheophrastuswas by all
standards a great scholar and scientist, his works reflect precisely the overall
attitude towards nature as well as the level of accumulated knowledge of the
Greek Society in general, during the end of Classical Antiquity.

REFERENCES

Baumann H., 1982. Die Griechische Pflanzenwelt in Mlthos. Kunst und

Literatur. Hirmer Verlag, Munchen.

Bristow A., 1980.The SexLife of Plants.New English Library London.

Evenari M., 1984.Seedphysiolog: its history from Antiquity to the beginning

of the 20th century.The Botanical Review50:119-142.

Morton A.G., 1981.Historv of Botanical Science.Academic Press,London.

Theoohrastus.Enouirv into Plants.Vols 1-2.Hort A.F. ftranslator).1916and

1926. William Heinemann Ltd, London and Harvard UniversiW Press.
Cambridge,Mass.

Theophrastus.De Causis Plantarum.Vols 1-3. Einarson B. and Link G.K.K.

(translators). 7976 and 1990.William Heinemann Ltd, London and Harvard
University Press,Cambridge,Mass.