CHAPTER 1 Aristotle and Theophrastus on plant-animal interactions COSTAS A. THANOS Institute of General Botany, University of Athens, Athens 15784, Greece Key words: Aristotle, Theophrastus, zoology, botany, plant-animal interactions, biology Abstract. Aristotle and Theophrastus, the last great philosophers and scientists of Greek Classical Antiquity, are the founding fathers of Zoology and Botany, respectively; they should also be honoured as the co-founders of Biology. They were close friends and life-long collaborators who evidently decided to pursue an organized study of the living world, probably in Lesbos at 344 BC (the landmark for the creation of the Science of Biology). The product of their division of labour, the voluminous zoological and botanical works of Aristotle and Theophrastus, respectively, were actually used contemporaneously as university textbooks by the students of the Lyceum. Besides numerous comparisons and analogies, mostly on general issues, between animals and plants, both Aristotle and Theophrastus deal with various cases of plant-animal interactions, covering virtually all aspects of the field. Their scientific approach is notable, although the barriers to knowledge imposed by their era did not permit a significant contribution on issues like plant sex and pollination. Their important accomplishments on plant-animal interactions include herbivory and poisonous plants, plant pests and use of manure, insect-repellence and gall formation, fig caprification and apiculture, seed dispersal and seed infestation. Introduction physician at the court of Amyntas C', father of Philip B'. At the age of 17, Aristotle moved to Aristotle (384-322 BC) and Theophrastus (371- Athens where he studied and subsequently taught 286 BC) should be considered as the last great in Plato's Academy. Although by far the most philosophers of Greek Classical Antiquity. They brilliant among Plato's pupils, Aristotle was not represent, in particular, the culmination of the appointed as the new director of the Academy natural philosophy of the Ionian scientific after Plato's death (347 BC). Apparently as a result tradition which was inaugurated on the Aegean of his non-designation and with a team of coast of Asia Minor, several centuries earlier, and colleagues and followers, he travelled to Assos of reached its climax on the opposite coast, at the Troad (Asia Minor) and founded a new school brightest cradle of Ionian civilization, Athens. It there. Unfortunately, the venture came to an should be borne in mind, however, that both abrupt end three years later with the assassination Aristotle and Theophrastus were not Ionians but of the patron of the school, Hermeias, the Macedonian (Dorian) and Aeolian, respectively; hegemon of Assos. accordingly, they never became full citizens of It is at that difficult moment in Aristotle's life Athens. During their lifetime they experienced the that Theophrastus seems to have played a decline of the City-State system of Classical prominent role; probably, at his suggestion Greece and its eventual replacement by a more or (Morton, 1981), the two men moved to the nearby less unified Greek State dominated by the northern island of Lesbos, in the North Aegean Sea. Greeks (the Macedonians), under the leadership of Theophrastus (his original name was Tyrtamos) Alexander the Great. was born at the Lesbian town of Eresus and his Aristotle was born at Stagira of Chalcidice father, Melantas, was a local fuller. It is probable (Macedonia) and his father Nicomachus was a that at an early age (ca. 355 BC) he went to 3 M. Arianoutsou and R.H. Groves, Plant-Animal Interactions in Mediterranean-Type Ecosystems, 3-11, 1994 1994 Kluwer Academic Publishers, Printed in the Netherlands. 4 Athens and enrolled in the Academy, where he Animals, in 10 Books), Περί ζώωv μoρίωv, PA was acquainted and associated with Aristotle. Ac- (De Partibus Animalium, Parts of Animals, in 4 cording to Morton (1981), Aristotle evidently took Books), Περί ζώωv κιvήσεως, MA (De Motu to the highly intelligent, industrious and good- Animalium, On the Movement of Animals, 1 natured young man, who became his close friend Book), Περί πoρείας ζώωv, IA (De Incessu and life-long collaborator. Animalium, Progression of Animals, 1 Book), After an apparent rest at Lesbos for two years Περί ζώωv γεvέσεως, GA (De Generatione (344-342 BC), Aristotle received an invitation by Animalium, Generation of Animals, in 5 Books). Philip B', king of Macedonia, to serve as a tutor The first work could be compared to what is for his teen-aged son, the future Alexander C' the currently considered a General Zoology text, Great. So Aristotle returned to his native country whilst the second is the earliest treatise on Animal where he spent several years at the royal court, in Physiology (the latter works covering more the capital Pella, teaching the young prince ‘τo ευ specialized fields). The also extant work Περί ζηv’ (in free translation ‘the quality of life’). φυτώv (De Plantis, On Plants, in 2 Books) that is Although there is no direct information, it seems included in Aristotle's minor works, is definitely highly probable that during his sojourn in Pella not by him. It was almost certainly written more and afterwards in Stagira (when Alexander came than 3 centuries later by Nicolaus of Damascus of age), Aristotle was accompanied by (1st century BC) and according to Morton (1981) Theophrastus. reflects the level to which Peripatetic science was In 335 BC, immediately after the struggle over later reduced. domination of Greece was finally decided in Theophrastus, though not admired as a major favour of Macedonia, Aristotle returned to Athens, philosopher (of the stature of Aristotle, Plato or together with Theophrastus, and founded the Democritus), was also a voluminous writer and is Lyceum. After Alexander's death (323 BC), credited by Diogenes Laertius (3rd century AD) Aristotle had to flee Athens for Chalcis, Euboea with 227 treatises. Apart from his well known where he died a few months later. Theophrastus Characters, his only other extant works are: Περί became the second director of the Lyceum and the φυτώv ιστoρίας (Historia Plantarum, Enquiry into Peripatetic School (the name of the school was Plants, HP, in 9 Books) and Περί φυτώv αιτιώv derived by the habit of lecturing while strolling (De Causis Plantarum, Causes of Plants, CP, in 6 around the gardens of the Lyceum) which reached Books). These works are the first, truly scientific its apogee of success during his 37 years of botanical writings and correspond roughly to administration. The function of this institution was modern textbooks of General Botany and Plant to train the leaders, officials and experts of the Physiology, respectively. Among his non-extant new era (Morton, 1981). Like its rival Academy, works, 6 Books on the behaviour of animals are the Lyceum was a true University of its epoch: an also included. up-to-date curriculum emphasizing the Peck (in his 1965 Introduction to HA) observational sciences, numerous lecturers, as discusses all the relevant bibliography (in many as two thousand students and a spacious, particular Thompson, 1910) concerning the dates well designed campus with buildings and open-air of the treatise and concludes that Aristotle's facilities, a very important library, a museum and natural history studies were carried out, or mainly the first botanical garden. carried out, between his two periods of residence Aristotle is generally considered one of the in Athens, and especially during the 2-year stay at greatest Ancient Greek philosophers and during Lesbos (344-342). A similar conclusion is reached his rather short lifespan he wrote on virtually by Morton (1981) who suggests that Aristotle had everything (in possibly as many as 400 works). not studied animals systematically until 344 BC Besides his philosophical treatises, he has left a when he moved to Lesbos. Kiortsis (1989) cites as number of voluminous works on natural history, writing dates the following: for HA 347-342 BC, the most important among them being the for PA 330 BC and for GA 330-322 BC. following (all of them fortunately extant): Περί τα Mitropoulos (in his Introduction to HA) believes ζώα ιστoρίαι, HA (Historia Animalium, History of that Aristotle's zoological works have been written 5 in collaboration with several colleagues and well, the origins of the two men must have played disciples (Theophrastus, Strato, Eudemus and a role in that decision. Aristotle's father was a other Peripatetics). In his opinion, Theophrastus physician and he himself had some medical may have contributed much and, in particular, the knowledge; Theophrastus, on the other hand, was spurious 10th Book of HA may belong exclusively closer to agriculture and forestry and evidently to him. On the other hand, Balme (in his 1988 was aware, from his father, of many technical Introduction to the third volume of HA) concludes aspects of handling clothes and leather. I assume that there seems to be no compelling reason to that the outlines of their major biological treatises believe that HA was written before the other had been thoroughly discussed and worked out biological treatises. In his opinion, all the during their stay in Lesbos; the bulk of their work available evidence suggests that Aristotle wrote would be already completed within the following HA I-IX as a study of animal differentiae; he decade, just in time for the inauguration of their collected the data initially from other treatises and Lyceum. If we accept that all these natural history then proceeded to complete the study from new books are simply University textbooks for the use reports, a process which was still unfinished at his of the Lyceum students, it is obvious that such death. The likeliest period for the bulk of his work treatises would continually be updated and is his stay at Lesbos (344-342 BC) and subsequent corrected (and this would account partially for the years (until 336 BC).