The Lagoon
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Modern life-history theory. For an introduction to life-history theory see ROFF (2002); for a typical paper on fish life history see WINEMILLER and ROSE (1993).
LXXXV
Greek life expectancy. Life expectancy at birth on Ikaria appears to be within the national range for Greece (C. Tsimabos pers. comm.), but a closer look at survival of the oldest old suggests that Ikarian women, at least, have a significant survival advantage at late age relative to Greece as a whole (M. Poulain pers. comm.).
The length and shortness of life. See KING (2001) for the authoritative account of A.’s theory of ageing. A. says we must investigate why some animals are long lived and others short lived at LBV 464b19. He tells of mayflies at HA 552b18 and how winged insects die at the end of summer at HA 553a12. He summarizes the comparative biology of lifespan at LBV 466a1 and says that the old are cold and dry at LBV 466a21. Is there a single explanation for death? JSVM 478b22. Is there one cause for the diversity of longevity? LBV 464b19. His account of the relative heat and moisture of different animals is given at LBV 5, 6. For the role of fat in promoting life see LBV 466a24, cf. PA 651b1; see FREUDENTHAL (1995) ch. IV. On the longevity cost of reproduction, LBV 466b7, HA 576b2 and GA 750a20; see LEROI (2010). For the same idea of a senescence cost of reproduction in modern evolutionary biology, see WILLIAMS (1966), ROSE (1991), LEROI (2001), ROFF (2002). On regeneration in plants see LBV 467a7, snakes and lizards HA 508b4, and Hydra BOSCH (2009). On death by failure of cooling systems, see JSVM 470b10; on the seizing up of cooling organs, JSVM 479a8 and JSVM 479a31; on the (false) etymology of earth/age, GA 783b7. On the vulnerability of old animals to variation in the environment, see JSVM 474b30, JSVM 478a15 and JSVM 479a16. On the role of the soul and ageing, see DA 415b25, DA 434a22 and throughout JSVM. Death is said to be in the nature of living things at JSVM 464b29, PA 644b23 and GA 731b24. For modern mechanistic theories of ageing see FINCH (2007) and, more recently, GEMS and PARTRIDGE (2013). For thermoregulation and ageing in humans see SOMEREN (2007). For evolutionary theories of ageing see WEISMANN (1889); the modern theory is due to MEDAWAR (1951/1981) and WILLIAMS (1957); see LEROI (2003) ch. IX for a popular review. For the general theory of the destruction and regeneration of natural objects: LBV 2, 3 and DC 288b15 and Ch. LXXX.
LXXXVI
The story of Daphnis and Chloe. I used the Loeb edition of Longus’ Daphnis & Chloe, translated by Jeffrey Henderson, 2009. The idyllic scene is described in I, 9–10; see MASON (1979), GREEN (1982) and GREEN (1989) ch. 3.
LXXXVII
Sponges and other plant–animal dualizers. On sponges, HA 487b10, HA 548a32, HA 548b8, HA 588b21, PA 681a10. VOULTSIADOU (2007) discusses the cultural role of sponges in antiquity. For the other plant–animal dualizers, sea anemones, sea squirts, etc., see HA 487b10, HA 547b12, HA 548a22, PA 681a10, PA 683b18. It is hard to say whether A. thinks that some or all of these creatures are animals, plants or something in between. For example, he says of sponges that they are ‘plant-like’ or even ‘plant-like in every respect’. I take it that A. thinks that they are, on balance, animals since they all seem to have at least one capacity of the sensitive soul (locomotion or sensation or appetite); that would be consistent with his polythetic approach to classification. But perhaps the most compelling reason to believe that A. thinks they are animals is that he discusses them in HA – where he might have discussed them elsewhere, e.g. in his lost On Plants (Peri phytōn). Similarly, in his Enquiries into Plants (HP IV, 6.10), T. touches on sponges but then says ‘they’re of a different character’ – presumably animal, to be dealt with elsewhere. The pseudo-Aristotelian text de Plantis, which is thought to be a commentary by Nicholas of Damascus on A.’s Peri phytōn, also struggles with this question since it asserts that animals have sensation, plants do not, shellfish have sensation but are at once both animals and plants: DP, 1; DROSSART LULOFS (1957). See LLOYD (1983) ch. I, 4, LLOYD (1996) ch. 3 and LENNOX (2001a) p. 301 for how A. deals with these creatures. For T. on corals and other sea ‘plants’ see St 38 and HP IV, 6 where the precious coral is the ‘sea palm’; HP IV, 7.2 on the growths in the Gulf of Heroes (Aqaba). See THOMPSON (1947) p. 250 for scepticism on sponge contraction; I thank Sally Leys, University of Alberta, for telling me about movement in sponges; see also NICKEL (2004).
‘Nature proceeds from the inanimate . . .’: HA 588a1, cf. PA 681a10, Meteor IV, 12 and GA 731a25. It may seem that there is a tension between this claim and A.’s belief that the world is composed of discrete kinds each possessed of its own inherited form and teleologically defined essence; however, by ‘continuity’ A. did not mean that the continuum of kinds is infinitely divisible, nor that kinds overlap so that the boundaries of one cannot be distinguished from the other, but only that they form a graduated series that progresses in small but discrete steps; see GRANGER (1985) contra LOVEJOY (1936).
LXXXVIII
Aristotelian themes in Darwin and vice versa. For the history of Natura non facit saltum and its use by Darwin see FISHBURN (2004). ‘Ever since Darwin’ was the title of Stephen Jay Gould’s 1977 anthology of essays from Natural History magazine, so named for the cliché used as the starting point of so many papers by evolutionary biologists. On the meanings of genos see Metaph V, 28. Some scholars (LENNOX (2001b) ch. 6 and PELLEGRIN (1986) ch. 2) have emphasized that in the zoology A. uses genos to designate a group of organisms that are related by descent. That’s not unreasonable so long as it is recognized that, in the case of a megista genos (e.g. birds) with subordinate genē (sparrows, cranes) we recognize that A. is not saying that sparrows and cranes are related to each other by descent since that would imply a common ancestor, i.e. evolution. His use of genos in the sense of common descent (definitions 1 and 2 in Metaph V, 28 which are very close) can, then, apply only to genē that are atoma eidē, i.e. that actually interbreed (e.g. humans). In general, he must be using genos in the third sense given in Metaph V, 28 which is purely classificatory and does not imply anything about ancestry. For a discussion of A.’s. anticipations of evolutionary themes (without being an evolutionist) see KULLMANN (2008).
LXXXIX
Darwin on A. See LENNOX (2001b) ch. 5 and GOTTHELF (2012) ch. 15 on A.’s indirect influence on Darwin. ‘Read A. . . .’: DARWIN (1838/2002–) p. 267. STOTT (2012) tells the story of how A. infiltrated himself into the Origin as an evolutionary precursor. Translations of The Parts of Animals: today LENNOX (2001a) is the authoritative English translation of PA. His superb commentary focuses on philosophical and theoretical aspects; the zoological facts still have to come from OGLE (1882), in this case p. 240, n. 36: ‘the camel, the cats, and many rodents including the hare are retromingent’. The German translation and commentary of KULLMANN (2007) are excellent on both philosophical and zoological matters.
XC
A., Linnaeus and the scala naturae. On the origin of Linnaean names see HELLER and PENHALLURICK (2007). In neither of the two passages – HA 588b30 and PA 681a10 – that are generally held to show the Ladder of Nature or scala naturae is the idea very explicit, LENNOX (2001a) pp. 300–1; however, in other passages, particularly in GA, it is clear that A. has a strong sense of how animals should be arranged in a continuum of increasing perfection. See, for example, GA 733a32 on the relative perfection of progeny, and GA 733a1 on the link between parental and parental perfection. The physiology of relative perfection is given in the following passages: hot animals have lungs, PA 669b1; tend to upright, PA 686b26; tend to be larger, GA 732a17; tend to live longer than cold animals (Ch. LXXXV). A.’s theory – PA 648a2 and PA II, 4 – of how the constitution of blood influences intelligence and temperament is in the same spirit, but more complicated. In brief, there are three properties of blood that influence intelligence and temperament: heat, thickness and purity. Although correlated, these properties vary, to some degree, independently in both blooded and bloodless animals, which allows A. to explain the various behaviours of various animals (bulls, bees and so on). Those with hot, thin and pure blood are best – for they are
both courageous and clever. Humans have the thinnest and purest blood of all animals; see LLOYD (1983) ch. I, 3.
The naturalists and the scala naturae. For the history of the scala naturae in Western thought see LOVEJOY (1936) ch. 2 who, p. 79, quotes Albert Magnus. Systema naturae, 1st edition, LINNAEUS (1735); 13th edition, LINNAEUS and GMELIN (1788–93). For a history of zoophytology see JOHNSTON (1838) pp. 407–37; ELLIS (1765). Cuvier classification, first given in 1812, is best known in the version given by CUVIER and LATREILLE (1817).
XCI
The great cuttlefish debate. For accounts of the debate see RUSSELL (1916) chs 3, 5, 6, APPEL (1987), GUYADER (2004) and STOTT (2012). For A.’s analysis of cephalopod geometry see PA IV, 9 and Chs XXIII, XXXVIII and XCVII. For Geoffroy and Cuvier on A. see GUYADER (2004) pp. 143, 155, 181. The terms ‘homology’ and ‘analogy’ have a complicated history but they were first distinguished by OWEN (1843), pp. 374, 378 and OWEN (1868); their meaning, however, has continued to evolve, HALL (2003). On Cuvier’s method see CUVIER (1834) vol. 1, pp. 97, 179–89; and why natural history should not have its Newton, CUVIER (1834) vol. 1, p. 96. ‘The form of the tooth . . .’: CUVIER (1834) vol. 1, p. 181; ‘Natural history has a rational . . .’: CUVIER and LATREILLE (1817) vol. I, p. 6, trans. OUTRAM (1986). For Geoffroy on the vertebrate breastbone and the loi de balancement see GUYADER (2004).
XCII
The evolution of concepts. For Cuvier’s relationship to other thinkers see RUSSELL (1916) ch. 3, OUTRAM (1986), RUDWICK (1997), GRENE and DEPEW (2004) ch. 5, REISS (2009) pp. 103–13. Cuvier’s Conditions of Existence appear in DARWIN (1859) p. 206 and PALEY (1809/2006) ch. 15. For the same idea in modern genetics see LEROI et al. (2003) on cancer in Xiphophorus hybrids and PHILLIPS (2008) on epistasis. Geoffroy’s loi de balancement appears as the correlation of growth in DARWIN (1859) p. 143 and as pleiotropies in LEROI (2001). A. speaks of recurring ideas at DC 270b16, Meteor 339b28, Metaph 1074b1 and Pol 1329b25.
XCIII
A.’s anti-evolutionism. On the Pre-Socratic zoogonies and transformism see CAMPBELL (2000), LLOYD (2006) ch. 11 and SEDLEY (2007). Plato’s transformism is evident from Tim 91D–92C; see SEDLEY (2007) ch. 4. A. considers the idea that all animals might be ‘earth-born’ at GA 762b23. With most commentators, I argue that A. is committed to the fixity of forms. BALME and GOTTHELF (1992) pp. 97–8, BALME (1987d) and GRANGER (1987) argue that he is not, but not convincingly; see LENNOX (2001b) ch. 6. Eternity of kinds/forms: DA 415a25, GA 731b31, Metaph VII, 8–9, GC II, 10–11. A. discusses the deleterious effects of congenital deformities at GA 771a12 and GA 772b35. HENRY (2006a) suggests that for A. the fit to the environment of a given kind is maintained by selection against mutations that fall outside some range so that any animal that bears them ‘will no longer be adapted to that environment and so possessing that feature will be detrimental to its ability to survive and reproduce’. That kind of selection, which is very similar to Empedoclean selection, is known as stabilizing or purifying selection, but I don’t think that A. invokes it. He only says that unconditionally unfit creatures (those lacking essential organs) die and he never ties this to the maintenance of forms, be it with respect to a particular environment or not.
Origin of new kinds/species by hybridization. On Linnaeus’ hybridism see MüLLER-WILLE and OREL (2007). A. discusses hybrids at Metaph 1033b33, GA 738b32, GA 746a29, HA 566a27, HA 606b25, HA 608a32; cf. Mirab 60. The question of whether A. thinks that new animal kinds can arise by hybridization is a tricky one. Scholars such as HENRY (2006b) who would not accept the dual-inheritance system that I do here, argue that both mother and father contribute their forms to the embryo. If so, then hybrids that are a stable mix of the parental forms would be possible. But there isn’t much evidence that A. thought this. In fact, at GA 738b28, where A. discusses dog × fox crosses, he claims that the hybrids will revert to the female’s form. This is incompatible with either exclusively paternal forms or biparental forms since it gives an unexplained priority to the mother’s form or matter. In fact, I think it’s an un-Aristotelian interpolation, probably by Theophrastus – the language of soil and seed suggest a botanist’s meddling with the texts; see CP I, 9.3, CP II, 13.3 and this text n. Ch. LXXI. Geoffroy’s teratological transformism, APPEL (1987) pp. 128, 130–42; GUYADER (2004).
Terato-transformism. On the relationship between the monstrous and natural in A. see GA 770b15 and GA 769b27. The following animals are said to be naturally deformed: seals, HA 498a33 and PA 657a22, and moles, HA 491b28, HA 533a1 and DA 425a10; for other examples and discussions of what exactly A. meant by ‘deformed’ or ‘warped’ see LLOYD (1983) ch. I, 4, GRANGER (1987) and WITT (2013). On how tetrapods came to walk on all fours see PA 686a32; cf. PA 686b21, Tim 91D–92C.
Evolutionism. There is a story, told by Ernst Mayr, David Hull and Arthur Cain, that Aristotle’s ‘essentialism’, via Linnaeus, held up the theory of evolution for 2,000 years. To refute this would require a detailed analysis of what these scholars thought Aristotle and Linnaeus said, and this book is not the place to do it. I shall do so in a future paper.
XCIV
Fossils. For Darwin’s predecessors see MAYR (1982) and STOTT (2012). Some scholars have argued that A. simply did not have evidence for evolution, e.g. BALME (1987d), BALME and GOTTHELF (1992) pp. 97–8; LENNOX (2001b) ch. 6. See Strab I, 3.19 for a theory of Lesbos’ geological history. For Xenophanes’ fossil fish see PEASE (1942); for Xanthus, Eratosthenes and Strato on fossils see Strab I, 3.3–4; Theophrastus on fossil ivory, St 37 and MAYOR (2000). The papers collected in DERMITZAKIS (1999) give an overview of Lesbos’ vertebrate palaeontology. SOLOUNIAS and MAYOR (2004) describe the elephants of Samos and their remains. Herod II, 75 speaks of the winged serpents of Arabia; see RADNER (2007). For Theophrastus on petrified reeds see HP IV, 7.3, and fossiles, Meteor 378a20. There are also a number of references to petrification in pseudo-Aristotelian works, e.g. Prob XXIV, 11, Mirab 52 and Mirab 95.
XCV
Theophrastus’ transformism. For assimilation to the country in wheat see CP IV, 11.5–9; new natures in plants, CP IV, 11.7 [trans. EINARSON and LINK (1976–90)]; degeneration of seed (reversion to wild), CP I, 9.1–3 and HP II, 2.4–6. On darnel see CP II, 16.3, CP IV, 4.5–5.5, HP II, 4.1 and HP VIII, 8.3, where T. recognizes that darnel might just be a weed. See THOMAS et al. (2011) for the evolution and cultural significance of darnel. T. is often said, on the basis of some fragments, to be more resistant to teleology than A. but there is no doubt that his biology is underpinned by a thoroughgoing teleology even if it is less of an ostensible concern for him; see LENNOX (2001b) ch. 12.
XCVI
Aristotelian and evolutionary explanations compared. ‘Nothing in biology makes sense except . . .’, DOBZHANSKY (1973). ‘Nature does that which, among the possibilities’, IA 704b11; cf. GA 788b20. For optimality thinking in A. see LEROI (2010); in evolutionary biology and its formal link to the theory of natural selection see GRAFEN (2007). Individuals are the beneficiaries of Darwinian adaptations, see DARWIN (1859) p. 186 and RUSE (1980); for a discussion on how to distinguish levels of selection from levels of adaptation see GARDENER and GRAFEN (2009). Living things participate in the eternal and the divine at DA 415a25, DA 415a22, GA 731b18, GC II, 10–11; cf. the dubiously Aristotelian MM 1187a30. When A. discusses the ultimate purpose of life, he generally frames it in terms of the purpose of the soul, i.e. the physiological system that controls nutrition, growth and reproduction, as well as other functions; see Ch. LIV. I claim that the features of Aristotelian creatures are ultimately for the sake of forms/kinds. Some scholars, e.g. BALME and GOTTHELF (1992) pp. 96–7 and LENNOX (2001b) ch. 6 and pers. comm., deny this and argue that eternal survival of kinds is just a secondary consequence of the desire of individuals to reproduce. However, DA 415b2 points out that there are two senses in which we can speak of ‘that for the sake of which’. The first is ‘that for the purpose of which’, the second is ‘that for the benefit of which’. He then clearly goes on to identif
y the ‘that for the benefit of which’ as the form/kind; see also DA 416b22. It is also worth pointing out that, when speaking of a particular adaptation, A. usually does not specify whether it (say, horns) is ‘good for the individual’ or ‘good for the species’. He doesn’t need to – it is good for both. Sometimes, however, he does explicitly say that some feature is good for the species, as when discussing the features of fish life history, GA 755a30; Ch. LXXXV. In this respect, his teleology is different from Darwin’s insofar as, for Darwin, individuals are beneficiaries of adaptation; for a Neo-Darwinian, genes are; see also Ch. CI. On reductionism in Aristotelian explanations see GOTTHELF (2012) ch. 3. A. claims that it is better to exist than not exist at GA 731b30.