The Wonderful Mr Willughby

by Tim Birkhead

  I’m convinced that Willughby and Ray missed a trick by not paying more attention to bird-keeping. The English edition of the Ornithology, as we’ve seen, did include a considerable amount of bird-keeping information – mainly from Aldrovandi’s Ornitholigae, Olina’s Uccelliera and Gervase Markham’s Hunger’s Prevention – added at Martin Lister’s suggestion, but only after the main research and writing was complete. This means that even though the Ornithology includes references to some intriguing aspects of bird biology derived from cage birds, including the acquisition and function of song, this information is incidental and is not fully integrated into the book’s main themes of identification and classification.28

  You can tell from their correspondence that Ray was reluctant to include this extra material, but Lister was convinced it would increase the book’s popularity, and Willughby’s widow, it seems, was hoping that the book might yet yield a profit. Lister clearly recognised bird-keeping as an important aspect of popular culture – why else suggest including a section on this? – and he was sufficiently perceptive to realise that it could reveal some new insights. But Ray either failed to see the point, or was in a hurry to complete the English edition.

  Hidden away in the pages of the Ornithology are other questions posed by Willughby. Here are just two examples of queries that do not appear on Ray’s list.

  The first relates to moult: the annual replacement of feathers. Ray writes in the Ornithology:

  It may be worth the while to enquire, why birds do yearly moult their feathers? Mr Willughby supposes that there is the same cause of the casting of feathers in birds, that there is of the falling off of the hair in men and other animals…29

  So far so good. Two facts, both correct, are established: that the moulting of feathers and hair is an annual event, and that it is the same in birds and mammals. But then, as Ray tells us, Willughby goes on to say:

  … upon recovery from fever or other disease, or upon resection after long abstinence. For in cock-birds the heat and turgency of lust is, as it were, a kind of fever, and so in the spring-time their bodies being exhausted by the frequent use of venery, they become lean: but in the hens the time of sitting and bringing up their young answers to a disease or long abstinence, for at that time they macerate themselves by hunger and continual labour. When these times are over both sexes returning to mind their own bodies and feed for themselves, do in short time recover their flesh and grow fat again, whereupon the pores of the skin being dilated the feathers fall off.30

  Willughby’s idea – and it is a rather strange one – is that the loss of feathers (and hair in humans) occurs when an individual recovers its health following a period of stress, which in turn could be a consequence of disease; too much sex in the case of males; or among female birds the energetic cost of incubation, brooding and chick-rearing. What Willughby means by ‘upon resection after long abstinence’ could either be the resumption of eating after fasting, or the resumption of sex after abstinence. His statement seems to be a badly remembered or misconstrued account – and certainly not a quote – of something Aristotle said. If it is, I have been unable to find it. Instead, what I discovered in Aristotle’s books on Generationand the History of Animals was a presumed link between the growth and loss of hair with lust, but also the similarity between the falling off of hair and the falling of autumn leaves in Aristotle’s desperate attempt to understand the appearance, disappearance and greying of human hair.31

  Willughby is correct, however, in stating that moulting follows reproduction – although this was already well known by numerous medieval writers. Such a link is particularly obvious in male ducks when they exchange their bright breeding plumage for more cryptic garb, as they, and the females, shed their flight feathers, thereby becoming flightless for a few weeks in the autumn until their new feathers have regrown.

  Willughby was not the first to discuss moult. Frederick II, in his thirteenth-century account of falconry, has quite a lot to say about it, and is actually much more perceptive than Willughby. But it is extremely unlikely that Francis could ever have seen Frederick’s account because it was not published until long after Willughby and Ray had died.32 Nonetheless, other falconry treatises that would have been available to Willughby (and Ray) mention moult, not least because it was such an important aspect of maintaining raptors in good condition. But it seems that neither Willughby nor Ray originally consulted any of these, and Ray did so only after Francis was dead when he added an account of falconry to the English translation of the Ornithology. Even then, Ray writes very little about the loss and replacement of feathers.

  The absence is curious because moulting, or mewing, as it was known, was a critical period in the falcon’s annual cycle. Falconers were very familiar with the pattern of feather loss and replacement in their birds: in fact, the terms ‘moult’ and ‘mew’ both come from the Latin muto, meaning ‘to change’, hence the mews as a place where hawks were housed while they moulted. One of the books that Lister encouraged Ray to cite in the English edition of the Ornithology was Latham’s Falconry or the Falcons Lure or Cure, published in 1615. Latham’s account includes one fascinating piece of advice that I’m surprised Ray did not comment on. It was the falconers’ practice of feeding their moulting birds on ‘kirnels’, the thyroid glands of sheep that were thought to help birds through the moult. This sounded like another old wives’ tale to me, one that Thomas Browne might have considered a ‘vulgar error’ and been keen to debunk, but in fact it works. The sheep’s thyroid gland contains the hormone thyroxin that in sheep controls their metabolism, but in birds stimulates moult and feather growth. Latham of course had no notion of hormones, but this was clearly an effective bit of folklore.33

  I suspect that because moulting is a process lasting weeks, or sometimes months, it was a subject that Willughby and Ray were unable (or unwilling) to investigate because it would have entailed keeping birds. Had they had cage birds, or even falcons, they would have had a better grasp of the process. Perhaps also if Willughby had appreciated that the pattern of moulting – the order in which wing feathers, for example, are shed and regrown – differs so much between different groups of birds, and if he had recognised that these patterns may have been a potential clue to the classification of birds, he might have taken more of an interest. Certainly, this is exactly what he did with insect metamorphosis, but then keeping insects was rather easier than keeping birds. It was the idea that moulting patterns might reveal hitherto unidentified taxonomic relations that attracted Erwin Stresemann and his wife Vesta to this area of ornithology in the 1950s. But, as must quickly have become apparent, moult did not inform classification, and their 1966 book Die Mauser der Vogel (The Moult of Birds) includes only a few pages on taxonomy.34

  Willughby’s list of ornithological queries must have once been longer than that which appears in the Ornithology. In the section on the common (or grey) heron, Ray states that ‘Gessner counts but eleven vertebres [sic] in the neck; I observed fifteen, of which the fifth has a contrary position, viz, is reflected upwards.’ Ray adds a note in the margin, saying ‘In another place Mr Willughby puts it among his queries, whether the five upper vertebres [sic] in the neck of a heron be reflected the contrary way.’ This sounds like a gentle admonishment that asks whether Willughby means just the fifth vertebra, or the upper five vertebrae that are reflected upwards? It may be because of that ambiguity that Ray did not include this particular query in the main list of queries. Given that several of Willughby’s other queries elicited responses from Ray, it is interesting that he doesn’t tell us the true answer, which makes me wonder whether he knew, and therefore whether anyone else knew at the time. Indeed, I think that Willughby may have been the first to describe these unusual vertebrae in the necks of herons. It is also intriguing that neither Willughby nor Ray say anything about the inflected vertebra being responsible for the distinctive kink in the heron’s neck, both of which seem such obvious questions, albeit in hindsight.

 
The numbering of cervical vertebrae in the grey heron by Willughby is actually correct, except that like some other authors, he did not count the first rather small vertebra, the atlas – the one immediately adjacent to the skull. He therefore identifies the special vertebra as the fifth one, whereas if you count the atlas as number one, it is the sixth. We also know that by being longer and articulating with the next vertebra at a different angle, it is this sixth vertebra that is responsible for the neck’s distinctive kink. And that, acting rather like a spear-thrower, allows the heron (and several of its close relatives, including egrets, bitterns and the darter) to more effectively stab its prey.35 I am intrigued to know how Francis Willughby made this vertebral discovery, for in a routine dissection, that is, without removing the muscle, the orientation of the neck vertebrae is not obvious. Rather, I suspect that he noticed the unusual bones while examining some of the mounted skeletons he and his colleagues encountered during their continental travels.36

  The kink in the heron’s neck, resulting from a ‘reversed’ vertebra, identified by Francis Willughby.

  Overall, those two pages of Willughby’s questions allow us to detect someone with an eager, enquiring mind; someone pushing the boundaries and a little careless perhaps of staying within the limits of his and Ray’s original aims. As today’s researchers know all too well, being curious about the natural world and constantly asking questions is an essential aspect of being a biologist. It is what Darwin identified in his brief autobiography as one of the attributes that made him what he was. I still find it puzzling that John Ray was so obviously vexed by Willughby veering into unknown territory, and I can only think that it was because, in his own inimitable way, Ray had a very clear vision of what the Ornithology had to do, and was worried that his colleague was trying to fly before either of them could walk. Those questions, however, suggest to me that had Willughby lived longer he would have gone on to think and write about birds in an extraordinarily novel way, with much more emphasis on their lives than their bodies.

  Throughout his adult life Francis Willughby seems to have had a weak constitution. At Trinity College his tutor, James Duport, was acutely aware of this and penned poems urging him to slow down and take care of himself. Although those poems were written after Francis had left Cambridge, they were clearly based on his time there and I can only presume that Duport also told Francis to his face that he should work less hard. But of course, if that’s the way one is, it is hard to stop doing what one loves.

  Willughby’s first bout of serious illness that we know of occurred in June 1662 when at the age of twenty-seven he was returning from Wales with John Ray and Philip Skippon. At Malvern, Francis became unwell and needed two days of bed rest and fasting before the ‘heat of the distemper’ subsided sufficiently to allow him to return home.

  We next hear of Francis being ill on several occasions between 1668 and 1671, in his daughter Cassandra’s account of his life written long after her father’s death. She relates, presumably based on family diaries, how he suffered violent fevers, and sought relief by taking the waters both at Astrop and at Cleehills near Ludlow while staying with the politician Sir Job Charlton.

  Francis was taken seriously ill again in 1669 while he and Ray were visiting John Wilkins in Chester. We know little about this particular bout of illness other than its debilitating nature. The following year there was another bout and on 28 April 1670, Ray wrote to Martin Lister saying that ‘Mr Willughby is I fear, fallen into a tertian ague.’ It may have been Willughby’s recovery from this malady, or possibly another one, that Ray told Lister about, in a letter dated December 1670, to which Lister replied saying: ‘I am very glad Mr Willughby is neer well again, and I thank God for his recovery, and doe heartily pray a continuance of good health to him. Methinks he is very valetudinary, and you have often alarmed me with his illness …’37

  A year or so of reasonable health followed his recovery, but all was not well. On 10 February 1671 a distant relative, Sir William Willoughby of Selston, died, leaving Francis as part-heir to a considerable estate at South Muskham and Carlton in Nottinghamshire, lands worth £1,000 a year (equivalent to around £80,000 today). It wasn’t, however, straightforward. Part of the family estate, by earlier legal arrangements, descended to Sir William’s sister and her husband, Beaumont Dixie. They had expected to inherit the bulk of Willoughby’s lands and money, and argued that Sir William had not been of sound mind when he made the will. Worse still, and revealing the man he was, Dixie went into the house before his brother-in-law had even died, removing papers and goods and helping himself to several thousand pounds in cash. As soon as Sir William died, Dixie placed padlocks on the doors to prevent the executors from making their inventory.

  Francis was summoned, and went to Selston with John Ray where they met the executors and sought legal advice. In a massive understatement, Ray described Dixie as being ‘of a contentious spirit’, and in a letter to Martin Lister he described how Francis was so preoccupied by this conflict with Dixie that he had little time for his studies: ‘The estate bequeathed him, will create him a great deal of trouble, and cost much money, and yet what the issue may be is uncertain, by reason of an errour in the will.’38

  Ray was correct: the situation was complicated, but one whose long-term advantages were considerable for Francis’s family, so it isn’t surprising that he committed a huge amount of time and emotional energy to resolving the case. At one point in 1671, knowing of Dixie’s ‘designs to get the will from him’, Francis was forced to ride the 140 miles to York at high speed to ‘get the will proved’. Later that same year he had to go to York again, to collect the will and take it to London. Travelling by coach via Nottingham and Haslington in Cambridgeshire, Francis picked up his brother-in-law, Sir Thomas Wendy, on the way, to help him defend his case. It was physically exhausting and financially crippling. When the accounts were drawn up later, it became clear that Willughby had spent £1,509 9s 2d – over £100,000 in today’s money – on the case. Yet Dixie didn’t give up, the case dragged on, and it was not until the eighteenth century that the inheritance was eventually secured.39

  Francis and Emma’s second son, Thomas, was born in April 1672. This was an occasion for celebration and a wonderful distraction – albeit briefly – from the stress of contesting the will, running the estate, and failing to be able to get on with his studies.

  In early May, Willughby was in London at the Royal Society, planning another botanical tour to the West Country, and able to comment on a letter sent in by Martin Lister on parasitic worms. Francis told the Society how he had regularly encountered such worms in many of the animals he had dissected – including a cormorant chick in the Netherlands that contained a selection of black worms – adding that he had found them both in the gut and also ‘lying loose in the cavity of the belly’. Parasitic worms, we now know, most often inhabit the gut, but when the fish, bird or quadruped host dies, the worms attempt to escape, burrowing through the gut and into the abdominal cavity. Francis promised to send the Society the names of all those animals in which he had found worms, but illness intervened, the plant-hunting trip was abandoned, and the list of parasite hosts was never sent.40

  On the third day of June 1672, Francis awoke next to his beloved wife, knowing that he was sick once more. Dr Anthony Hewitt, a Padua-trained physician, called from Lichfield, would have taken Francis’s pulse, examined his urine, removed several ounces of blood, and probably prescribed a purge and an emetic – all in the hope of restoring the imbalance between Francis’s humours. Seventeenth-century medicine knew no better and had precious little to offer in the way of either relief or reassurance.

  Two weeks after the first attack, Francis acknowledged to John Ray that he was not going to recover. They discussed what was to become of their joint endeavours, with Ray reassuring Francis that he would pull everything together and ensure that all would be published. Francis told him that the insect work – the caterpillars, the little butterfly maggots, th
e beautiful little bees in the willow wood – were what, above all else, he wanted in print. He also asked Ray to look after his sons’ education, and Ray agreed.

  On 24 June Francis signed his will, which among other things declared him a Protestant ‘utterly detesting and abhorring all the idolatory, abominable errours and ignorance that is the Popish religion’, and barring inheritance by any descendant who was a Catholic. Francis made provision for his family, optimistic that the Dixie affair would be resolved in their favour, but prudently making provision in case it wasn’t.

  He must have known the end was near. Every day the family – the children now dressed in black, anticipating the inevitable – and John Ray came to his room, and each day they must have later walked away with a sense of hopelessness. The maids sat in attendance day and night. Emma’s drawn and tear-stained face must have upset Francis, but as though to reassure his wife, he admitted to her and Ray that if it pleased God he was content to leave this world.

  And so he did, on the morning of 3 July 1672, thirty days after first becoming ill. He was just thirty-six.

  Many years later Emma recounted the events of those dreadful days to her daughter Cassandra, who recorded them thus:

  He was seized … with a plurisie, which terminated in that sort of fever which phisitians [physicians] call catarhalis, which illness he labored under a month. He very soon apprehended his own danger … expressed great concern about leaving his children and parting from my mother [Emma, his wife] who he had lived with about four years and a half – declaring that during this entire time no unkind word had passed between them.41