Abstract
Although the contributions of James Petiver to the early development of systematic natural history are widely acknowledged, he is often criticized for scientific, curatorial and even social shortcomings. This rather dubious reputation is at odds with his standing among entomologists as ‘the father of British butterflies’. Shortly before his death in 1718, Petiver published a densely packed eight-page pamphlet entitled Papilionum Britanniae. Analysis of this work, which at first sight makes an apparently exaggerated claim of accounting for ‘above eighty English butterflies’, reveals that Petiver was an original, perceptive and truly systematic entomologist, in several important respects ahead of his time.
Keywords: seventeenth century science, eighteenth century science, polymorphism, polyphenism, forms, species
Introduction
James Petiver is widely acknowledged for his relentless accumulation of natural history objects and his prolific publications based on them. Some critics have drawn attention to a meanness of spirit: ‘[his] work on British insects [was] … propelled by a collecting mania directed almost wholly to private, rather selfish ends’.1 His contemporaries complained of the careless and chaotic state of his collections: ‘Everything is kept in true English fashion in prodigious confusion in one wretched cabinet and boxes’;2 ‘[he] put them into heaps … many of them injured by dust, insects, rain’.3 And it is the case that many of his published works consist of a befuddling melange of animals, plants and other artefacts seemingly flung together without rhyme or reason.
So why is it that Petiver, arguably selfish, careless and unsystematic, has been described as ‘the father of British butterflies’4—especially when others, such as his friend and contemporary Joseph Dandridge (1665–1747), could lay equal claim to the title?5 Is Petiver's celebration justified? One reason is that, whatever the supposed shortcomings of his oeuvre, Petiver published, whereas Dandridge, for example, did not—‘publish or perish’ in action more than two centuries before that phrase was coined.
However, as explored below, if we analyse Petiver's swansong on the British butterflies, Papilionum Britanniae, his special place in entomology is demonstrably based on far more than self-promotion or romanticized myth. Each butterfly morph or variant is given its own, separate name. But in all cases the sequence of presentation, coupled with Petiver's use of ‘generic’ vernacular names to group morphs together, results in a system fully congruent with modern species-level taxonomy. He also formulated an outline higher classification for the butterflies (Papilionoidea) that was not significantly improved until the late eighteenth century, elements of which survive to this day—perhaps most notably, the family Nymphalidae. Contrary to the typical characterization of Petiver as a haphazard and careless collector, the Papilionum reveals a discriminating and observant entomologist.
Early works on butterflies published in Britain
The first British work on insects, Insectorum sive Minimorum Animalium Theatrum, was published in 1634 by Théodore de Mayerne, a French physician to James I and later Charles I.6 This Latin treatise was based on a manuscript long in circulation that included contributions from Conrad Gesner, Edward Wooton, Thomas Penny and Thomas Moffet—and, without attribution, Ulisse Androvandi.7 An English translation, attributed to Moffet, was published 24 years later by Edward Topsell.8 The text and illustrations refer to as many as 20 native species of butterflies, but the work was not solely addressed to British fauna.9 The first work exclusively about British insects appeared soon after, by Christopher Merrett.10 This is considered to include 21 species of butterflies—but the identities of some are problematic. Moreover, Merrett evidently chose to include species that, even then, must have been uncommon, omitting others that were surely well known.11 So his account cannot be regarded as complete.
Merrett's work appeared about the time of Petiver's birth. Over his life, in addition to all his other achievements, and often working with leading naturalists of his day, Petiver developed a truly comprehensive knowledge of British butterflies. His entomological contacts included, in addition to Dandridge, William Vernon, Eleanor Glanville and Eleazar Albin. Most notable, however, was his friend John Ray (1627–1705), the outstanding naturalist and systematic botanist who, incidentally, had come to serious entomology relatively late in life.12 Ray's posthumous publication Historia Insectorum13 included many British butterflies (37 species according to Dunbar14)—several made known to him by Petiver, as well as Dandridge and others.
One of Petiver's first publications commences with Gonepteryx rhamni (Linnaeus, 1758), a yellow butterfly found in Britain for which Petiver adopted the name ‘Brimstone’—a fitting colloquial name still in use today.15 Petiver was the first to apply short, memorable English names to the British butterflies in his publications, and his efforts culminated in the remarkable eight-page pamphlet analysed here, Papilionum Britanniae, published the year before his death.16 The Papilionum consists of just two pages of text and six pages of figures (see appendix, figures A1–A6). It presents many points of interest, including differences in figure numbering and plate layout, and idiosyncrasies of typography. Here I focus on Petiver's systematic knowledge of British butterflies, as reflected in this mid-Enlightenment but pre-evolutionary work. The stimulus comes from its only general statement, the subtitle or ‘advertisement’ (figure 1). The most striking question that arises is the number of species implied by ‘above Eighty English Butter-flies’. In fact, Petiver gives English names (he was working 40 years prior to Linnaeus's introduction of the binominal system) for 79 ‘different’ British butterflies, and describes two others in the same manner without actually bestowing or mentioning colloquial names. For the purposes of this analysis a total of 81 is accepted. In addition, he named an exotic species he erroneously believed to be British (see below).
Figure 1.
Papilionum BRITANNIAE Icones, Nomina, &c. Containing the Figures, Names, Places, Seasons, &c. of above Eighty English Butter-flies, being all that have hitherto been obtained in Great Britain. By James Petiver, Fellow of the Royal Society, LONDON. (Reproduced with permission. Copyright © the Natural History Museum, London.)
Before going further it is necessary to confirm that all insects included are butterflies—species belonging to the superfamily Papilionoidea, a sub-group of the order Lepidoptera. This is so; not a single moth or other insect is included. The term ‘moth’ applies to all other Lepidoptera, comprising numerous superfamilies. We also have to question whether they are based on reliable British records. The answer is undoubtedly yes, with one remarkable exception: the oft-debated ‘Hampsted Eye’. Petiver's mysterious inclusion of an Indo-Australian butterfly, Junonia villida (Fabricius, 1787), was based on a specimen supposedly found on Hampstead Heath by Albin. This curiosity has been discussed at length over many years.17
To assess the notion of 81 British butterflies, I use two works as ‘baselines’: Richard South's Butterflies of the British Isles, published just over 200 years after the Papilionum, and the Millennium atlas of butterflies in Britain and Ireland, by Jim Asher, Martin Warren, Richard Fox, Paul Harding, Gail Jeffcoate and Stephen Jeffcoate, which appeared almost 300 years after Petiver's work.18 The Millennium atlas lists a total of 72 species from Britain and Ireland in two categories: residents and regular migrants (58), and rare migrants and extinct species (14). Two species not considered are the Cryptic Wood White (Leptidea juvernica, Williams, 1946), recently recognized as a separate species found in Ireland but not Great Britain, and the locally extinct Scarce Copper (Lycaena virgaureae), for which, despite many expressions of doubt,19 there is evidence that it formerly occurred in England.20 I have not included either in my calculations as both were unrecorded from Britain in Petiver's time.
South listed 68 species as ‘British’—but this included 16 species unknown in Great Britain until after Petiver's death. Of the total 72 species listed in the Millennium atlas, 20 were unknown from Britain until after his lifetime.21 Both of these enumerations indicate a total of 52 species of butterflies authentically known from Great Britain by 1717. Given that all insects listed and illustrated in the Papilionum are butterflies, and that all but one are British, how did Petiver arrive at 81? Was his account even complete for all British butterflies then known? To answer these questions, I first present an inventory of the species included in Petiver's pamphlet.
Inventory of species in the Papilionum Britanniae
Petiver was working within a framework of divine creation. Ideas such as organic evolution and extinction were not considered. With the exception of Ray's notion that species can be determined by constant features that appear when organisms are propagated from ‘seed’, modern concepts of biological or phylogenetic species were unavailable to him. Despite this, by structuring the inventory below by what we currently consider to be biological species, his remarkable perspicacity at ‘species level’, and even foresight as a systematic lepidopterist, is revealed.
Forty-nine species of British butterflies as now recognized are listed below, numbered P1–P49, in their order of appearance in the Papilionum. The first line gives the current English name followed by its formal binomen, and an indication of sex(es) or form(s) illustrated. Then, indented, come the plate and figure numbers, the colloquial name(s) given, with (in square brackets) any earlier date than 1717 at which Petiver gave the name (spelled out in full if different), and finally Petiver's English commentary. Not included are Petiver's brief Latin descriptions and cryptic references to his own and other works (e.g. Ray). Figure 2 is a facsimile of two typical, successive entries.
Figure 2.
Text for plate 5, figures 9 and 10 of the Papilionum, giving separate names to females and males, respectively, of the Meadow Brown (Maniola jurtina). Note the short Latin descriptions, and cryptic references to Musei Petiveriani (‘Mus. nost.’) and Ray's Historia Insectorum. Here Petiver is uncertain whether these two butterflies relate to each other, as male and female, or not. (Reproduced with permission. Copyright © the Natural History Museum, London.)
The six plates of the Papilionum (figures A1–A6) present 166 figures of adult butterflies, in almost every case consisting of a pair of images linked by a dotted line (see figure 3, top). Each pair shows the upper- (dorsal) and under- (ventral) wing surfaces (with two minor exceptions). If we regard each pair as one ‘specimen’, and exclude the two images for the exotic ‘Hampsted Eye’ ([P50]), the plates depict 84 specimens of British butterflies—all but five of which are given colloquial names.
Figure 3.
Above: Petiver's Golden Meadow-eye (9, i) and Brown Meadow-eye (10, k), from a section of the Papilionum (pl. 5—see figure A5). Below: specimens of the Meadow Brown (Maniola jurtina) that survive in Petiver's collection (vol. 1, f. 25), pressed between mica sheets: left, female; right, male. (Photographer of specimens: Jonathan Jackson; reproduced with permission. Copyright © the Natural History Museum, London.) (Online version in colour.)
In preparing this list, I made extensive use of the work of Maitland Emmet, who in 1989 enumerated the British butterflies described by Petiver during 1695–1717.22 The inventory below is completed with three species recorded from Britain by 1717 that Petiver did not include in the Papilionum, and a separate entry for the exotic. These last four species ([P50–P53]) are not included in the analyses below.
British species of butterflies included in the Papilionum
P1. Black-veined White, Aporia crataegi. Female.
Plate I, figs 1 (up), 2 (un), White Butterfly with black Veins [1699], ‘Found in Meadows about June.’
P2. Large White, Pieris brassicae. 1st generation male (3,4); 1st gen. female (5,6).
Plate I, figs 3 (up), 4 (un), Great white Cabbage Butterfly [1703: The Greater White Cabbage Butterfly], ‘Appears in May, June &c.’; Plate I, figs 5 (up), 6 (un), Great Female Cabbage Butterfly [1706], ‘These are very common with the last, in Kitchen Gardens where Cabbages grow, on which they lay their eggs.’
P3. Small White, Pieris rapae. 2nd gen. male (7,8); 1st gen. female (9,10); 2nd gen. female (11,12); 1st gen. male (13,14).
Plate I, figs 7 (up), 8 (un), Lesser white Cabbage Butterfly [1703], ‘Very common in Fields and Gardens most part of the Summer’; Plate I, figs 9 (up), 10 (un), Lesser white double Spotted Butterfly, ‘This and the next are found promiscuously with the rest’; Plate I, figs 11 (up), 12 (un), Lesser white treble spotted Butterfly, ‘Differs from the last in having a third spot on the parting of the Wings’; Plate I, figs 13 (up), 14 (un), Lesser white unspotted Butterfly,‘The under [= hind or posterior] Wings of these 6 last are frosted below with a pale yellow and without Veins, which the 3 next have’.
P4. Green-veined White, Pieris napi. 2nd gen. female (15,16); 1st gen. male (17,18); 2nd gen. male (19,20).
Plate I, figs 15 (up), 16 (un), Common white, veined Butterfly, with 2 or 3 Spots [1699], ‘Common with the next in Fields and Meadows most part of the Summer’; Plate I, figs 17 (up), 18 (un), Lesser, white, veined Butterfly, ‘This has no Spots’; Plate I, figs 19 (up), 20 (un), Lesser, spotted, white, veined Butterfly, ‘Differs from 15 in having but one Spot on each upper [= fore or anterior] Wing’.
P5. Wood White, Leptidea sinapis. 1st gen. male (21,22); 1st gen. female (23,24).
Plate I, figs 21 (up), 22 (un), White small tipt Butterfly [1706], ‘I have observed this in Hampsted and other Woods in June, &c.’; Plate I, figs 23 (up), 24 (un), Small white Wood Butterfly [1706], ‘This is sometimes tipt below, but scarce seen above, N.B. I have observed amongst all these white Butterflies, the most spotted to be the Females.’
P6. Brimstone, Gonepteryx rhamni. Male (1); female (2).
Plate II, fig. 1 (up & un), The Brimstone Butterfly [1695], ‘This and the next appear amongst the first in the Spring and again in the Autumn.’; Plate II, fig. 2 (up & un); The Male Straw Butterfly [1695: The pale Brimstone Butterfly], ‘This being so near white, it often escapes as common.’ [Petiver had the sexes of this species reversed.]
P7. Clouded Yellow, Colias croceus. Male (3); female (4).
Plate II, fig. 3 (up & un), The Saffron Butterfly [1703], ‘Its Ground Saffron, with broad black Edges, seen about Deptford, Peckham, &c. from June till September.’; Plate II, fig. 4 (up & un), Spotted Saffron Butterfly [1706: Saffron Butterfly with Spotted Tips], ‘Found with the last, and differs from it in having a spotted edge.’
P8. Swallowtail, Papilio machaon. Male.
Plate II, fig. 5 (up & un), Royal William [1699], ‘Its Beauty, Size, and Tail differs it from all others. This has been caught about London and divers Counties in England, yet rarely.’
P9. Orange-tip, Anthocharis cardamines. Male (6); female (7).
Plate II, fig. 6 (up & un), White, marbled Male Butterfly [1699], ‘Found with the Female, but more particularly observed by its Saffron Tips, by which it is easily known flying.’; Plate II, fig. 7 (up & un), White marbled Female Butterfly [1699, A.305, P.33], ‘This appears in April and May, but not common.’
P10. Bath White, Pontia daplidice. Male (8); female (9).
Plate II, fig. 8 (up & un), The slight greenish Half-Mourner [1699: greenish marbled half-Mourner], ‘This is somewhat less than the next, and has fewer Spots, especially above.’; Plate II, fig. 9 (up & un), Vernouns greenish Half Mourner [1702: Vernon's half-Mourner], ‘This has also been found about Hampsted in July or August.’
P11. Marbled White, Melanargia galathea. Female.
Plate II, fig. 10 (up & un), Common Half-Mourner [1695: Our half Mourner], ‘These have near and equal Mixture of white and black, they appear about Midsummer and after in moist Meadows.’
P12. Red Admiral, Vanessa atalanta. Female.
Plate II, fig. xi (up & un), The Admiral [1699], ‘A beautiful Fly, and eminently distinguisht by a red List [= line or stripe] cross the upper Wing. Often seen in Gardens and Fields from the End of July till Autumn.’
P13. White Admiral, Limenitis camilla. Male.
Plate II, fig. 12 (up & un), White Admiral [1703: White Leghorn Admiral], ‘Found about Dullidge and Wickham near Croyden [sic]. as also at Henly [sic] upon Thames.’
P14. Silver-washed Fritillary, Argynnis paphia. Male (1,2); female (3,4).
Plate III, figs 1 (up), 2 (un), Great Silverstreakt Orange Fritillary [1699: The greater silver-streaked fritillary], ‘Differs from the next in having Streaks above on an Orange Ground.’; Plate III, figs 3 (up), 4 (un), Great Silver-streakt Golden Fritillary, ‘The Spots above more and larger, without any black Streaks, on a pale golden Ground. These appear about the beginning of July.’
P15. High Brown Fritillary, Argynnis adippe. Female.
Plate III, figs 5 (up), 6 (un), Great Silver-spotted Fritillary [1699: The greater silver-spotted Fritillary], ‘In this the Spots above are larger than the next, and below the ground between the Spots pale.’
P16. Dark Green Fritillary, Argynnis aglaja. Male.
Plate III, figs 7 (up), 8 (un), no common name given, ‘In this the Spots are less, and the Ground below greenish. This and the last appear about the midst of July.’
P17. Heath Fritillary, Mellicta athalia. Male (9,10); female (11,12).
Plate III, figs 9 (up), 10 (un), White May Fritillary [1699: May Fritillary], ‘Very like the next, but whiter below.’; Plate III, figs 11 (up), 12 (un), Straw May Fritillary, ‘Both these pretty common in Cain-Wood [Kenwood], and differ from the next in being Checker'd above.’
P18. Small Pearl-bordered Fritillary, Boloria selene. Female.
Plate III, figs 13 (up), 14 (un), April Fritillary [1717, not 1699—see P20], ‘This has many white and Silver Spots below, by which it chiefly differs from Fig. 18.’
P19. Duke of Burgundy, Hamearis lucina. Male.
Plate III, figs 15 (up), 16 (un), Vernons, small Fritillary [1699: Mr Vernon's small Fritillary], ‘It's the least of all the Fritillaries yet known. Found in several Woods round London.’
P20. Pearl-bordered Fritillary, Boloria euphrosyne. Female.
Plate III, figs 17 (up), 18 (un), April Fritillary with a few Spots [1699: April Fritillary], ‘This has but one oval Silver Spot in the midst of each Wing beneath, whereas Fig. 14 [see P18 above] has divers. Perhaps Male and Female. Frequent in Cain-Wood.’
P21. Marsh Fritillary, Euphydryas aurinia. Female (19,20); male (21,22).
Plate III, figs 19 (up), 20 (un), Mr Dandridge's midling Black Fritillary, ‘Mr. Dandridge has observed this and the next at the end of May and beginning of June in Cain Wood and the Oak of Honour Woods near Dullidge.’; Plate III, figs 21 (up), 22 (un), Small black Fritillary, ‘Differs chiefly in being less than the last.’
P22. Queen of Spain Fritillary, Argynnis lathonia. Female.
Plate III, figs 23 (up), 24 (un), Lesser Silver-spotted or Riga Fritillary [1704: Riga Fritillary], ‘Observed about Cambridge.’
P23. Glanville Fritillary, Melitaea cinxia. Female.
Plate III, figs 25 (up), 26 (un), White Dullidge Fritillary [1703: Lincolnshire Fritillary], ‘Found in the Wood, thereabouts in May.’
P24. Large Tortoiseshell, Nymphalis polychloros. Female.
Plate IIII, figs 1 (up), 2 (un), Great Tortoise-shell Butterfly [1699: The greater Tortoise-shell Butterfly], ‘A large Fly, I have observ'd them both in Spring and Autumn, they often settle on Trees, and commonly the Elm in the Wood.’
P25. Small Tortoiseshell, Aglais urticae. Male.
Plate IIII, figs 3 (up), 4 (un), Lesser or Common Tortoise-shell [1699], ‘Very frequent in Summer, and often found in Houses in the Winter.’
P26. Comma, Polygonia c-album. Overwintering (2nd gen.) female (5,6); 1st (spring) gen. male (f. ‘hutchinsonii’) (7,8); overwintering female ab. ‘variegata’ (9,10); overwintering male var. (11,12).
Plate IIII, figs 5 (up), 6 (un), The Silver Comma, ‘This and the 3 following appear about Midsummer.’; Plate IIII, figs 7 (up), 8 (un), The pale Comma, ‘This below is of an Oker Marble, and paler than the last.’; Plate IIII, figs 9 (up), 10 (un), Jagged wing'd Comma, ‘These Wings are deeper cut and more vivid; it's finely marbled underneath, with small greenish Eyes specky with black.’; Plate IIII, figs 11 (up), 12 (un), Small Comma, ‘It's very dark below, and in all Parts less.’
P27. Small Copper, Lycaena phlaeas. Male.
Plate IIII, figs 13 (up), 14 (un), Small Tortoise shell [1699: The small golden black-spotted Meadow Butterfly], ‘In Meadows and about Hedges from July till Autumn.’
P28. Grizzled Skipper, Pyrgus malvae. Male? (15,16); female? (17,18).
Plate IIII, figs 15 (up), 16 (un), Brown Marsh Fritillary [1699: Our brown Marsh Fritillary], ‘The Spots in this are larger and fewer than the next, and Found with it.’ Plate IIII, figs 17 (up), 18 (un), Small-spotted brown Marsh Fritillary, ‘The Spots in this are smaller and more distinct.’ [Sexes difficult to separate based on Petiver images; see also below.]
P29. Dingy Skipper, Erynnis tages. Male.
Plate IIII, figs 19 (up), 20 (un), Handley's small brown Butterfly [1704: Handley's brown Butterfly], ‘It's brown above and paler below, and dully marbled.’
P30. Painted Lady, Vanessa cardui. Female.
Plate IIII, figs 21 (up), 22 (un), Painted Lady [1699], ‘It usually settles on Banks and dry Ground.’
P31. Brown Hairstreak, Thecla betulae. Male (23); female (24); underside female (25).
Plate IIII, fig. 23 (up), The brown Hair-streak [1703: brown double Streak], ‘Mr. Ray has seen this and 26 in Coitu on Nettles, July 8. 1692’; Plate IIII, fig. 24 (up), Golden Hair-streak [1703: Golden brown double Streak], ‘Found with the last about Croydon.’; Plate IIII, fig. 25 (un), ‘This Figure represents the Under-side of the 2 last, which differ not below.’
P32. Purple Hairstreak, Quercusia quercus. Male (26); female (27); underside male (28).
Plate IIII, fig. 26 (up), Ray's blew Hair-streak [1702: Mr. Ray's Purple-Streak], ‘The Ground above is a dark hair brown, with a blewish Cast over it.’; Plate IIII, fig. 27 (up), Our blew Hair-streak, ‘The Ground the same as the last, but with the Inside of its upper Wings has a broad Field of vivid blew.’; Plate IIII, fig. 28 (un), ‘The 2 last are the same below.’
P33. Peacock, Aglais io. Female.
Plate V, figs 1 (up), a (un), The Peacock's Eye [1699], ‘This often appears early and continues long, it's pretty common in Gardens and Fields.’
P34. Grayling, Hipparchia semele. Female (3, c); male (4, d).
Plate V, figs 3 (up), c (un), The Tunbridge Grayling [1699: Black-ey'd marble Butterfly], ‘Because first caught in these Parts. This I take to be the Female, and appears about Midsummer.’ Plate V, figs 4 (up), d (un), Brown Tunbridge Grayling, ‘Much darker on the upper side and somewhat smaller.’
P35. Speckled Wood, Pararge aegeria. 1st gen. male (5, e); 2nd gen. female (6, f).
Plate V, figs 5 (up), e (un), The Enfield Eye [1704], ‘It being the Place I first observed them in. These appear from April till July.’; Plate V, figs 6 (up), f (un), Brown Enfield Eye, ‘Mr. Dandridge first observed this about Watford towards the End of July.’
P36. Wall, Lasiommata megera. Female (7, g); male (8, h).
Plate V, figs 7 (up), g (un), The London Eye [1699: The golden marbled Butterfly, with black eyes], ‘These I have met with from May till August, but not very common.’; Plate V, figs 8 (up), h (un), London Eye, with a brown List, ‘Found with the last, and may be Male and Female.’
P37. Meadow Brown, Maniola jurtina. Female (9, i), male (10, k).
Plate V, figs 9 (up), i (un), Golden Meadow-Eye [1699: The golden Meadow, ey'd-Butterfly], ‘Common in Meadows before Mowing, and perhaps Female to the next.’; Plate V, figs 10 (up), k (un), Brown Meadow-Eye [1699: The brown Meadow, ey'd-Butterfly], ‘Found frequently with the last in May and June.’
P38. Gatekeeper, Pyronia tithonus. Male (xi, l), female (12, m).
Plate V, figs xi (up), l (un), no common name given [1699: The lesser double-ey'd Butterfly], ‘Differs from the next in having a brown List. Perhaps Male and Female.’; Plate V, figs 12 (up), m (un), Hedge-Eye with double Specks [1699: The brown Meadow, ey'd-Butterfly], ‘Seen about Hedges in August.’
P39. Ringlet, Aphantopus hyperantus. Female (13, n), male (14, o).
Plate V, figs 13 (up), n (un), Brown 8 Eyes [1699: The brown ey'd-Butterfly with yellow circles], ‘These rarely appear before August and mostly near Rivers.’; Plate V, figs 14 (up), o (un), Brown 7 Eyes, ‘This has often underneath one Eye fewer than the last.’
P40. Small Heath, Coenonympha pamphilus. Female (15, p), male (16, q).
Plate V, figs 15 (up), p (un), Golden Heath-Eye [1699: The small Heath Butterfly], ‘Very common on Heaths, from May till Autumn.’; Plate V, figs 16 (up), q (un), Selvedg'd Heath-Eye, ‘Differs than the last, in having a brown brim or edge and can be found with it.’
P41. Chalk Hill Blue, Polyommatus coridon. Male.
Plate VI, fig. 1 (up, un), Pale blue Argus [1704], ‘I first observ'd this in Thickets near Banstead Downs, about July.’; Plate VI, fig. 2 (un), ‘Nicely differs from the last, by a small line underneath.’
P42. Common Blue, Polyommatus icarus. Male (3), female (4–7).
Plate VI, fig. 3 (up, un), Blue Argus [1699: The little Blew-Argus], ‘Very common on Heaths from June till August.’; Plate VI, fig. 4 (up, un) Selvedg'd blew Argus, ‘Found with the last, but more rare.’; Plate VI, fig. 5 (up, un) Mixt Argus, ‘It's brown above mix'd with blew.’; Plate VI, fig. 6 (up, un), no common name given, ‘Differs from fig. 4. in being less and with smaller eyes.’; Plate VI, fig. 7 (up, un), no common name given, ‘This has fewer Spots underneath than fig. 5.’
P43. Adonis Blue, Polyommatus bellargus. Female.
Plate VI, fig. 8 (up, un), Lead Argus, ‘Eyed underneath like the foregoing, and found with them.’
P44. Silver-studded Blue, Plebejus argus. Female.
Plate VI, fig. 9 (up, un), Small Lead Argus, ‘Less than the last.’
P45. Brown Argus, Aricia agestis. Female.
Plate VI, fig. 10 (up, un), Edged, brown Argus [1704: Edg'd brown Argus], ‘Its ground darker than the 2 last.’
P46. Holly Blue, Celastrina argiolus. Male (11), female (12).
Plate VI, fig. 11 (up, un), Blue speckt Butterfly, ‘I have observed this on Holly Trees.’; Plate VI, fig. 12 (up, un), Blue speckt Butterfly, with black tipps, ‘Found with the last.’
P47. Green Hairstreak, Callophrys rubi. Male.
Plate VI, fig. 13 (up, un), Holly Butterfly [1702: unnamed; 1706: Holly under green Butterfly], ‘Because I first observed it on that tree.’
P48. Small Skipper, Thymelicus sylvestris. Female (14), male (15).
Plate VI, fig. 14 (up, un), The spotless Hog [1704], no commentary in English (but see fig. 17); Plate VI, fig. 15 (up, un), Streakt Golden Hog [1704], no commentary in English (but see fig. 17).
P49. Large Skipper, Ochlodes venata. Male (16), female (17).
Plate VI, fig. 16 (up, un), Streakt cloudy Hog [1704: The chequer-like Hogg], ‘Differs from the last in being clouded with brown.’; Plate VI, fig. 17 (up, un), Cloudy Hog [1704: The Chequered Hogg], ‘This has no Slants. All these Varieties are found in the Woods, &c. about Hampstead.’
Exotic species erroneously included
[P50]. Meadow Argus, Junonia villida. Male [Indo-Pacific species].23
Plate V, figs 2 (up), b (un), Albin's Hampsted Eye, ‘Where it was caught by this Curious Person, and is the only one I have yet seen.’
Three British species known or probably known to Petiver but not included
[P51]. White-letter Hairstreak, Satyrium w-album. [Petiver: The Hair-Streak.24]
Widespread but uncommon through much of England.
[P52]. Mazarine Blue, Polyommatus semiargus. [Ray: no common name.25]
Possibly not known to Petiver—but, given the relationship between the two naturalists, he could have been aware of this rarity. Doubt has often been expressed about Ray's record, but Bretherton agreed with Ford with respect to the identification.26
[P53]. Purple Emperor, Apatura iris. [Petiver: Mr Dale's Purple Eye.27]
In Petiver's time almost certainly more widespread in England than now, but apparently only known to him from a single specimen in the Dale collection. This belonged to Samuel Dale, a neighbour of Ray, and thus the butterfly was perhaps captured in Essex.28
In addition, based on the works of Moffet and Merrett,29 it has been suggested that the Silver-spotted Skipper (Hesperia comma) was also known from Britain prior to the Papilionum.30 This, however, is now discounted.31
Summary of inventory
The Papilionum presents authentic records of 49 British species (discounting the ‘Hampsted Eye’, P50), enumerated as P1–P49 above. In addition, three species already known from Britain by 1717 were omitted, one of which, the Mazarine Blue (P52), recorded by Ray, could have escaped Petiver's attention. The other two (White-letter Hairstreak, P51, and Purple Emperor, P53) Petiver named in his earlier works, and it seems possible that their omission in 1717 was an oversight. Although this seems at odds with the care that Petiver evidently gave to this publication, currently there is no other explanation. To the 49 British butterflies in the account Petiver applied 79 colloquial names, but failed to give one for the Dark Green Fritillary (Argynnis aglaja).
Analyses: variation and butterfly diversity
In many species of insects the sexes differ in colour pattern, size and/or shape; some vary with the seasons; and other types of variation in external appearance also occur. To confirm that diverse individuals do belong to the same species ultimately depends on rearing broods from eggs laid by individual females. By the eighteenth century very little life-cycle work of this sort had been done. For English butterflies the pioneer was Eleanor Glanville (ca 1654–1709), known to have been encouraged by Petiver. Unfortunately, it is not clear how much use of her work he made in preparing the Papilionum.
The following subsections analyse variations in each of the 49 British species in the Papilionum now known to be due to polymorphism (including sexual dimorphism), polyphenism (seasonal changes) and individual variation (both continuous and discontinuous). The section continues with some observations on Petiver's illustrations and distribution records, and is completed with an analysis of his qualities as a systematist in light of the Papilionum.
Polymorphism and sexual dimorphism
Polymorphism ‘is the occurrence together in the same habitat of two or more distinct forms of a species in such proportions that the rarest of them cannot be maintained by recurrent [genetic] mutation’.32 This includes sexual dimorphism: differences between the sexes readily appreciable by superficial inspection. Such variations are usually due to genetic differences. An integrated classification for polymorphism and sexual dimorphism has recognized nine potential categories, including monomorphism as the limiting case.33 A tenth category, weak sexual dimorphism, was subsequently added.34 The British butterflies of the Papilionum represent four of these ten categories.
1. Monomorphism
The sexes of many British butterflies are hard to separate visually—discounting the fact that in many species the forewings of the sexes differ a little in shape and the abdomens in girth, and that there are differences in overall mean size. Of the species listed in the Papilionum, 18 are considered monomorphic in this sense: P8, P11–P13, P15, P16, P18, P20, P22–P25, P27, P29, P30, P33, P35 and P39. In only two of these cases does Petiver apply more than one name: the Speckled Wood (P35), which, although the sexes look very similar, exhibits seasonal variation, and the Ringlet (P39), which exhibits discontinuous variation.
2. Weak sexual dimorphism
Nine species in the Papilionum differ only slightly in outward appearance of the sexes: P1, P5, P17, P19, P21, P40 and P45–P47. Petiver gave two names to five of these, but not to the Black-veined White (P1), the Duke of Burgundy (P19), the Brown Argus (P45) or the Green Hairstreak (P47). This group thus has five names more than needed to name the separate species.
3. Strong sexual dimorphism
In this class, as in the last, both sexes have only one morph, but the sexes are more strikingly distinct. Of the Papilionum species, 17 fit here: P2–P4, P6, P9, P10, P31, P32, P34, P36–P38, P41, P43, P44, P48 and P49. Petiver gave two names to all of these, with three exceptions: the Chalk Hill Blue (P41), the Adonis Blue (P43) and the Silver-studded Blue (P44), for which he either did not have the opposite sex, or perhaps misidentified one of the sexes (the ‘blues’ are arguably the most difficult British butterflies for accurate identification). In one case he mistakenly reversed the sexes: the Brimstone (figure 4). Petiver gave more than two names to two species in this class (Small and Green-veined whites, P3 and P4), butterflies that also exhibit seasonal variation. Strong sexual dimorphism results in 14 names more than is necessary to name the species.
Figure 4.
English specimens of the Brimstone (Gonepteryx rhamni) that survive in Petiver's collection (vol. 1, ff. 9 and 10), pressed between mica sheets: left, female; right, male. The former was thought by Petiver to be the male, which he called ‘The Male Straw Butterfly’ (see entry P6). (Photographer: Jonathan Jackson; reproduced with permission. Copyright © the Natural History Museum, London.) (Online version in colour.)
4. Multiple female dual polymorphism
This category encompasses a few British butterflies with two or more female morphs not one of which looks the same as the single male morph. This applies to three Papilionum species: P7, P14 and P42. For the first two (Clouded Yellow and Silver-washed Fritillary), one might have expected Petiver to have applied three names: one for the male and two for the two distinct female forms that both species have. However, it seems that Petiver did not know of the alternative female f. ‘valesina’ of the fritillary (only common in the New Forest area), or the rarer pale ‘alba’ form of female Clouded Yellow. Thus Petiver applied only two names to both.
The third species is the Common Blue (P42), in which females range from almost all blue (male-like) to dark brown. Females have marginal bands of bright orange spots on both wings that are absent in males. In reality, females show more or less continuous variation rather than discrete morphs, but it is convenient to include this species here. Petiver illustrates the male, to which he gave one name, and four females, to two of which he gave colloquial names. Overall, this category results in four names more than needed for the species.
The Comma (P26)
The Comma (Polygonia c-album) is a remarkably variable species that cannot readily be accommodated by the above system. It exhibits seasonal variation (spring and overwintering generations of adults), slight sexual dimorphism (wing margin shape) and probably continuous variation (upper and underside colour patterns) and even discontinuous variation (possibly bimodal in adult size). This ‘cocktail’ of variation Petiver addresses by illustrating four individuals, with four names—three more than is necessary for the single biological species represented.
Seasonal variation (polyphenism)
Many butterflies produce two or more generations in a year. In temperate regions this can result in more or less subtle differences in pattern between early (spring) and later (summer) broods. Such polyphenism is an expression of phenotypic plasticity in response to different climatic conditions (e.g. light, temperature) during larval development.35
About one-third of species in the Papilionum regularly or occasionally produce multiple generations. Of these, eight are widely regarded as polyphenic: P2–P5, P10, P26, P35 and P46. Petiver gave additional names reflecting seasonal forms to four of these: both sexes of the Small White (P3), the male Green-veined White (P4), the Comma (P26) and spring and summer forms of the Speckled Wood (P35). For the Wood White (P5), in which the second generation is only partial, Petiver illustrated only the first (summer) generation. For the Large White (P2) he also illustrated only the first brood. The Bath White (P10) is a rare migrant, with the spring form exceptionally rare in Britain (it is also sexually dimorphic, and Petiver introduced two names with respect to that difference). Petiver did not illustrate the subtle seasonal differences of the Holly Blue (but he did name the two sexes, which themselves differ). Overall, seasonal polyphenism is ‘responsible’ for five more names than necessary to designate the species—but one is already counted above (P26).
Individual variation
Most butterflies also exhibit much individual variation in appearance that cannot be regarded as polymorphism or polyphenism. This can be divided into continuous and discontinuous, and is generally thought to be of genetic and/or environmental origin. Both sorts are at work in Petiver's account. Of the four names applied to the Comma (P26), variation in size is responsible for one of them. In the case of the Ringlet (P39), which varies discretely in both sexes with respect to the number of conspicuous border ocelli (‘eyes’) expressed,36 Petiver gave different names to seven- and eight-‘eyed’ individuals.
Another case is the Grizzled Skipper (P28) in which, based on the wings, the sexes can only reliably be separated by a male structure called the ‘costal fold’, absent in females. This difference cannot be appreciated in Petiver's figures. What is striking, however, is that, whichever sexes his figures really represent, they illustrate two different expressions of the variable spotting pattern of this species that affects both sexes. Richard South suggested that one (figure A3: 15,16) was a variant named ‘taras’37—but these figures show a phenotype closer to another named variant, ‘scabatella’.38 Thus in this case the additional name is due to individual variation, not sexual dimorphism. In total, individual variation is the ‘cause’ of three additional names—one of which has already been counted above (P26).
Phenology
Rather little regarding phenology of the British butterflies can be gleaned directly from the Papilionum (significantly more information can be found in Petiver's earlier publications). Two factors confound literal interpretation: the 11-day change to the Gregorian calendar in 1752 (when 2 September was immediately followed by 14 September) and climate change. Changes in plant phenology over the past 150 years are extremely complex, with many plants coming into leaf even weeks sooner than in the eighteenth century.39 A recent study on British butterflies suggests, however, that changes in insect phenology may be much more modest.40 Thus for the Grizzled Skipper (Pyrgus malvae) Petiver noted: ‘I first observed this April 30 1696 in a bog on Hampstead-Heath.’41 Apparently the first ever British record for this species, when corrected for calendar change this equates to 11 May—still well within its currently accepted flight period.42 A meaningful analysis of Petiver's phenological data is beyond the scope of this paper—but would be worth undertaking.
Illustrations
Although entomological illustration, a subject in its own right,43 is not the focus here, because Petiver's plates are so fundamental to the Papilionum, it seems appropriate to say something about them. The earliest entomological works, such as those of Ulisse Aldrovandi (1522–1605) and Moffet, were illustrated by woodcuts—with the obvious limitations of that technique. Many of the crude illustrations in Moffet's Theatrum are based on the work of Thomas Penny, whose original illustrations have been described as ‘exquisite’.44 By the end of the seventeenth century, adoption of copperplate intaglio printing led to significant improvements in quality. Eleazar Albin's A Natural History of English Insects (1720) has been said to be the first entomological book with coloured plates produced in England.45 Many of these were engraved from Albin's originals during 1713 and 1714 by H. Terasson—additionally known as a draughtsman. Terasson also engraved plates I, III and IIII of the Papilionum. Plate I is signed ‘H. Terasson delin. et sculp. London, 1715’, which suggests the possibility that, for this plate at least, he might also have created the originals himself. Closer examination of Petiver's earlier works might resolve this. The other three plates were engraved by Sutton Nicholls (1668–1729). Although plate II is unsigned, Nicholls's style is unmistakable.
On all six plates, as already noted, corresponding upperside and underside images are linked by pecked lines. Presumably this was Petiver's design, perhaps his own innovation. Although effective, this convention does not appear to have been adopted by others widely, if at all. In Albin's Natural History, for example, including those engravings by Terasson, each separate image on a plate is denoted by a lower-case letter, a convention still widely used today.
The individual Papilionum images have been described as ‘passably accurate copper-plate engravings’,46 and it has been remarked that Petiver, ‘as an artist, though careful and accurate … could not hold a candle to Albin, Dandridge or Wilkes’.47 Although realistic in relative scale, pattern and shape, close examination of the printed engravings reveals that most are more akin to excellent caricatures than fine artwork. However, I have not seen the originals—assuming they still exist.
There are coloured versions of the Papilionum. In an account published in 2007, this is discussed at some length, but inconclusively regarding whether or not the colourings, or at least some of them, were original, or only stem from Millan's 1767 second re-issue.48 Since then, Dunbar has pointed to copies with ‘pretty colouring’ which he suggests might be original, and others with ‘poor-quality’ colouring which could be subsequent.49 More research is needed. I have not seen copies with good colouring, only poor. In my opinion the poor colouring detracts from the utility of the engravings for identification.
Distribution records
The Papilionum gives particular localities for 17 species that Petiver evidently considered noteworthy, the majority around London. Most are now locally extinct in the London area, including the Grayling at ‘Tunbridge’—although three are now relatively common in suitable parts of the metropolis (Large and Small skippers, Speckled Wood). For the other 32, Petiver gave nothing specific, suggesting that he considered most to be common around the capital and home counties. This includes the Black-veined White (P1; see figure 5) and Large Tortoiseshell (P24), both now nationally extinct. More information is to be found in Petiver's earlier publications. Specific localities included in the Papilionum are listed in table 1, in alphabetical order.
Figure 5.
An English specimen of the Black-veined White (Aporia crataegi) in Petiver's collection (vol. 1, f. 1). It has been extinct in Britain since the 1920s; Petiver simply remarked of it that it was ‘found in meadows about June’ (see entry P1). (Photographer: Jonathan Jackson; reproduced with permission. Copyright © the Natural History Museum, London.) (Online version in colour.)
Table 1.
Fifteen butterfly localities mentioned in the Papilionum.
| Location | Species |
|---|---|
| Banstead Downs | Chalk Hill Blue (P41) |
| Cambridge | Bath White (P10), Queen of Spain Fritillary (P22) |
| Croydon | Brown Hairstreak (P31) |
| Deptford | Clouded Yellow (P7) |
| Dulwich | White Admiral (P13), Glanville Fritillary (P23) |
| Enfield | Speckled Wood (P35) |
| Hampstead | Wood White (P5), Bath White (P10), Large and Small skippers (P48 and P49) |
| Henley on Thames | White Admiral (P13) |
| Honor Oak | Marsh Fritillary (P21) |
| Kenwood | Heath Fritillary (P17), Pearl-bordered Fritillary (P20), Marsh Fritillary (P21) |
| ‘London’ | Swallowtail (P8), Duke of Burgundy (P19), Wall (P36) |
| Peckham | Clouded Yellow (P7) |
| Tunbridge [sic] | Grayling (P34) |
| Watford | Speckled Wood (P35) |
| West Wickham | White Admiral (P13) |
Systematics
The fundamental task of taxonomy is to provide an account of order among the diversity of organisms. According to one account, this involves enumerating the kinds of living things that exist and have existed in the past, and determining the patterns of difference and connection among them, entailing five major tasks or functions: (1) primary discrimination (recognition of basic units of biological diversity, including formal description of species and varieties); (2) systematization (assessing relationships among taxa); (3) classification (production of summary schemes that encapsulate current knowledge of taxa); (4) symbolization (creation and application of names); and (5) identification (secondary recognition: providing means to match unidentified material to the established system).50 To varying degrees all five are manifest in Petiver's Papilionum, and are assessed here under four headings: primary discrimination, higher classification (tasks 2 and 3 combined), vernacular names and identification.
1. Primary discrimination
Although at first sight it would seem that Petiver was misled by variation to establish more names than necessary for the biological species represented in the Papilionum, unerringly in every case his names are introduced in strict succession. Thus the four names for the Small White (Pieris rapae), reflecting sexual dimorphism and seasonal variation, relate to eight upper/underside figures, 7–14, on plate I. The four names for the Comma (Polygonia c-album), reflecting sexual dimorphism, polyphenism and individual variation, likewise relate to eight figures on plate IIII, 5–12. This sequential presentation is so in all cases—including the Brown and Purple hairstreaks (P31 and P32), despite Petiver noting Ray's potentially misleading account of seeing these different butterflies mating.
Thus Petiver's primary discrimination involved sorting to the level of variety, morph or form. These were named and then grouped into what we now recognize as biological species—or sorts—by the sequence of presentation and ‘generic’ naming. Hence all four name phrases referring to Pieris rapae include the generic ‘Lesser White’, and all Polygonia c-album include ‘Comma’.
Anthocharis cardamines (the Orange-tip, P9) and Maniola jurtina (the Meadow Brown, P37) are instructive. As Petiver notes of the ‘white’ (P9), when flying the male is instantly recognizable by its orange (‘saffron’) forewing tips, wholly lacking in the female. But the underside hindwings of both sexes are beautifully and similarly ‘marbled’. Thus Petiver's ‘White Marbled Male Butterfly’ is ‘found with the female’—the ‘White Marbled Female Butterfly’. The butterfly that Petiver called the Golden Meadow-eye he listed immediately before his Brown Meadow-eye, noting that the former is ‘perhaps female to the next.’ Indeed, these are, respectively, female and male Maniola jurtina (see figure 3).
The dorsal and ventral wing surfaces of many butterflies are strikingly distinct, the undersides often giving the best clues to natural affinity. Petiver clearly realized this, the Papilionum being the first work to show both wing surfaces for practically every butterfly presented—an ideal frequently not fulfilled even today. As an aside in this context, it is interesting to consider the debate regarding whether or not the mica mounts in which many of Petiver's surviving butterfly specimens are preserved are original (see figures 3–5), or the product of later curatorial effort.51 Mica mounts allow ready examination of both wing surfaces. Given Petiver's close attention to this need, it would make great sense if these mounts were indeed original. It has also been pointed out to me that the handwriting on the mount surrounds is unmistakeably Petiver's, surely confirming that these were his handiwork.
From these observations we see that Petiver gave different names to the sexes if they were distinct in appearance, even when it is quite clear that he knew they were the same ‘sort’ (e.g. the Orange-tip). Where only one form was perceived (only sometimes, owing to wanting the opposite sex or seasonal forms etc.), then the single name applied corresponds directly to our modern species. Thus, for example, the monomorphic Limenitis camilla receives just one name in the Papilionum—‘White Admiral’, the very common name by which this butterfly is still known today.
2/3. Higher classification
Each plate of the Papilionum (see appendix, figures A1–A6) has a separate title (current butterfly families represented indicated within square brackets):
Pl. I English white butterflies [Pieridae]
Pl. II English yellow, white & other mixt butterflies [Pieridae, Papilionidae, Nymphalidae]
Pl. III English spotted butterflies [Riodinidae, Nymphalidae]
Pl. IIII English spotted & streak'd butterflies [Hesperiidae, Lycaenidae, Nymphalidae]
Pl. V British eye-wing'd butterflies [Nymphalidae]
Pl. VI Small English, blue, golden & brown butterflies [Hesperiidae, Lycaenidae]
Like most early butterfly classifications,52 species are grouped inconsistently on the basis of wing colour, pattern and overall size—which has proved a wholly insufficient basis for recognizing natural higher taxa. Even so, two plates (I and V) successfully group members of a single modern family (although Nymphalidae occur mixed with other families on three other plates). Strong echoes of this six-part classification recur in the tenth edition of Linnaeus's Systema Naturae53—which for butterflies frequently refers to both Petiver and Ray.
One of Linnaeus's major subdivisions was that of the Nymphales (etymological origin of Nymphalidae)—with these in turn divided into two subgroups, one being the Nymphales Gemmati (‘in which family the wings are adorned with eyes’54)—in perfect equivalence to Petiver's ‘eye-wing'd butterflies’. The ‘eyes’ are border ocelli,55 complex homologous wing-pattern elements that only occur in, and can thus be used to group, one major subsection of the Nymphalidae.56 Petiver's eye-winged butterflies were based on a feature still very important in butterfly systematics and biology today.
4. Vernacular names
Undoubtedly Petiver is best appreciated by lepidopterists for his consistent use (and coining where required) of short, mostly very memorable, colloquial names.57 It can be argued that this was a crucial step from the old, unwieldy and certainly unmemorable Latin descriptions that passed for insect ‘names’ prior to Petiver, resolved by adoption of Linnaeus's now universal system of binominal nomenclature. Four names adopted by Petiver survive unaltered: Brimstone, White Admiral, Painted Lady (already an existing folk-name58) and Brown Hairstreak, together with various arguses and tortoiseshells. The descriptor ‘fritillary’ calls for comment. It was a name used by early lepidopterists for any butterfly with a densely spotted or chequered pattern; even today this applies—and ‘fritillary’ does not pick out a single, natural group. We can only regret that many of the other intriguing names used by Petiver, such as half-mourners, royal williams, eyes and hogs, have not passed into modern usage.
5. Identification
As already noted, Petiver's figures for the Papilionum are only ‘passably accurate’.59 For example, the wing venations for white butterflies on plate I clearly exhibit numerous inaccuracies. Reproducing good-quality figures at that time was expensive, and Petiver often seemed short of money. The need for minute detail only became apparent when exotic insects started to flood into European museums in the nineteenth century. Taking the engravings made for Petiver together with the short descriptions for each figure, an entomologist working in south-east England today would still have a high success rate in matching most butterflies encountered to Petiver's system of names—with the exception of the ‘blues’ (Lycaenidae: Polyommatinae) and true skippers (Hesperiidae: Hesperiinae). To give just one example, Petiver's description and images of the Small Skipper (P48) are insufficient to differentiate it from the Essex Skipper (Thymelicus lineola), not recognized as native to England until 1889.
Most insect taxonomists, even in a world of DNA barcodes, still rely on dichotomous keys (expert systems) for identification. In contrast, lepidopterists rarely do this, preferring instead the ‘atlas’ approach of numerous illustrations brought together to depict all the species under consideration. In this regard, Petiver's plates are contemporary in form and function. The extent to which the rare Papilionum is a model for future faunistic studies on British and other butterflies is nevertheless uncertain. The most famous eighteenth-century British butterfly and moth book, the celebrated and beautifully executed Aurelian, does not conform in any way to Petiver's ‘atlas’.60 In this respect, Petiver was far ahead of his time.
Conclusion
As stated at the outset, the single most striking question posed by the Papilionum is the excessive number of species suggested by ‘80’ British butterflies. Analysis of the inventory reveals that 49 British species are included, for which Petiver gave or introduced in effect 81 colloquial names (79 explicit ones, plus P18 and P38 male without names). The total of 32 more than the number required to denote 49 species equals the aggregate of supernumerary names reflecting weak sexual dimorphism (5), strong sexual dimorphism (14), multiple female polymorphism (4), complex variation in the Comma (3), seasonal variation (4) and individual variation (2). So what was Petiver naming, if not species?
At the time that Petiver started working, concepts of ‘species’ remained closer to Aristotle's logical system of categories than expressions of biological reality.61 Petiver died only 50 years after Redi demonstrated that maggots do not develop in dead flesh by spontaneous generation,62 and 40 years before the publication of Linnaeus's landmark tenth edition of the Systema Naturae. Another century was to pass before Darwin and Wallace's theory of evolution by natural selection revolutionized our understanding of the living world. Even then, it was decades before a clear concept of the biological (and later still, the phylogenetic) species emerged—a process in which butterflies again played a significant role.63
In the Papilionum, Petiver does not use the word ‘species’ at all. But if we look to the work of his great friend, senior and mentor, John Ray, we find something very interesting. According to Wilkins,
Ray is responsible for formulating the first explicit entirely biological notion of species, but this is not the same as saying that he presented what we would now consider a biological species concept. For him, this was a sense of ‘species’ that applied to reproducing forms—that is, living things. It was the first time a concept was proposed that applied only to the classification of living things. Prior to this, all such concepts were general-duty concepts of classification that were then applied equally to, say, books or rocks as to life.64
In my opinion this fits exactly with what Petiver was doing with the British butterflies. He named discrete forms of living organisms—one, two or more per modern ‘species’—regardless of whether these were due to sexual dimorphism (which I think he very clearly understood, but was not always sure about in the case of particular butterflies—e.g. the Meadow Brown, figures 2 and 3; the Wall, P36) or other forms of variation (such as seasonal forms, which he perhaps did not understand). Moreover, by his sequence of presentation and naming, and attention to the wing undersides, he grouped his forms into similar ‘sorts’, with a 1:1 correspondence of these groups to modern biological species. These sorts were then grouped into higher categories, corresponding very roughly to modern families with a level of success about equal to that expressed by Linnaeus 40 years later.
In conclusion, the Papilionum demonstrates that Petiver was a careful and discriminating systematic entomologist—an impression quite contrary to that created by reports of his chaotic collections and unstructured content of his early publications. His accolade as ‘the father of British butterflies’ is surely well deserved, and is also scientifically verifiable.
Acknowledgements
My most sincere thanks are due to Sue Ryder and John Chainey for access to and information about Petiver's butterfly collection, Jonathan Jackson for the images of Petiver specimens, and Hellen Pethers and her colleagues for access to literature in the Natural History Museum, London. I am also greatly indebted to Richard Coulton and Charlie Jarvis for their help, patience and encouragement, and to them and two anonymous referees for suggesting many improvements to the manuscript. Finalization of this paper has been facilitated by an Emeritus Fellowship (2017–2019) awarded to the author by the Leverhulme Trust for studies on the Nymphalidae (‘eye-wing'd butterflies’), which is gratefully acknowledged.
Appendix
Figure A1.
Plate I of the Papilionum: ‘English white butterflies’. Engraved (and drawn?) by H. Terasson, London, 1715. (From the Natural History Museum, London.)
Figure A2.
Plate II of the Papilionum: ‘English yellow, white & other mixt butterflies’. Unsigned [by Sutton Nicholls]. (From the Natural History Museum, London.)
Figure A3.
Plate III of the Papilionum: ‘English spotted butterflies’. Engraved by H. Terasson, London. (From the Natural History Museum, London.)
Figure A4.
Plate IIII of the Papilionum: ‘English spotted & streak'd butterflies’. Engraved by H. Terasson. (From the Natural History Museum, London.)
Figure A5.
Plate V of the Papilionum: ‘Brittish eye-wing'd butterflies’. Engraved by Sutton Nicholls. (From the Natural History Museum, London.)
Figure A6.
Plate VI of the Papilionum: ‘Small English, blue, golden & brown butterflies’. Engraved by Sutton Nicholls. (From the Natural History Museum, London.)
Footnotes
D. E. Allen, The naturalist in Britain: a social history (Penguin Books, Harmondsworth, 1976), p. 14.
Z. K. von Uffenbach, Merkwürdige Reisen durch Niedersachsen, Holland und Engelland (Ulm, 1753–1754), vol. 2, p. 583; translated in W. H. Quarrell and M. L. Mare, London in 1710: from the travels of Zacharias Conrad von Uffenbach (Faber & Faber, London, 1934), pp. 126–127.
H. Sloane, quoted in A. MacGregor, ‘The life, character and career of Sir Hans Sloane’, in Sir Hans Sloane: collector, scientist, antiquary (ed. A. MacGregor), pp. 11–44 (British Museum Press, London, 1994), p. 23.
E. B. Ford, Butterflies (Collins, London, 1945); M. A. Salmon, The Aurelian legacy: British butterflies and their collectors (Harley Books, Colchester, 2000).
Allen, op. cit. (note 1), pp. 14–15.
Ford, op. cit. (note 4); D. Dunbar, British butterflies: a history in books (British Library, London, 2010).
J. Neri, The insect and the image: visualizing nature in early modern Europe, 1500–1700 (University of Minnesota Press, Minneapolis, 2011).
T. Moffet, ‘The theater of insects: or, lesser living creatures as, bees, flies, caterpillars, spiders, worms, &c. a most elaborate work’, in E. Topsell, The history of four-footed beasts and serpents (2nd edn, Topsell, London, 1658), vol. 3, supplement.
Dunbar, op. cit. (note 6).
C. Merrett, Pinax rerum naturalium britannicarum (Pulleyn, London, 1666).
Ford, op. cit. (note 4).
Salmon, op. cit. (note 4).
J. Ray, Historia insectorum (Royal Society, London, 1710).
Dunbar, op. cit. (note 6).
J. Petiver, Musei Petiveriani Centuria Prima, Rariora Naturae (Smith & Walford, London, 1695), p. 3.
J. Petiver, Papilionum Britanniae (Royal Society, London, 1717). Available from: https://www.biodiversitylibrary.org/page/55873901#page/260/mode/1up (accessed 28 March 2019).
R. I. Vane-Wright and W. J. Tennent, ‘Whatever happened to Albin's Hampstead Eye?’, Entomologist's Gazette 58, 205–218 (2007); R. I. Vane-Wright and W. J. Tennent, ‘Colour and size in Junonia villida (Lepidoptera, Nymphalidae): subspecies or phenotypic plasticity?’, Syst. Biodivers. 9, 289–305 (2011).
R. South, The butterflies of the British Isles (2nd edn, Warne, London, 1921). J. Asher, M. Marren, R. Fox, P. Harding, G. Jeffcoate and S. Jeffcoate, The millennium atlas of butterflies in Britain and Ireland (Oxford University Press, 2001).
A. M. Emmet and J. Heath (eds), The moths and butterflies of Great Britain and lreland, volume 7, pt 1: Hesperiidae-Nymphalidae: the butterflies (Harley Books, Colchester, 1989).
M. J. Perceval, ‘The Lepidoptera of Henry Seymer (1767–1785)’, Entomologist's Gazette 34, 215–227 (1983); M. J. Perceval, ‘Our lost coppers (Lepidoptera: Lycaenidae)’, Entomologist's Gazette 46, 105–118 (1995); R. I. Vane-Wright and H. W. D. Hughes, The Seymer legacy (Forrest Text, Tresaith, Ceredigion, 2005); A. Barker and R. I. Vane-Wright, ‘Did Henry Seymer obtain Lycaena dispar (Lepidoptera: Lycaenidae) from Hampshire?’, Entomologist's Gazette 58, 119–125 (2007).
Emmet and Heath, op. cit. (note 19).
A. M. Emmet, ‘The vernacular names and early history of British butterflies’, in Emmet and Heath, op. cit. (note 19), pp. 7–21.
Vane-Wright and Tennent (2007), op. cit. (note 17).
J. Petiver, Gazophylacii naturae et artis: decas secunda (J. Petiver, London, 1703), pl. 11, fig. 9.
Ray, op. cit. (note 13).
R. F. Bretherton, ‘Cyaniris Dalman’, in Emmet and Heath, op. cit. (note 19), pp. 166–168. Ford, op. cit. (note 4).
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