[page] 300

J Ornithol (2004) 145: 300–320 DOI 10.1007/s10336-004-0043-8

ORIGINAL ARTICLE

Frank D. Steinheimer

Charles Darwin's bird collection and ornithological knowledge during the voyage of H.M.S. "Beagle", 1831–1836

Received: 19 April 2004/ Revised: 10 May 2004/ Accepted: 10 May 2004 / Published online: 25 June 2004

© Dt. Ornithologen-Gesellschaft e.V. 2004

Abstract This paper analyses Charles Darwin's bird collection and the ornithological knowledge he derived from it during the voyage of H.M.S. "Beagle". Darwin collected 468 bird skins, 10 detached parts of the lesser rhea, and the nests and eggs of 16 different taxa as well as 14 whole birds and 4 parts of birds which he preserved in spirit. He labelled these specimens with a number tag only, cross-referring the number to a notebook entry. Partly because of his limited orni-thological knowledge and partly because he was con-fronted at times with entirely unknown birds, Darwin was often unable to apply the correct generic designations and gave his South American specimens English and Spanish names from literature and the local tongues, as well as the scientific generic names of European birds. Back home, it was John Gould, the prominent ornithologist of the Zoological Society of London, who made sense of Darwin's collection, among his many other scientific achievements correctly identifying the Galapagos finches as a group of closely related birds. Darwin's bird collection did not receive much attention in the latter part of the 19th century. Most of the specimens had their original labels removed and replaced by ones of the custodian institution. Today, original Darwin specimens stemming from the "Beagle" voyage are to be found in at least eight different institutions, but almost half of the bird specimens Darwin collected on the "Beagle" voyage are not accounted for. The appendix to this paper lists for the first time all the birds which Darwin collected during the voyage. Darwin's famous book On the origin of species hardly draws upon any ornithological examples from his voyage on the "Beagle". Nevertheless, Darwin contributed much to ornithology. His collection contained 39 new species and subspecies of birds, mainly described by Gould, and some birds from populations now extinct, and he also made a few very good field observations, published in the sections of The Zoology of the Voyage of H.M.S. Beagle dedicated to birds.

Keywords Bird collection Charles Darwin Ornithology Voyage of the "Beagle"

Introduction

Charles Darwin (1809–1882) is often seen as an experi-enced ornithologist with an expert knowledge of birds. In particular, his discovery of a group of 'finches' (Family Emberizidae) on the Galapagos Islands in September and October 1835, now commonly named 'Darwin's finches',¹ cemented his reputation as a field ornithologist par excellence (inter alia Heberer 1959, pp. 33–35; Lack 1961, pp. 9-10; Barlow 1963, p. 261 (footnote 1); de Beer 1963, p. 132; Moorehead 1969, pp. 196–205; Grinnell 1974, pp. 260–261; Schmitz 1983, pp. 56–60; Wehner and Gehring 1990, p. 559; Curio 1993,

F. D. Steinheimer

Bird Group, Department of Zoology,

Natural History Museum, Tring, UK

¹The term 'Darwin's finches' was first used by Lowe 1936 (cited in Sulloway 1982b, p. 45) and established by Lack (1961, p. 14), with the argument that "the term 'Galapagos finch' is less satisfactory, since one species, namely Pinaroloxias inornata, occurs not in the Galapagos, but on Cocos Island [...]." In fact, Darwin himself had only encountered some of the finch species. In this paper, both Galapagos finches and Darwin's finches are used, according to prevailing usage.

Dedicated to Prof. Dr. Ernst Mayr

Communicated by F. Bairlein

F. D. Steinheimer (&)

Museum fur Naturkunde der Humboldt-Universitat zu Berlin,

Germany, Sylter Strasse 18, 90425 Nuremberg, Germany

E-mail: franksteinheimer@yahoo.co.uk

Tel.: +49-30-20938512

Fax: +49-30-20938528

p. 344; cf. Sulloway 1982b, pp. 6–7). This is reflected in passages such as the following by Lack (1961, p. V), who wrote in the preface to Darwin's Finches: "...in 1835, Charles Darwin collected some dull-looking finches on the Galapagos Islands. They proved to be a new group of birds and, together with the giant tortoises and other Galapagos animals, they started a train of thought which culminated in the Origin of Species, and shook the world." Moorehead (1969, p. 202) added: "...it was the number of different species of finch, and the variety of their beaks, that so amazed Darwin."

The general public still sees birds as Darwin's major clue to evolution. Rediscoveries in museum collections of original 'finches' from Darwin's collection made headline news 160 years after the voyage (Anonymous 1996; Clausen 1996; Field 2003), while Darwin's own Galapagos finches were the biggest visitor attraction at a recent touring exhibition (1999–2002) on the early voyages of discovery (Rice 1999).

However, since Kottler (1978) and Sulloway (1979, 1982a, 1982b), the myth of 'Darwin and his finches' has been shown to be just that. We now know that in the field Darwin did not appreciate the close relationship between the different species of finches on the Galapagos Islands. In fact, he actually believed he had collected specimens of five different (sub-) families—finches, grosbeaks, American blackbirds, wrens and war-blers—although he did not rule out that the first three at least might be closely related. Darwin did not discuss evolution during his visit to the Galapagos Islands, nor did he subsequently use 'Galapagos finches' in his book (Darwin 1859) On the Origin of Species (Kottler 1978; Sulloway 1979, 1982a, 1982b; Futuyma 1990; Mayr 1994; Quammen 1997; Larson 2001; Keynes 2002, 2003). To him, these birds remained even in latter years "undistinguishable from each other in habits" (Darwin in Gould et al. 1841, p. 99).

Therefore, the question as to what Darwin actually knew about, and contributed to, ornithology seems to be justified (cf. Bourne 1992). Moreover, how much did Darwin actually draw on ornithology from the "Beagle" voyage in developing his theory of evolution?

While undertaking a database project for the Natural History Museum, London/Tring, the author had the opportunity to assemble data on Darwin's entire avian collection. The information which this brought to light on the taxa involved, combined with Darwin's ornitho-logical notes (Barlow 1963; Keynes 2000), permitted a thorough evaluation of Darwin's ornithological knowledge during the voyage of H.M.S.² "Beagle" in 1831– 1836. More than half of Darwin's bird specimens were

²H.M.S. stands for His Majesty's Ship (more exactly Surveying Sloop; cf. Bourne 1992). The "Beagle" was built in 1820 and named after the dog. The ship had already been used for a previous voyage to Tierra del Fuego and Patagonia in 1826–1830 before Darwin's famous 'Voyage of H.M.S. "Beagle". Any use of the term 'Voyage of H.M.S. "Beagle" in this paper always refers to this latter, second journey of the "Beagle".

301

traced during this research; data for additional birds were added from literature, since all of Darwin's orni-thological notes have survived as well. "Lucky we are that Darwin kept up [...] collecting throughout his life. If he had not collected and saved specimens, notes, manuscripts, letters, and annotated books, our under-standing of his life and motivations would be close to nil" (Porter 1985, p. 974).

Darwin and ornithology before the voyage

No Darwinian bird collection of any kind which pre-dates the voyage of the "Beagle" has survived. In all likelihood there never was one, except for a few eggs and perhaps nests. One can only surmise what ornithological knowledge Darwin would have acquired during his teens and subsequent studies in Britain by looking at his notes and collections made during the early stages of the voyage, and from autobiographical sources (Barlow 1958; Burkhardt and Smith 1985, 1986). Darwin started collecting natural history objects, including birds' eggs, as a young boy (Steinmuller and Steinmuller 1987, p. 29). He apparently read Gilbert White's (1789) The natural history and antiquities of Selborne and wondered why every gentleman did not become an ornithologist (Barlow 1958, p. 45). During his studies, he certainly collected plants, beetles, fossils and geological objects (Junker 2001), focussing, from when he attended Cambridge, on beetles and plants (Porter 1985). He was also a keen hunter and kept a record book on what he had seen and shot (Steinmuller and Steinmuller 1987, pp. 31, 47).

Darwin learned how to prepare bird specimens at the museum in Edinburgh (Desmond and Moore 1995, pp. 42, 58). His teacher there was the former travel servant of Charles Waterton (1782–1865), a famous British collector, ornithologist and taxidermist. Darwin spent some time studying the bird galleries of the Edinburgh, Cambridge and London museums. He wrote retrospectively that he also read bird books housed in the university library of Edinburgh during this period (Steinmuller and Steinmuller 1987, p. 45). While at Edinburgh University in 1825–1827, Darwin attended lectures given by John James Laforest Audubon (1785– 1851) on the habits of North American birds (Keynes 1997, p. 461). One can therefore assume that Darwin had all the tools needed in the field by an ornithologist of his day, most importantly being able to shoot a bird and preserve its skin. However, Darwin was never taught, not even by his great mentor Prof. John Stevens Henslow (1796–1861) in Cambridge, what data should be noted down on bird specimen labels. This lack was later to prove a problem.

By the time of his departure a few weeks short of his 23rd birthday, Darwin would probably have known most common British and some exotic birds. He had not, however, acquired thorough identification skills for all British birds, nor would he have encountered all

[page] 302

British species, especially those not commonly shot on hunts and offered at markets, such as nocturnal birds and rare passerines. For example, Darwin was unable to identify a short-eared owl (Asio flammeus) when con-fronted with the taxon on East Falkland Island. Even worse were Darwin's identification skills for the different plumages of seabirds and waders, which he sometimes attributed to the wrong genus. He did not know the scientific names of all British birds and had no special knowledge of the scientific names of exotic species, lacking the necessary years of training in the field as well as in reference collections.

Generally, one can say that during his youth and student years Darwin's naturalist's eyes were turned more often to the soil then to the sky (cf. Desmond and Moore 1995). Nevertheless, he was well equipped to record, but not to identify, the avifauna of the regions to be visited during the voyage of H.M.S. "Beagle". Henslow commented in a letter to Reverend George Peacock (1791–1858) that Darwin is not "a finished Naturalist, but as amply qualified for collecting, observing, & noting anything new to be noted in Natural History" (Barlow 1967, p. 30). Or as Porter (1985, p. 977) phrased it: "Charles Darwin was as prepared to be the naturalist of the "Beagle" as any contemporary university graduate could have been." Darwin, however, had two further advantages: from his youth onwards he had been an avid collector of any kind of object, and his lack of specialization meant that he was not narrow-mindedly interested in only one single subject. Secondly, Darwin had learned Spanish in his enthusiasm to follow Alexander von Humboldt's (1769–1859) example and explore Tenerife (Steinmuller and Steinmuller 1987). Thus he was probably able to understand at least parts of the stories told to him by locals in South America regarding the native bird life, although his recourse to interpreters during the voyage (Barlow 1933, p. 30) suggests that his Spanish may not have been particularly good.

When Darwin accepted a place on H.M.S. "Beagle" "more as a companion [of the Captain] than a mere collector" (Barlow 1967, p. 30), he nevertheless busily assembled and acquired the books, tools and material for preparation which he believed he might need for the long journey (Keynes 2002). While Captain Robert FitzRoy (1805–1865) might have viewed Darwin more as the accompanying gentleman, his former Cambridge professors were keen to stress "the importance of his collections [...]" (Porter 1985, p. 977). The ship's library contained a wonderful set of the most modern travel guides and natural history books of the time. The following (partly) ornithological works would have been most useful for identifying birds in the field (cited from Burkhardt and Smith 1985, pp. 553–566; some books were subsequently sent out to the "Beagle" during the voyage): Blainville et al. (1834); Burchell (1822, 1824); Cuvier (1817, 1829); Griffith et al. (1828); Forster (1778); King (1827, MS to King et al. 1839); Labil-lardiere (1799–1800); Lesson (1828); von Wied-Neuwied

(1820); Quoy and Gaimard (1824); Saint-Vincent et al. (1822–1831); and Spix and Martius (1824). Darwin also had access to Molina (1809)³ not listed in Burkhardt and Smith (1985), since there are references to the work in his zoology notes (Barlow 1963; Keynes 2000). Unfortunately, the library lacked the book of Azara (1802–1805), which would have been of much help for identifying some birds of the Rio Plata region (cf. Darwin 1870; Sulloway 1982c).⁴ Darwin was also missing some additional publications on the avifauna of South America such as Lichtenstein (1823), Vieillot (1816) and Wagler (1827), but their implication for field ornithology would have been small.

Darwin's ornithology during the voyage of H.M.S. "Beagle"

I am spending September in Patagonia, much in the same manner as I should in England, viz. shooting (Darwin in Barlow 1933, p. 100).

H.M.S. "Beagle" set off on 27 December 1831 and returned nearly 5 years later on 2 October 1836 after having charted the coast of South America and cir-cumnavigated the globe. Darwin's interest in birds during the voyage was marginal (Bourne 1992). A letter home to his distant cousin William Darwin Fox (1805– 1880) on 23 May 1833 reveals his attitude towards ornithology: "You ask me about Ornithology; my labours in it are very simple.- I have taught my servant to shoot & skin birds, & I give him money.- I have only taken one bird which has much interested me [...]" (Burkhardt and Smith 1985, p. 316, cited in Bourne 1992). The bird Darwin was referring to was the least seed-snipe Thinocorus rumicivorus⁵; Barlow 1963, pp. 211, 278; Bourne 1992). While in the vicinity of Rio de Janeiro on 4 June 1832, Darwin wrote in his diary of the local hunters, whom he joined in shooting deer and other mammals, that "they shoot parrots & Toucans &c. [between hunting mammals]. I soon found this very stupid & began to hunt my own peculiar game" (Barlow 1933, pp. 66–67). Darwin prepared only eight birds for his collection, and there was not a parrot or a toucan among them.

Darwin himself wrote in his autobiography (Sobol 1959, p. 71) that his love for hunting diminished after the first 2 years of the journey, and he began more and more often to entrust his gun to his servant Syms Covington

³The version Darwin consulted was the English translation of the Italian original (cf. Sulloway 1982b). The taxonomically important first descriptions of birds are found in Molina (1782). ⁴The references to Azara (1802–1805) in Darwin's notebooks and Gould and Darwin (1838, 1839a, 1839b, 1839c) and Gould et al. (1841) were added after the voyage (Sulloway 1982c). ⁵See Appendix: 'Charles Darwin's bird collection from the voyage of the H.M.S. "Beagle" at http://www.do-g.de, 'journals '– Supplementary information to published papers.

[page] 303

(c. 1816–1861; cf. Armstrong 1992, p. 135). Darwin believed that shooting would take too much of his attention away from investigating the geology of the countries visited. While Darwin concentrated on geol-ogy, invertebrates and plants, Covington collected only vertebrates. Covington's birds were entered into Dar-win's specimen lists without any indication as to who the actual collector was. It is probable that from mid May-1833 onwards Syms Covington shot most of the birds.

Darwin's Zoology Notes (published by Keynes 2000) show the same bias. While much space is dedicated to ornithology in the first two-thirds of the journey (espe-cially during his stay in Uruguay and Argentina), later notebook entries speak much more about marine invertebrate life such as corals, crustaceans and mol-luscs. However, Darwin's real love, despite his zoologi-cal background, was geology: "Geology & invertebrate animals will be my chief object of pursuit through the whole voyage" (Barlow 1967, p. 54; cf. Darwin 1851, 1890a, 1890b). Except in 1832, he wrote from two to eleven and a half times as many pages of geological notes per year as biological notes (Porter 1985, p. 984).

Darwin and Covington collected 327 dry specimens (mainly skins) on the outward journey to South America and along its east coast to the southern tip between 1832 and May 1834 (Cape Verde Islands: 5 bird specimens; St. Paul Rocks: 2; Brazil: 8; Uruguay: 115⁶; Argentina: 115; Tierra del Fuego: 49; and Falkland Islands: 33), but only 167 specimens during the second half of the journey from June 1834 to 1836 (Chile: 92 bird specimens; Peru: 4; Galapagos Islands: 65; New Zealand: 1; Cocos Island: 1; Ascension Island: 1; and Cape Verde Island: 3), not collecting any bird specimens from such locations as Tahiti, where the "Beagle" stopped for 11 days, Aus-tralia, where it anchored 28 days in Sydney and 11 days at St. George's Sound, Tasmania, where the "Beagle" crew stayed 15 days, Mauritius (10 days), and St. Helena (7 days). Apparently, Darwin hardly had the chance to get on shore in some places (Darwin cited in Barlow 1967, p. 114): "In our passage across the Pacifick [sic], we only touched at Tahiti & New Zealand: at neither of these places, or at sea had I much opportunity of working." Nevertheless, comparisons of Darwin's "Beagle" notes on entomology (and to some extent on mammalogy) show that, as regards zoology, Darwin "was possibly even fonder of birds than of the beetles generally regarded as his favourites" (Keynes 1997, p. 462).

Darwin's nomenclature

Darwin designated new birds he found in three different ways. He used their correct English generic/family

⁶Darwin noted about Maldonado: "My collection of the birds [...] of this place is becoming very perfect" (Darwin in Barlow 1933, p. 153).

names when he knew them or could learn them from others, e.g. grebe, petrel, cormorant, hawk, caracara, oven bird, wren, sparrow. In some cases, however, Darwin also cited the local Spanish names, such as carpintero for woodpeckers of the genus Dendrocopos, or avestruz(s) petise for the lesser rhea (Pterocnemia pennata). Darwin sometimes mixed up the names of closely related birds or misspelled the name slightly. He made just one real error, labelling the rufous hornero Furnarius rufus with the name for its nest, which is 'casita', meaning little house.

Most errors occurred when Darwin applied scientific (genus) names. Keynes (2002, p. 147) summarizes the problems involved in this approach as follows: "In some cases this was fine—an owl is an owl both in England and in Patagonia—but for many of the passerines [...] there was no directly equivalent South American family, although there were many species that, as he [Darwin] was careful to point out, occupied an ecological niche very similar to those of their European cousins." One may add that some of the species which Darwin encountered were not known to science at the time.

Darwin hardly made any use of the literature on board the "Beagle" in identifying or learning more about birds. Had he done so, he would have found Cuvier (1817, 1829), Griffith et al. (1828), Forster (1778), King (1827, MS to King et al. 1839) and Lesson (1828) most useful. Lesson (1828) is cited just six times in Darwin's ornithological notes, in connection with the correct name for Thinocorus rumicivorus Eschscholtz, 1829 (Barlow 1963, p. 211), when Darwin correctly criticized Lesson's views on the feeding behaviour of the black skimmer Rynchops nigra (Barlow 1963, p. 223), when referring to the nest of Furnarius rufus (Keynes 2000, p. 158), when writing about the different species of penguins in the region (Barlow 1963, p. 246; Keynes 2000, p. 212) and twice in connection with the tameness of birds, especially of the dark-bellied cinclodes Cincl-odes patagonicus (Barlow 1963, pp. 232, 266). However, in Chile Darwin referred frequently to Molina (1809), which increased his ornithological understanding considerably.

Darwin used the following names in his Zoology Notes and Specimen Lists from H.M.S. Beagle (ed. Barlow 1963; Keynes 2000). The names and sequence follow Peters et al. (1934–1987). The term 'in errore' has been added wherever an obvious misidentification is present. A question-mark indicates that a cross-reference between Darwin's note and a current taxon is uncertain.

– RHEIDAE: ostrich, Struthio rhea (for Rhea); ostrich

petise, petise, avestruz petise [sic: Esp. Avestruz petizo

= dwarf ostrich] (for Pterocnemia). – TINAMIDAE: Perdrix [sic: Perdix; in errore ], partridge

[in errore ] (for Rhynchotus, Nothoprocta, Nothura). – PROCELLARIIDAE: Procellaria gigantea, nelly, quebranta

huesos [sic: Esp. quebrantahuesos] (for Macronectes);

[page] 304

Procellaria (for Fulmarus, Procellaria); Puffinus [in errore] (for Procellaria cinerea); petrel (for Puffinus).

– HYDROBATIDAE: Thalassidromus (for Oceanites).

– PELECANOIDIDAE: petrel (for Pelecanoides).

– SPHENISCIDAE: penguin (for Spheniscus).

– PODICIPEDIDAE: fresh water grebe, grebe (for Rollan-dia); fresh water grebe, Podiceps (for Podiceps).

– FREGATIDAE: frigate bird (for Fregata) (not collected).

– PHALACROCORACIDAE: cormorant (for Phalacrocorax).

– SULIDAE: booby (for Sula).

– ARDEIDAE: heron, Ardea (for Ardea); bittern (for Nyctanassa, Nycticorax).

– THRESKIORNITHIDAE: ibis (for Plegadis, Theristicus).

– PHOENICOPTERIDAE: flamingo (for Phoenicopterus).

– CATHARTIDAE: Vultur aura (for Cathartes aura); condor (for Vultur) (not collected).

– ACCIPITRIDAE: hawk, Falco [in errore] (for Circus); hawk, caracara [in errore] (for Buteo).

– FALCONIDAE: caracara, hawk (for Phalcoboenus); chimango, vulture [in errore], Caracara chimango, Falco, P. [= Phalcoboenus] Pezoporus⁷ (for Milvago); Falco (for Polyborus); hawk [in errore] (for Falco).

– ANATIDAE: duck (for Anas, Amazonetta, Chloephaga, Tachyeres).

– RALLIDAE: land rail (for Rallus), "bird from summit of barren arid mountain of Ascension" (? for Porphyru-la).

– HAEMATOPODIDAE: Ostralogus [sic: Ostralegus] (for Haematopus).

– ROSTRATULIDAE: [?] Limosa [in errore] (for Nycticry-phes).

– CHARADRIIDAE: Vanellus (for Belonopterus); Charadrius (for Pluvialis, Oreopholus, Charadrius); Arenaria [in errore] (for Charadrius); plover (for Oreopholus); Tringa [in errore] (for Zonibyx).

– SCOLOPACIDAE: Numenius, curlew (for Numenius); Scolopax, godwit, [?] Tringa (for Limosa); Tringa (for Tringa, Heteroscelus, Tryngites); plover [in errore], Charadrius [in errore] (for Arenaria); Scolopax, snipe (for Capella); Tringa, Pelidna minutilla⁸ (for Erolia).

– RECURVIROSTRIDAE: Himantopus, [?] Charadrius [in errore] (for Himantopus).

– THINOCORIDAE: ptarmigan [in errore], partridge [in errore] (for Attagis); Scolopax-Perdrix [sic: Perdix; in errore], Perdrix-Scolopax [sic: Perdix; in errore], [?] vaginalis, Tinochorus [sic] Eschscholtzii⁹ (for Thino-corus).

– LARIDAE: gull, Larus (for Larus); Larus (for Gabianus); Sterna (for Gelochelidon); tern (for Sterna, Anous); noddy (for Anous).

– RHYNCHOPIDAE: Rhyncops [sic; ? error in Barlow 1963, p. 221 or Darwin's notes; taxon not in Keynes 2000, p. 387] (for Rynchops).

– COLUMBIDAE: palomba [sic: palomo, paloma = Chil-ean Esp./Esp. dove] (for Columba, Zenaidura, Col-umbina); dove (for Columba, Zenaidura, Columbina, Nesopelia, Metriopelia); pigeon (for Columba).

– PSITTACIDAE: Psittacus (for Cyanoliseus, Myiopsitta).

– CUCULIDAE: "small flocks, very noisy chattering bird"(for Guira).

– TYTONIDAE: owl (for Tyto).

– STRIGIDAE: owl (for Speotyto, Strix, Asio).

– CAPRIMULGIDAE: Caprimulgus (for Caprimulgus).

– APODIDAE: swallow [in errore] (for Apus).

– TROCHILIDAE: Trochilus (for Chlorostilbon, Patagona, Sephanoides); humming bird (for Patagona).

– ALCEDINIDAE: Alcedo (for Ceryle, Chloroceryle, Hal-cyon); kingfisher, Alcido [sic; ? error in Barlow 1963, p. 249 or Darwin's notes] (for Ceryle).

– PICIDAE: woodpecker, pituı [old Chilean Esp. for pit-ıo,], Picus (for Colaptes); woodpecker, carpintero [Chilean Esp. = carpenter] (for Dendrocopos).

– FURNARIIDAE: Furnarius (for Geositta, Upucerthia, Eremobius, Cinclodes, Furnarius¹⁰); long-billed casaia (for Upucerthia); oven bird, casita [sic: Esp. little house = nest; casaro = house-builder in Gould and Darwin 1839c], house maker (for Furnarius); Certhia [in errore] (for Limnornis, Aphrastura, [?] Phleocryptes, Limnoctites, Spartonoica, Synallaxis, Asthenes, Pha-cellodomus); creeper [in errore] (for Aphrastura, Asth-enes, Pygarrhichas); Furnarius-Certhia (for Anumbius); long-tailed creeper (for Leptasthenura); red-throated creeper (for Asthenes anthoides), Turdus [in errore] (for Cinclodes).

– FORMICARIIDAE: Lanius [in errore] (for Thamnophilus).

– RHINOCRYPTIDAE: Myothera,¹¹ turco [Chilean Esp. = turca] (for Pteroptochos megapodius); guid-guid [sic: Esp. huet-huet],¹² barking bird¹³ (for Pteroptochos tarnii); Myothera, tapacola [sic: tapaculo] (for Pter-optochos albicollis); Myothera [in errore], cheucau [sic: Esp. chucao] (for Scelorchilus); [unidentified] bird (for Rhinocrypta); [unidentified] bird, wren [in errore] (for Scytalopus); Myothera [in errore], cheuqui [sic: Esp. churrın] (for Eugralla).

– TYRANNIDAE: [unidentified] bird (for Suiriri, Xolmis, Pyrocephalus); Parus [in errore] (for Serophaga, Anairetes, Tachuris); long-tailed tit [in errore] (for Anairetes); Muscicapa [in errore] (for Polystictus, ? Myiophobus, Ochthocea, Hymenops, Xolmis, Neo-xolmis, Muscisaxicola); Motacilla [in errore] (for Lessonia); scarlet-breasted Tyrannus (for Pyrocephalus

⁷See Barlow 1963, p. 278: added after the journey.

⁸Added after the voyage.

⁹See Barlow 1963, p. 211: added after 7th September 1835, during

or after the voyage.

¹⁰Darwin correctly identified Furnarius rufus to species level. ¹¹Myothera is normally synonymised with Pteroptochos albicollis (in Wytsman 1905–1914, fide J. Torres-Mura, personal communication, April 2004), but Sulloway (1982c) synonymized Myothera with all species of Pteroptochos.

¹²Today the Chilean people call Pteroptochos tarnii 'huez-huez.' ¹³The call of this species could be seen as similar to the 'barking' of a very small dog in a high voice; another Chilean bird, the black-necked stilt Himantopus mexicanus has also a 'barking' call, as already noted by Darwin as well (Barlow 1963, p. 217).

[page] 305

rubinus nanus); Tyrannus (for Pyrocephalus, Ochth-ocea); scissor tail (for Tyrannus savana); Lanius [in errore] (for Xolmis, Agriornis); Alectrurus (for Alec-trurus); yellow-breasted Tyrannus (for Myiarchus magnirostris); Sylvia (for ? Muscisaxicola, ? Satrapa); "habits resembling our field-fares" (for Neoxolmis); "like the Butcher bird [...] resembles a hawk [...] & [...] like a kingfisher", bien te veo [Esp.: I see you well] (for Pitangus); "little bird with crest" (for Elaenia); [Dar-win's name not traced for Platyrinchus].

– PHYTOTOMIDAE: finch, Phytotoma rara (for Phytotoma rara).

– ALAUDIDAE: [unidentified] bird (for Eremopterix, Ammomanes).

– HIRUNDINIDAE: swallow (for Tachycineta, Progne, Notiochelidon); Hirundo (for Progne).

– MOTACILLIDAE: Anthus, Alauda [in errore], lark [in errore] (for Anthus).

– TROGLODYTIDAE: Sylvia [in errore], "bird lives near the beach" (for Cistothorus); wren (for Troglodytes).

– MIMIDAE: Lanius, thenca [Esp.], callandra [sic: Esp. calandria] (for Mimus); thenca [Esp.] (for Nesomimus).

– TURDIDAE: thrush, Turdus (for Turdus).

– EMBERIZIDAE: Fringilla [in errore] (for Ammodramus, Zonotrichia, Phrygilus, Diuca, Poospiza, Embernagra, Geospiza, Camarhynchus, Thraupis); sparrow [sensu American sparrows] (for Zonotrichia); blue sparrow (for Phrygilus alaudinus); Emberiza (for Melanodera, Sicalis); Icterus [in errore] (for Geospiza); grosbeak [in errore] (for ? Camarhynchus); Sylvia [in errore], wren [in errore] (for Certhidea); [Darwin's names not traced for Aimophila, Donacospiza, Sporophila, Gubernatrix, Volatinia, Saltator and Pipraeidea].

– PARULIDAE: Sylvia [in errore] (for Dendroica, Geothly-pis).

– VIREONIDAE: Lanius (for Cyclarhis).

– ICTERIDAE: Xanthornus (for Xanthopsar); Icterus (for Agelaius, Pezites, Pseudoleistes, Amblyramphus, Curaeus, Molothrus); thili, chili [from Molina 1809, Chilean Esp. trile] (for Agelaius);¹⁴ Sturnus ruber [in errore] (for Pezites militaris [nec Sturnella rubra = Amblyramphus holsericeus]); Anthus [in errore] (for Dolichonyx); [unidentified] egg (for egg of Molothrus).

– FRINGILLIDAE: Fringilla, siu [from Molina 1809, Esp. jilguero] (for Carduelis).

– PLOCEIDAE: sparrow (for Passer).

a rapid ride across the pampas. (Hudson 1870a, about Darwin).

Waders posed the biggest challenge to Darwin's identification skills. He misidentified, for example, the common turnstone Arenaria interpres as a plover of the genus Charadrius, and vice-versa. Darwin lacked any real interest in waders, and barely mentioned their behaviour, habitat or geographical range. Accordingly, the chapters on species of Charadriiformes in the Zool-ogy of the Voyage of H.M.S. Beagle (Gould et al. 1841) are quite poor and contain very little data beyond the collecting localities.

Furthermore, Darwin had immense problems in applying any sensible name to species of bird confined to America, such as Tinamidae,¹⁵ Furnariidae, Tyrannidae and Rhinocryptidae.

A notebook entry from the Patagonian coast near Santa Cruz in April 1834 reveals the problems Darwin faced when writing about undescribed bird species (Barlow 1963, p. 247): "[No.] 2011 Callandra.- see supra. No ? (1216).- S. Cruz / [No.] 2012 Lanius; rare (unless ?) [del.] both on coast & in the interior plains; chases insects very quickly half flying, half running.- do [S. Cruz] / [No.] 2013 Lanius.- rare.- female.- Hab. do. [S. Cruz]—N. B. [added about a year later in summer 1835] One of these birds is brownish with a white tail; I saw it in the lofty & arid valleys on the Eastern slope of the Cordillera at Chili.- The other of these, or one shot at P. Desire or St Julians, is brown all over & with a broarded [sic: broader] bill like true Lanius, this bird is found also in Chili as far North as Copiapo. I am as-sured that it commonly kills young birds." The first species Darwin was referring to is the Patagonian mockingbird Mimus patagonicus, described by Laf-resnaye and d'Orbigny in 1837. The other birds are shrike-tyrants, Agriornis montana leucura (Gould, 1839) and Agriornis microptera microptera (Gould, 1839) both described by Gould on the basis of Darwin's material, and the species that kills young birds is Agriornis livida livida (Kittlitz, 1835) (cf. Darwin in Gould and Darwin 1839c: p. 56: Agriornis gutturalis). Darwin used the substitute name Lanius (genus of old world shrikes) for the shrike-tyrants.

Labelling of specimens

Identification problems

However close an observer a naturalist may be, it is not possible for him to know much of a species from seeing perhaps one or two individuals in the course of

¹⁴Darwin wrote in Gould et al. 1841, p. 106 that the name of the country would derive from the song of Agelaius thilius, which is indeed one of the theories as to where Chile got its name from (J. Torres-Mura, personal communication, April 2004).

Trust nothing to the memory; for the memory becomes a fickle guardian when one interesting object is succeeded by another still more interesting. (Darwin 1839, p. 598).

Darwin recorded the important information regard-ing each specimen, such as locality, date and habitat, in his notebook only, not on the specimen label. Even after

¹⁵Subsequent authors (Steinmuller and Steinmuller 1987, p. 153) still referred to the tinamous in Darwin's words, as grey partridges.

[page] 306

Fig. 1 Original field label by Darwin on a specimen of Gallinago paraguaiae (BMNH 1860.1.16.74). The species name is in Gould's hand, the number "1203" is written by Darwin. Photo: Harry Taylor, Natural History Museum, London/Tring

the voyage he still considered this the most reliable method of recording data (Darwin 1839, pp. 598–600). His specimen labels were made out of scrap paper, tied with a cord to the foot or neck of the bird (birds in alcohol got a metal tag; Figs. 1, 2). These bird labels contained nothing except for a number ranging from 185 to 3907 (all dried zoological items were numbered in the same sequence), cross-referring the individual specimen to the corresponding notebook entry. Darwin kept separate notebooks for geological (cf. Herbert 1980) and biological specimens (plants and animals: Zoology Notes, Keynes 1997), the latter numbered in two different sequences for dry and wet collections (Porter 1985, p. 979; cf. Darwin 1839). During the last months of the voyage, separate lists of the mammals, birds,¹⁶ insects, shells, plants, reptiles and amphibians, crustaceans, and fishes, were copied out by Darwin and Covington for the benefit of the specialists to whom they were to be given for classification. During this procedure, Darwin marked each original 'bird entry' with a capital 'B' (Chancellor et al. 1988). Some original bird entries, though, were missed out, so that one has to consult the original Zoology Notes (ed. Keynes 2000) rather than the bird notes (ed. Barlow 1963) when completeness is important. The numbers on the skin labels were mainly written in pencil, and to a lesser extent in Indian ink. Only one label is known to have survived on which the locality is also cited. It belongs to a Spanish sparrow Passer hispaniolensis (BMNH¹⁷ 1881.5.1.2117) collected

on the Cape Verde Islands, collection number 189, and reads "loc S Jago" [in pencil; collected at Praia (14°55'N, 23°31W) on the island of Sao Tiago]. Later, John Gould (1804-1881), who worked on these specimens, sometimes added a species name to the original label. All Darwin's birds would have carried these labels originally, but later owners of the bird collection did not appreciate the importance of these reference numbers, replacing Darwin's labels with proper museum labels after the sale of the Zoological Society Museum in 1855. Thus, most of Darwin's data was used only once, in the writing of the section of the Zoology of the Voyage of H.M.S. Beagle which deals with birds (Gould and Darwin 1838, 1839a, 1839b, 1839c; Gould et al. 1841). Most subsequent museum labels do not quote any actual collecting locality and rarely make reference to Darwin's notebook numbers. Unfortunately, just 16 original labels have survived (Sulloway 1982a knew only one; see Appendix). The numbers of 499 bird items were entered into the notebooks of Darwin published by Barlow (1963) and Keynes (2000), but, as comparisons with the text of Gould and Darwin (1838, 1839a, 1839b, 1839c) and Gould et al. (1841) have now revealed, a total of 512, perhaps even 515, items had probably been collected. Two notebook numbers are shared by two and, respectively, three different egg clutches. The birds not traced in the notebooks are two specimens in alcohol, five skins from Rio de Janeiro (April to July 1832) and

¹⁶The ornithological notes had been compiled between 12 April–19

July 1836 (Sulloway 1982c).

¹⁷For any museum's abbreviation see Appendix.

¹⁸The Zoological acquisition date

Society labels recorded the donor's name, the and Darwin's specimen numbers: front "C.

Darwin Esq. Jan 4 1837", back (number); but most of these labels

were also replaced.

[page] 307

Fig. 2 A set of all but one specimen of Darwin's collection with original field labels attached. Photo: Harry Taylor, Natural History Museum, London/Tring

one skin each from Tierra del Fuego (southern summer 1832/1833 or 1834), Buenos Aires (October 1833) and East Falkland Island (March 1833 or 1834).

Darwin used a method for recording data known to geologists and palaeontologists. It is difficult to tie a label to a stone, and writing a number on the specimen cross-referring the item to a notebook entry is the only practicable solution. However, most bird collectors would put any important information directly on the specimen labels, entering additional data into a notebook only if space were short. Darwin's contemporar-ies Johann Natterer (1787–1843), Alfred Russel Wallace (1823–1913) and Eduard Ruppell (1794–1884), to name but a few, used minute paper labels, but still wrote down collecting locality, date, sex and a reference number.

Observations and investigations

I do not doubt that every traveller must remember the glowing sense of happiness he experienced, from the simple consciousness of breathing in a foreign clime, where the civilized man has seldom or never trod. (Darwin 1839, p. 606).

Despite the foregoing criticism of the view that Darwin was a good field ornithologist, hedid make some excellent

ornithological observations and carried out innovative experiments, noting down every minute detail of his research. His ornithological notes run for 86 numbered and 10 additional pages. Darwin's real strength was that he never took anything for granted and was aware that either he or other scientists might have made errors in certain observations. Darwin was also good at asking the right questions, many of which he could then answer himself during the voyage through his own field studies. Take for example his investigations into the olfactory abilities of the Andean condor Vultur gryphus, which had been triggered by a report by Audubon that vultures can smell (Darwin in Gould and Darwin 1838, pp. 3–6; cf. Barlow 1963, pp. 240–245):

I tried [...] the following experiment. The condors were tied, each by a rope, in a long row at the bottom of a wall. Having folded a piece of meat in white paper, I walked backwards and forwards, carrying it in my hand at the distance of about three yards from them; but no notice whatever was taken of it. I then threw it on the ground within one yard of an old cock bird; he looked at it for a moment with attention, but then regarded it no more. With a stick I pushed it closer and closer, until at last he touched it with his beak: the paper was then instantly torn off with fury, and at the same moment every bird in the long row began struggling and flapping its wings. Under the same circumstances, it would have been quite impossible to have deceived a dog.

[page] 308

What Darwin observed ties in what we know about these birds today: while the Cathartes species studied by Audubon have a superb olfactory sense, condors find their food optically (Houston 1994).

Darwin also gave a vivid account of the lesser rhea Pterocnemia pennata (Keynes 2000, pp. 188–189):

There is some notice about a second species of Rhea, which is very rarely found N of the R. Negro [Pata-gonia]. Mr Martens [Conrad Martens (1801–1878), the official artist on board] shot one at Port Desire, which I looking slightly at it pronounced to be a young one of the common sort, that is it appeared to be 2/3 in size of the common one. I also [saw] some live ones of the same size, but entirely forgot the Petises. I have since reclaimed the Head, Legs & several feathers [...]. The scales on legs are of a different shape, & is feathered below the knees, this accounts for their being said to be shorter in the legs & perhaps for being feathered to the claws. (it is a bird which the R. Negro Gauchos have only seen once or twice in their lives). [...] Whatever Naturalists may say, I shall be con-vinced from such testimony as Indians & Gauchos that there are two species of Rhea in S. America.

Darwin was correct. Two species of rheas inhabit South America. These parts of several different individuals of rhea were later mounted to form a nearly perfect composite specimen, now considered lost, for the museum of the Zoological Society of London (Rice 1999). John Gould, the curator of the Society's museum, described Darwin's, or, as it is called today, the lesser rhea as Rhea darwinii in 1837 (Gould 1837d). The fa-mous French traveller Alcide Dessalines d'Orbingy (1802–1857), however, got the credit for the discovery and naming of the bird, which he described as Rhea pennata in 1834, the same year in which Darwin wrote the above note, away from civilisation and out of contact with the scientific community (cf. Herbert 1980, pp. 107–110).

Darwin was excellent in his observations when it came to game birds. His account of tinamous, for example, reads like a hunting manual:

Does not live in covies [pl. of covey = flock of par-tridges], but generally in pairs, runs more & does not lie so close as the English Partridge; not a high shrill[,] chirp or whistle—When riding through the country great numbers, (generally in pairs) may be seen, as when at a short distance they take no pains to conceal themselves. They are silly birds; a man on horse-back by riding in a circle or rather in a spire, round & round, each time closer & closer, so bothers the bird, that it may be knocked on the head, or noosed by running knot, at the end of a long stick, with the greatest ease. Flesh most delicately white, when cooked more so than of Pheasant, but rather dry & flavourless (Barlow 1963, p. 218).

Darwin gave some hints that where he indicated the sex of a bird, these had in fact been sexed internally by

dissection. He did not believe that birds could always be sexed on size or plumage characters, which was common practice among the naturalists of the day. In his discussion of the different colour morphs and sizes of striated caracara Phalcoboenus australis from the Falk-lands, Darwin checked for the first time during the voyage the reproductive organs (Darwin in Gould and Darwin 1838, p. 16). It is unclear whether somebody influenced Darwin, and subsequently Covington, to adopt internal sexing as a normal procedure when preparing bird skins. Neither do we know whether Darwin was taught the differences in the sexual organs of birds or if he found out how testes differ in appearance from ovaries himself (cf. nothing is mentioned in Armstrong 1992). From March 1834 on, though, Darwin and Covington consistently dissected birds to sex them. In a notebook entry for birds collected around Valparaiso (Chile) Darwin stated (in Barlow 1963, pp. 253-254): "The following birds were shot at Valparaiso, during months of August & September [1834].- The sexes were distinguished by S. Covington by opening their bodies, & judging chiefly from the granulated state of the Ovarium: it being the Spring [of the Southern hemisphere], probably this means is correct." Darwin, an amateur ornithologist, would not have had the necessary knowledge of sex-related plumages, and therefore had to dissect.

Careful investigations were made into the geographical range of each species and its general habitat. Furthermore, Darwin was interested in what the birds ate, frequently dissecting the stomachs of birds and reporting on the contents. Most of his thorough bird accounts close with a passage on the bird's calls and songs. Where Darwin was able to gather information on breeding, this information has also been added. The following is a good example of a detailed notebook entry:

[No.] 1222 Furnarius [i.e. common miner Geositta cunicularia cunicularia (Vieillot, 1816)].- This bird has a considerable distribution. On the east coast, it occurs from 30° (& perhaps in this & many other cases those bird[s] which frequent plains, reach much further to the North, to the extreme limit of the great central plains) to 40°. I never saw a specimen further South than this. On the west coast, it occurs from Concepcion (where open country commences) 37°, to as far (at least) as Lima in 12°.- It constantly haunts the driest plain parts, away from bushes & trees. Sand dunes, near the sea coast afford a very favourite resort. At Maldonado [Uruguay] & at Bahia Blanca [Patagonia] it is very abundant, it is very tame, most quiet solitary little bird; being disturbed only flies to a very short distance. Is active early in the mornings & late in the evenings (like Robin): is fond of dusting itself in a road: walks, but cannot run very quickly; in stomach small Coleoptera [beetles], chiefly Carabi-dous insects. At certain times it frequently utters a peculiar shrill, but gentle, quickly reiterated cry (so quickly reiterated as to make one running sound). In

[page] 309

this respect resembles the Oven bird [Furnarius rufus], but as widely differs in its quietness, from that active bird.- It builds its nest at the bottom of a narrow cylindrical hole, which is said to extend horizontally to nearly six feet long. Several country men, told me, that when boys they had attempted to dig out the nests, but from their depth had nearly always failed.-The bird chooses any low little bank of firm sandy soil, by the side of a road or stream. (Barlow 1963, pp. 217–218).

Without the personal anecdotes, this species description could not be beaten by any in a modern handbook (cf. e.g. Ridgely and Tudor 1994, pp. 26–27). Most unusual for the time, but extremely helpful, were Darwin's reports on the pedal locomotion of birds, which is sometimes characteristic for the whole family or genus. He very often noted down if a bird hopped, walked or ran on the ground. Even the geographical range of the species is given more or less correctly,¹⁹ which is astonishing when one takes in account that this would entail Darwin having to check for the bird in all the places he visited. Darwin was intrigued by the isolation factor constituted by the Andes and, probably towards the end of the voyage, compiled a list of birds occurring on both sides of the mountain range (Keynes 1997, p. 466).

Some notes, however, were extremely poor. Sulloway (1982a, 1982b) has already explored this fact with regard to the Galapagos finches. No attention was paid to which island which specimen came from. During the Galapagos visit itself, Darwin showed no great interest in this group of birds, nor in any other bird from the Galapagos Islands, perhaps with the exception of the mockingbirds (Keynes 2003). The same is apparently true for his visit to Isla de Chiloe (Willson and Armesto 1996).

A letter from Darwin to George Robert Gray written in the summer of 1839 (Burkhardt and Smith 1986, vol. 2, p. 196) witnesses the problems faced by the scientists who worked on Darwin's collection when confronted with Darwin's scanty notes: "With respect to the Falkland Emberiza, I can give little assistance 1919, 1920, 1922 female 'Shot in same large scattered flock on the hills'[.] But 1920 & 1923 & 1879 'I think more commonly occur on the plains' I put mark of ? to 1923 whether different species or not.- In another part of my catalogue I say I saw 1046 & 1047 together[.] My specimens were shot in March, corresponding to September, & this is all the very little inform[ation] I can tell you[.]" Gray (in Gould and Darwin 1839c, Gould et. al. 1841) never-theless managed to correctly identify one group of birds as the black-throated finch Melanodera melanodera (Fig. 3) and described the other taxon as a new species, the yellow-bridled finch Melanodera xanthogramma.

Darwin's contribution to ornithology

I shall relate [...] the few facts, which I have been enabled to collect together; and these, if not new, may at least tend to confirm former accounts. (Darwin's preface of Gould and Darwin 1838).

There is no question that of all Darwin's works, his evolution theory has had the biggest impact on orni-thology (cf. Stresemann 1951). However, Darwin also kept important notes and collected a small number of very significant birds, among them birds new to science at the time or from nowadays extinct populations. Just how important were Darwin's bird collection and notes for the ornithology of the countries he visited?

Essentially, Darwin's bird collection only became ornithologically significant through the involvement of other people. Luckily, acknowledging that he was no expert in ornithology, Darwin donated his collection to the most senior experts in the field, to the anatomically interested Thomas Campbell Eyton (1809–1880), to George Robert Gray (1808–1872) and to John Gould (cf. Sauer 1998). The latter busily went about describing Darwin's new bird species in lectures given to the Zoo-logical Society of London between January 1837 and January 1838 (Gould 1837a, 1837b, 1837c, 1837d, 1837e, 1837f, 1838), while Darwin added some comments dur-ing two of these meetings (Darwin 1837a, 1837b). Eyton compiled the anatomical appendix for the Zoology of the Voyage of H.M.S. Beagle (Gould et al. 1841), but also used some data gathered by Darwin in his Synopsis on the Anatidae (Eyton 1869, pp. 29, 31–32).

John Gould later summarized the ornithological knowledge which had been accumulated by Darwin during his journey in part 3 (Birds) of the Zoology of the Voyage of H.M.S. Beagle (Gould and Darwin 1838, 1839a, 1839b, 1839c; Gould et al. 1841), a book mainly financed by the government (Lords Commissioners of Her Majesty's Treasury; cf. Darwin in Owen and Darwin 1840, pp. II-III). Gould left a draft of the manuscript and the lithographs of Elizabeth Gould (1804–1841) with Darwin when he and his wife left for their own expedition to Australia on 16 May 1838. Some doubts remained with regard to certain birds and their taxonomy, so Darwin consulted George Robert Gray for advice. Gray was asked to work over Gould's manuscript, and he apparently double checked each name in issues 2–5, which had not yet been published (cf. e.g. Burkhardt and Smith 1986, vol. 2, p. 280).²⁰ It took Gray so long to finish issue 5 that Darwin was forced to ask him in 1840 if he would "oblige [him] by kindly finishing the remaining

¹⁹The species does, however, occur along the east coast to the top of Tierra del Fuego.

²⁰The first issue (pp. 1–16, pls. 1–10) of the bird section of Zoology of the Voyage of H.M.S. "Beagle" was apparently published in July 1838, the second (pp. 17–32, pls. 11–20) in January 1839, the third (pp. 33–56, pls. 21–30) in July 1839, the fourth (pp. 57–96, pls. 31– 40) in November 1839, the fifth (pp. 97–164, pls. 41–50) in March 1841 (Burkhardt and Smith 1986, vol. 2, pp. 432–437; Zimmer 1926, p. 159).

[page] 310

Fig. 3 Darwin's specimens of the black-throated finch Melanodera melanodera (BMNH 1855.12.19.50, BMNH 1885.12.14.806) in comparison to Gould and Darwin (1839c): plate 32. Photo: Harry Taylor, Natural History Museum, London/Tring

MS" (Darwin cited in Porter 1985, p. 1003). Darwin was not very pleased to see that Gray had added his name to any new combination of names, disregarding 'Strick-land's laws'²¹ (letter to Leonard Jenyns, cited in Burk-hardt and Smith 1986, vol. 2, p. 317), but Gray's name nevertheless appears in the printed version, which was published from July 1838 to March 1841, heavily edited by Darwin himself. For this task Darwin had worked on both his bird collection and that of the BMNH from mid 1837 onwards (Brandon-Jones 1996, p. 504). His notes are in fact equally important as Gould's and Gray's descriptions as they add much on habitat, behaviour and even ecology for some species not found elsewhere at the time. Furthermore, the expert work on avian taxonomy by Gould and Gray provided Darwin with some raw material for his thoughts on evolution (see below; cf. Sulloway 1982 b, 1982c).

Darwin twice resumed his ornithological notes in later life (Darwin 1870, 1881). In 1870, Darwin wrote a paper

²¹Strickland et al.: Report of a committee appointed "to consider of the rules by which the nomenclature of zoology may be estab-lished on a uniform and permanent basis." Report of the 12th meeting of the British Association for the Advancement of Science held at Manchester 1842:105–121 (cited in Burkhardt and Smith 1986, vol. 2, p. 500). Darwin was much in favour of referring the species name to the first author, but not to the author of any new combination.

on the habit of the campo flicker Colaptes campestris, of which he once collected two specimens in Uruguay and Argentina (cf. Barlow 1963, p. 219, Nos. 1238; and p. 225, No. 1428). When Darwin studied the campo flicker in the field near Maldonado, Uruguay, he wrote in his notebook that it "alights horizontally, like any common bird, on the branch of a tree: but occasionally I have seen it clinging to a post vertically [...] frequent the open plains." (Barlow 1963, p. 219). Darwin explored the example also in On the Origin of Species:

On the plains of La Plata, where hardly a tree grows, there is a woodpecker (Colaptes campestris) which has two toes before and two behind, a long pointed ton-gue, pointed tail-feathers, sufficiently stiff to support the bird in a vertical position on a post [...], and a straight long beak. [...] Hence this Colaptes in all the essential parts of its structure is a woodpecker [...], yet, as I can assert, not only from my own observations but from those of the accurate Azara [1802–1805], in certain large districts it does not climb trees, and it makes its nests in holes in banks! (Darwin 1872, p. 183).

Darwin's paper from 1870 was triggered by the accusation of William Henry Hudson (1841–1922) that these woodpeckers, as published by Darwin in the Origin of Species, do not breed in mud banks, but in tree holes like other woodpeckers (Hudson 1870a, 1870b). It is now common knowledge that the campo flicker does both (Winkler et al. 1995, p. 326).

Eleven years later Darwin published on the parasitic behaviour of the cowbirds Molothrus, four specimens of two species of which were to be found in his collection.

John Gould described a total of 37 new bird species on the basis of Darwin's collection. G.R. Gray added one further species.²² Darwin collected another 12 birds previously described by Molina (1782), 10 which had been described from d'Orbigny's collection in 1837 and 1838, 7 birds already named by Lesson in 1826–1839, 8 described by King in 1828–1831 and 13 described by Kittlitz mainly in his 1830 publication (see Appendix for type specimens in Darwin's collection).

Darwin's contribution to the avifauna of Chile has already been discussed (Jaksic and Lazo 1994; Willson and Armesto 1996). He followed Giovanni Ignazio Molina (1740–1829), Rene Primevere Lesson (1794– 1849), Prosper Garnot (1794–1838), Philip Parker King (1791–1856), Baron Friedrich Heinrich von Kittlitz (1799–1874), Jeremiah Reynolds (1799–1859) and Al-cide Charles Victor Marie d'Orbigny (1802–1857) as a collector of birds in Chile (cf. Steinheimer 2002, p. 35). Hardly any new (sub-)species were left to discover,²³ though Darwin contributed many useful ornithological observations and range extensions.²⁴ The same is true for the greater part of the east coast of South America. In the plains of Patagonia, however, Darwin encoun-tered 11 undescribed taxa,²⁵ despite the fact that d'Orbigny and Captain Philip Parker King had also collected there.²⁶ Other new species came from Maldonado in Uruguay²⁷ and from the Cape Verde

Islands.²⁸

²²G.R. Gray also described Tyto alba punctatissima (G.R. Gray in Gould and Darwin 1839b, pp 34–35), but the description had been based on a specimen of Captain FitzRoy's collection. ²³Three new Chilean taxa are based on Darwin's collection: Cap-rimulgus longirostris bifasciatus Gould 1837c: Valparaiso; Melano-dera xanthogramma xanthogramma (Gray in Gould and Darwin 1839c): Tierra del Fuego and on Falkland Islands; Ochthocea parvirostris (Gould in Gould and Darwin 1839b): Valparaiso and Santa Cruz, Argentina.

²⁴Following Jaksic and Lazo (1994), Darwin made notes of about 40 Chilean species out of 440, and added 11 new country records. ²⁵Aimophila strigiceps strigiceps (Gould in Gould and Darwin 1839c): Santa Fe; Agriornis microptera microptera Gould in Gould and Darwin 1839c: Patagonia; Agriornis montana leucura Gould in Gould and Darwin 1839c: Port Desire; Asthenes pyrrholeuca flavogularis (Gould in Gould and Darwin 1839c): Patagonia; Buteo ventralis Gould 1837b: Santa Cruz; Caprimulgus parvulus parvulus Gould 1837c: Santa Fe; Coturnicops notata notata (Gould in Gould et al. 1841): Rio de la Plata; Eremobius phoenicurus Gould in Gould and Darwin 1839c: Patagonia; Myiophobus fasciatus auriceps (Gould in Gould and Darwin 1839b): Buenos Ayres; Ochthocea parvirostris (Gould in Gould and Darwin 1839b): Santa Cruz and Valparaiso, Chile; Phalcoboenus megalopterus albogularis Gould 1837b: Santa Cruz.

²⁶King's birds are mainly housed in the BMNH collection. See d'Orbigny and Gervais (1835–1847) for details of d'Orbigny's collection.

²⁷Ammodramus humeralis xanthornus Gould, 1839; Limnoctites rectirostris (Gould, 1839); Limnornis curvirostris Gould, 1839. ²⁸Ammomanes cincturus cincturus (Gould, 1839); Eremopterix nig-riceps nigriceps (Gould, 1839); Passer iagoensis iagoensis (Gould, 1837).

311

Ornithologically the most fruitful was Darwin's visit to the Galapagos Islands, where no proper collection-based survey of birds had ever taken place before the "Beagle" crew came ashore. Twenty-two new (sub-)species were among the birds which Darwin brought back to Britain. They were all described immediately by John Gould, who also, as discussed in Sulloway (1982a, 1982b), was the first to recognize the close relationship among the Galapagos finches. Gould received Darwin's specimens on 4 January 1837 (see below), and had already presented the Galapagos fin-ches as new to science 6 days later at a meeting of the Zoological Society (10 January 1837). At that time, Darwin had not yet given Gould the necessary notebook data so he was forced to rely on descriptions only, even in sexing the birds (cf. Gould 1837a). It was only later that Gould placed the genus Certhidea within the group (probably during March 1837), but this was covered up by the fact that before the proceedings of the meetings of the Zoological Society were published later the same year, Gould had corrected his paper (1837a) of the 10 January meeting accordingly (Sullo-way 1982b, pp. 21–22).²⁹ What Gould (1837a), unlike most 19th century ornithologists, understood after several months of research (cf. Sulloway 1982b, pp. 21– 22) was that within this group of birds, the different beaks, which for continental birds were believed to be one of the stable family characters, were highly modi-fied whereas the plumage, otherwise most variable, remained almost identical.

One can therefore say that it was mainly John Gould who made Darwin's collection and notes into a signifi-cant contribution to ornithology. The Galapagos finches subsequently proved to be more interesting than any other specimens in Darwin's collection. Swarth (1929, 1931), followed inter alia by Lack (1947, cited 1961), Steadman (1982) and Rosemary and Peter Grant (e.g.

²⁹Although contested by Warren and Harrison (1971) and Sullo-way (1982a, 1982b, 1982c), only Darwin's specimens were used in Gould's first descriptions, and therefore only these specimens can be considered types of the Geospizinae (Galapagos finches). If birds collected by such a high-ranking person as a Captain of the Royal Navy had been available, this would have been mentioned in Gould's publication. Darwin travelled as private and self-financed naturalist on the "Beagle" and was free to dispose of his specimens as he liked. FitzRoy, an employee of the Navy, was more restricted. He gave, probably on order by Sir William Burnett (1779–1861), the physician-general of the navy, his bird collection to the BMNH on 21 February 1837, where they did not get the same attention accorded to Darwin's at the Zoological Society. Sulloway (1982a) claims that Gould also used a Geospiza specimen from Fuller's collection, which was given to him by Eyton. However, there is no correspondence to corroborate this, and it is therefore believed that the missing taxon was not in Eyton's collection, but was among those of Darwin's Geospiza specimens not traced. Darwin later communicated the locality data relating to the collections of FitzRoy and other shipmates to John Gould, and it is even assumed that Gould later saw FitzRoy's birds in the BMNH, but as there is no reference to FitzRoy in Gould's first description, any other Galapagos finches than those of Darwin can be at best seen as Paratypes (cf. Gould 1837a).

[page] 312

Grant et al. 1985; Grant 1986), showed their adaptive radiation and on-going evolution, similar to that of the Hawaiian honeycreepers, which make them one of the best examples of Darwin's evolutionary theory (cf. Weiner 1994).

Five bird populations encountered by Darwin are today believed to be extinct. His specimens count among the few remaining records we have of the existence of these populations, which comprise the large-beaked population of the large ground-finch Geospiza magni-rostris magnirostris (cf. Fuller 2001, p. 297), believed by Darwin to occur on Santa Marıa (Charles) and San Cristobal (Chatham) Islands,³⁰ the Santa Marıa (Charles) Island population of the sharp-beaked ground-finch Geospiza nebulosa (cf. Lack 1961, p. 23; Sulloway 1982b, p. 30), the population of the Charles mocking-bird Nesomimus trifasciatus trifasciatus from Santa Marıa (Charles) Island, though the same subspecies still occurs on neighbouring islands (Mayr and Greenway 1960, p. 447), the yellow-bridled finch Melanodera xan-thogramma xanthogramma from the Falkland Islands³¹ and the Andean tapaculo Scytalopus magellanicus ma-gellanicus, also from the Falklands.³² Last but not least, Darwin recorded a "gull" in the "neighbourhood of Porto Praya [i.e. Sao Tiago] from 16th of January to 7th of Feb. [1832]" (Keynes 2000, p. 371; Darwin No. 185). There is hardly any other record of any gull species from the Cape Verde Islands (Hazevoet 1995). Unfortunately, the gull specimen, which Darwin shot on the Cape Verde islands and which he entered under number "185" in his specimen list, has not been traced. A specimen of rail from the Ascension Islands, also of unknown where-abouts, might also have proved interesting. Taylor (1998, p. 478) listed the record under the species of Al-len's gallinule Porphyrio alleni.

Darwin's bird collection

Let [the collector] work hard from morning to night, for every day and every hour is precious, in a foreign

³⁰Sulloway (1982b) raises the possibility that Darwin's birds did not come from these two islands at all, but from San Salvador (James) Island. Still, there is no bird with such an enlarged beak anywhere in the Galapagos Islands today.

³¹On 6–8 July 1998, a resident of the Falkland Islands, Lynda Anderson, probably saw two birds of this species on San Carlos, East Falkland (R. Woods, personal communication, April 2004, cited from a fax to Hay Miller, July 1998). However, there is no evidence that the species is still resident in the Falklands (Woods 1988; R. Woods, personal communication, April 2004). ³²It seems conclusive that Darwin indeed collected a specimen of that species on the Falkland Islands (Keynes 2000, p. 385, Darwin No. 1144, see Appendix). However, the bird was more likely to be a rare vagrant from mainland South America than a breeding bird on the Falklands, which was what Darwin first believed (in Gould and Darwin 1839c, p. 74; cf. Woods 1988, p. 213). Darwin might have confused the nests of Cistothorus platensis (W.R.P. Bourne, personal communication, February 2000; R. Woods, personal communication, April 2004).

clime; and then most assuredly his own satisfaction will one day well repay him. (Darwin 1839, pp. 601– 602).

Darwin's comprehensive geographical notes on certain birds' distributions were only possible because he would, at least during the first 3 years of the voyage, always collect any bird, when time permitted, regard-less of whether he already had a specimen of the same species in his collection or not. This might have been due to the fact that he was not actually able to tell the difference in the field, but it may more charitably be believed that his collection of small series³³ of the same species was actually inspired by his intense nterest in biogeography and geographic variation. Darwin may often have been thwarted, though, by the space problem in the tiny cabins and store rooms of H.M.S. "Beagle",³⁴ which probably did not permit constant access to his collection when he wanted to make com-parisons.

Obviously, contemporary field collectors of the likes of Alcide Dessalines d'Orbingy in 1826–1833 and the early Alfred Russel Wallace in 1848–1852 did more comprehensive collecting in South America than Darwin. Johann Natterer brought back more than 12,000 bird specimens from his exploration of Brazil in 1817– 1835. Even Captain Robert FitzRoy assembled a collection of 188 birds, not bad when one considers that he was in command of the "Beagle" and chiefly occupied with meteorology, hydrography and charting the coast line of the South American continent (Sharpe 1906, p. 323).

Darwin's 468 bird skins, 10 parts of the lesser rhea, the nests and eggs of 16 taxa, and his 14 whole birds and 4 parts of specimens in alcohol from the voyage of H.M.S. "Beagle" seem nothing in comparison (cf. Appendix; Eyton in Gould et al. 1841).^35,36 "But it must be borne in mind how large a proportion of time, during a long voyage, is spent on the water, as com-pared with the days in harbour [...besides] other losses [...] are the want of room, of seclusion, of rest; the jading feeling of constant hurry [...] " (Darwin 1890a, pp. 532, 533).

It is fairer, then, to compare Darwin's collection with those of other travellers on national circumnavigations, such as Baron Friedrich Heinrich von Kittlitz (1799– 1874) during the voyage of the "Senjawin" in 1826–1829 and Andrew Bloxam (1801–1878) on H.M.S. "Blonde" in 1824–1825, whose bird collections are indeed of a

³³It has to be stated, though, that the largest series of any one and

the same species numbers just six individuals (cf. also Sulloway

1982c).

³⁴Darwin had a very small cabin under the forecastle at his disposal

for storing specimens (Porter 1985, p. 986).

³⁵Darwin made some additional field notes on birds not collected

(cf. Darwin 1839; Jaksic and Lazo 1994).

³⁶Corrections of the total number of Darwin's avian specimens

have been made since Steinheimer (2003).

similar size to Darwin's. Nevertheless, most voyages and most field collectors collected many more birds than Darwin.

Nonetheless, as shown above, Darwin's contribution to ornithology was not insignificant. Many of the specimens collected by Darwin are now the types for new names proposed by John Gould. Furthermore, "there has probably been more published on Darwin's orni-thological collections than on all the other groups combined" (Porter 1985, p. 1002). One might therefore wonder why we do not actually know much more about Darwin and his bird collection, other than those from the Galapagos Islands. Nobody has ever comprehen-sively listed, for example, which birds Darwin collected or encountered during the voyage of the "Beagle". There are several explanations for this. One is the subsequent dispersal of Darwin's bird collection. When Darwin was still out in South America, he would send back bird specimens to Prof. John Stevens Henslow at Cambridge, who was paid by Charles' father, Dr. Robert Waring Darwin (1766–1848), to store the boxes until Darwin's return.³⁷ Henslow received four boxes and eight casks sent from Montevideo and the Rio de la Plata in August 1832, November 1832, July 1833 and November 1833, one box from Buenos Aires sent in May 1834 and four boxes from Valparaiso sent in January and June 1835 (Porter 1985). All additional specimens collected later than June 1835, such as birds from Lima and the Galapagos Islands, were stored in the "Beagle" until her return to England.

Once Darwin was back in Britain, he and Covington spent some time in Cambridge sorting out the birds from the rest of Darwin's natural history collection in late December 1836 and early January 1837 (Sobol 1959, p. 126). Darwin had already been concerned before the journey as to whom he would present the zoological specimens he was going to collect during the circumnavigation. He had originally tended towards the British Museum, but could "not feel [...] any great respect even for the present state of that establishment", while the Zoological Society Museum was considered "nearly full" (Porter 1985, pp. 979, 991). Upon his return Darwin learned that his bird collection was neither wel-comed with open arms nor much valued. "I am out of patience with the Zoologists, not because they are overworked, but for their mean quarrelsome spirit" (Darwin cited in Barlow 1967, p. 121).

³⁷During the voyage, Darwin prepared rough bird skins, just like the ones known as study skins in modern museum collections. He always applied arsenical soap on the skins, and a corrosive sublimate on the beaks and legs (Darwin 1839, pp. 600–601). Darwin stuffed the specimens with dry grass or moss and left the incision un-sewn. Back home, the larger proportion of the birds presented to the Zoological Society Museum and all of those given to the BMNH before 1857 (see below) were immediately mounted for display purposes. Nevertheless, several dozen original skins of Darwin have survived.

313

In the end, the 14 birds in alcohol, more than a dozen skins and probably four anatomical specimens were forwarded to Darwin's friend and Cambridge colleague, Thomas Campbell Eyton,³⁸ among them birds later found in the BMNH and, in the case of one steamerduck Tachyeres patachonicus (CUMZ 12/Ana/60/a/5),³⁹ in Cambridge. Eyton wrote the appendix "anatomical description" in Gould et al. 1841. The remainder of the bird skins⁴⁰ was presented to the Zoological Society of London on 4 January 1837, the same day as Darwin delivered a talk to the Geological Society entitled Observations of proofs of recent elevation on the coast of Chili (Darwin 1838).⁴¹ These Zoological Society skins attracted much interest among Darwin's contemporar-ies, as shown above. Darwin subsequently withdrew some, mainly duplicate, skins from the Society museum to forward them on loan to George Robert Gray at the BMNH. In a letter dated between June and early August 1839,⁴² Darwin wrote that he would try to negotiate with the Zoological Society for the donation of a specimen of the scale-throated earthcreeper Upucerthia dumetaria to the BMNH. The bird was subsequently given to the BMNH in early August 1839, and is now registered as BMNH 1839.8.4.1 (see Appendix).

In total, about 34–39 skin specimens from the "Beagle" voyage were subsequently given by the Zoo-logical Society to the BMNH in 1839, 1841 and 1856, on Darwin's initiative, which got him entered into BMNH registers as a donor. It seems certain that the

³⁸Since his return Darwin would have met Eyton personally at the latest towards the end of 1837 (Burkhardt and Smith 1986, vol. 2, pp. 54, 64), but it is believed that by then he had already sent the skin specimens to Shropshire, where Eyton lived. It is equally possible that the two met during the previous winter (December 1836 to March 1837) in Cambridge. Later, in November 1839, Darwin forwarded the specimens in spirit, numbered 388, 630, 650, 707, 721, 722, 728, 1037, 1043, 1050, 1157, 1309 ('Specimens in spirits of wine') and two unnumbered items, altogether 14 whole birds (cf. Keynes 2000; Burkhardt and Smith 1986, vol. 2, pp. 243– 244). It is likely that Darwin had also sent the four parts of birds to Eyton numbered 620 (tongue of woodpecker), 576 and 577 (tra-cheae of ducks, 'specimens in spirits of wine') and 3362 (stomach contents of flamingo, 'specimens not in spirits'). ³⁹It was always believed that this duck came from the Falkland Islands, but Darwin did not collect any duck during his two visits to East Falkland (cf. Barlow 1963; Keynes 2000). The anatomical specimens might have been forwarded to the Royal College of Surgeons as suggested by Darwin in a letter to Eyton on 6 January 1840 (Burkhardt and Smith 1986, vol. 2, pp. 249–250). ⁴⁰The Morning Herald, London, from 12 January 1837, p. 5, reported that Darwin presented 450 bird specimens to the Zoological Society of London on 4 January 1837. The number might be correct considering that Eyton received at least 12, but perhaps even 18, specimens of Darwin's 468 bird skins. ⁴¹Sulloway (1982b, p. 20) cited an unpublished minute of the Council of the Zoological Society, which recorded that the Society had received Darwin's "Beagle" birds on that date, accompanied by a letter in which Darwin asked the Society to dispose of any duplicate specimens, and to mount and describe the rest. ⁴²In Burkhardt and Smith (1986, vol. 2, p. 196) the letter is dated from June-October 1839, but the BMNH had already received the specimen in question in early August.

[page] 314

1839 and 1841 donations of at least nine birds were mainly triggered by Darwin asking George Robert Gray of the BMNH for help in finishing the bird chapters of the Zoology of H.M.S. Beagle (Gould and Darwin 1838, p. IX). Gray re-classified three of these birds for the book. In 1855, the Zoological Society Museum was broken up. The BMNH had the first choice of specimens. George Robert Gray, then the assistant curator of zoology, was entrusted with the selection of important material for the BMNH (Sharpe 1906, p. 251). Many type specimens, though, were overlooked by Gray, or possibly not shown to him by John Gould. Whatever the case, some of the remaining skins not chosen by Gray were subsequently sold to other bidders, such as the private collectors John Henry Gurney (1819–1890; cf. Gurney 1884, 1894) and Henry Baker Tristram (1822–1906). Most of the Zoo-logical Society bird skins from the voyage of the "Beagle", however, went into the private collection of John Gould, and from there to Philip Lutley Sclater (1829–1913), Osbert Salvin (1835–1898), Frederick Du Cane Godman (1834–1919) and Gustav Adolph Frank (1808–1880).⁴³ Confirmed Darwinian bird specimens are now housed at the BMNH, LIVCM, MANCH, CUMZ, NMSE, RMNH, USNM and VMM (cf. Appendix; Steinheimer 1999, 2000, 2001, 2003). John Gould was not only a good ornithologist, natural his-tory artist and museum curator, but an even more successful businessman. Gould used bird skins as a 'special gift' for those subscribing to his bird books (Guntert et al. 2004), shipping Darwin's bird specimens to institutions all over the world.

When the second lot of specimens was transferred at Darwin's request from the Zoological Society to the BMNH, the museum was just about to move its nat-ural history collections from the old Montagu House at Bloomsbury to the department of zoology in Robert Smirke's new British Museum building. John Edward Gray (1800–1875), the older brother of George Robert Gray and curator of zoology at the BMNH, was busy keeping the collection in some kind of order, and therefore unable to cope effectively with the new specimens arriving at the museum, with correspon-dence, or with the maintenance of collections (Gunther 1975, 1980; see also Brandon-Jones 1996). The registration of Darwin's birds in the collection registers was therefore very poor, and mainly at genus level only. Moreover, Gray and his brother replaced all the original Darwin labels with the BMNH 'Handlist' labels (Sharpe 1906, p. 85). The collection therefore lost some of its authenticity and value. Its importance diminished

⁴³Specimens of Baron Guillaume Michel Jerome Meiffren-Laugier de Chartrouse (1772–1843), cited on the BMNH labels and in the BMNH registers as deriving from Darwin (cf. Steinheimer 2003), are more likely birds from the collections of Rene Primevere Les-son, Prosper Garnot or Alcide Charles Victor Marie d'Orbigny, which tends to be confirmed by matching Darwin's field numbers and the number of available specimens.

further when new, larger and better dated collections from the same regions arrived.⁴⁴ Other institutions holding Darwin specimens, perhaps with the exception of Gurney's Norwich Castle Museum and Tristram's specimens at Liverpool, also lost track of the fact that these birds had been collected by Darwin. By the time the remaining collection of the Zoological Society of London had been sold, most of the specimens were just referred to the Society, not to Darwin. This all hap-pened before Darwin made his name with the publication of On the Origin of Species (Darwin 1859), and we must remember that, in those days, Darwin was neither as famous as he is today, nor did his collection enjoy such a good scientific reputation (cf. Sharpe 1906).

It is therefore no wonder that the whereabouts of at least 243 bird skins from Darwin's collection are now not known, despite intensive research. It is possible that authentic Darwin birds could yet come to light, but their provenance will of course most likely no longer be evi-denced by their labels. Nevertheless it is hoped that the Appendix, which gives for the first time a comprehensive list of all the birds collected by Darwin, might help future researchers to locate missing birds from the voyage of the "Beagle".

Darwin's ornithological notes and the Origin of species

Sulloway (1982b) has already shown that the mocking-birds from the Galapagos Islands rather than the finches were used in Darwin's On the Origin of Species, though some later authors have still succumbed to confusion on this matter (Strathern 1999, pp. 43–44; Riedl 2003,

p. 59).⁴⁵

In a chapter entitled 'Geographical distribution', Darwin (1872, p. 414, pagination from Odhams Press reprint edition)⁴⁶ noted:

But we often take, I think, an erroneous view of the probability of closely-allied species invading each other's territory, when put into free intercommunication. Undoubtedly, if one species has any advan-tage over another, it will in a very brief time wholly or in part supplant it; but if both are equally well fitted for their own places, both will probably hold their separate places for almost any length of time.

In short, Darwin recognized the importance of building different ecological niches ("own places") for the

⁴⁴E.g., Sulloway (1982b, p. 40) reported that the next person to

collect on the Galapagos Islands was Dr. A. Habel, shooting 460

specimens in 1868, followed by many others.

⁴⁵Confusion occurred mainly because Darwin referred to the

Galapagos finches in the second edition of his Journal of Research

(Darwin 1845).

⁴⁶This and all further quotations are also found in the 1859 edition,

though differently worded (cf. Darwin 1859, pp. 402, 48, 390, 183,

184, 184–185, 243, 349)

[page] 315

Fig. 4 Four specimens of captive-bred pigeons from Darwin's collection. Photo: Harry Taylor, Natural History Museum, London/Tring

establishment of two very similar species in the same geographical region (cf. Gause 1934, pp. 19–20). As an example he used the Galapagos mockingbird Nesomimus parvulus, the Charles mockingbird Nesomimus trifascia-tus, and the San Cristobal mockingbird Nesomimus melanotis (cf. Barlow 1963, p. 262; Keynes 1997, p. 466), of which he collected one and, respectively, two specimens of each (Darwin's numbers 2206, 3307, 3349, 3350). In his Transmutation Notebooks (Notebook B from 1837–1838; Barrett et al. 1987, p. 195) he goes on to say that: "It may be argued representative species [such as the South American and Galapagos mockingbirds] [are] chiefly found where barriers [...] interrupt[ed...] communication," stressing the importance of geographical isolation for speciation processes. The birds of the Galapagos Islands are also cited, albeit in general terms, as an example of how vague the distinction is between what one sees as a species and what scientists refer to as subspecies (Darwin 1872, p. 69). The mockingbirds of the Galapagos Islands collected by Darwin have been clas-sified both as subspecies of a single species (Mayr and Greenway 1960, pp. 447–448), and as three different (allo-)species (Dickinson 2003, p. 649).

Darwin turned again to the Galapagos birds when explaining endemism. Darwin showed that of the 26 land birds known at the time, 21, or perhaps even 23, were believed to be endemic to the Galapagos Islands,

whereas of 11 marine birds only 2 were considered endemic (Darwin 1872, pp. 405–406).

However, Darwin (1872) also drew upon several additional ornithological observations from mainland South America for his book On the Origin of Species. As example of both diversified and changed habits, what would now be termed intraspecific variation in behav-iour, Darwin (1872, p. 183) mentioned the great kiska-dee Pitangus (Saurophagus) sulphuratus, which apparently both hovers and moves like a kestrel, and perches on twigs and darts into water like the kingfisher (cf. Ridgely and Tudor 1994). Geographical separation, different hunting preferences of divided populations and availability of food might modify this behaviour to either extreme. Darwin saw this as evidence that behaviour can be variable within one population, allowing diversification and the building of different behavioural niches after geographical separation. He encountered these birds near Maldonado in May and June 1833, and shot one specimen, numbered in his collection 1216 (cf. Keynes 2000, p. 153). Darwin (1872, p. 183) continued exploring different behaviour, moving on to the 'atypical' behaviour of one species within a genus as shown by the campo flicker Colaptes campestris (see above).

Darwin (1872, p. 184) went on in his investigation of behavioural differences on the family level by turning to the order of Procellariiformes: "Petrels are the most aerial and oceanic of birds, but in the quiet sounds of Tierra del Fuego, the Puffinuria berardi [i.e. Pelecanoides urinator berard], in its general habits, in its astonishing

[page] 316

power of diving, in its manner of swimming and of flying when made to take flight, would be mistaken by any one for an auk or a grebe." What Darwin noted was that the diving petrels, Family Pelecanoididae, catch most of their food by diving under water, either from the surface or by plunging in directly from the air, which contrasts with the feeding behaviour of most other members of the Procellariiformes, though some albatrosses, and of course shearwaters, can dive short distances. Darwin's field notes read: "This bird in its habits is a complete diver [...], drops from the air like a stone, & as quickly dives to a long distance." In February 1834, after having seen these birds on the open sea between the Falkland Islands and Tierra del Fuego, and off Patagonia, Darwin obtained one specimen of the common diving petrel for his collection (Barlow 1963, p. 230, No. 1782).

A major part of the ornithology drawn upon in On the Origin of Species concerned cuckoo behaviour (see below). Darwin's first hand experience was of the bay-winged cowbird Molothrus badius and the shiny cowbird M. bonariensis. He encountered both species several times around Maldonado and Montevideo in Uruguay. Of the latter species Darwin even possessed a single egg numbered 1592 in his notebooks, which was found within a clutch of the rufous-collared sparrow Zonotri-chia capensis (Darwin in Gould et al. 1841, pp. 108–109; Barlow 1963, p. 227). Darwin did not discover the bird's brood parasitic behaviour himself. It was Syms Cov-ington who obtained the mixed egg clutch and came up with the theory about its cause. Darwin learned more about the brood parasitism of the cowbirds by speaking to local people, however, and finally consulted Azara (1802–1805) who confirmed his and Covington's suspicions. For Darwin, brood parasitism was one of the major problems to be tackled in his theory, and the cowbirds showed how such behaviour might have evolved. Cowbirds show all stages of graduation from a fully developed brood parasite to self-breeding.

The next reference to birds from the voyage of H.M.S. "Beagle" in Darwin's On the Origin of Species concerns nest building behaviour (Darwin 1872, p. 282). Darwin aimed to explain inborn behaviour in closely related but ecologically separated species, such as the song thrush Turdus philomelos and the austral thrush Turdus falcklandii. Darwin saw the American thrush on the Falkland Islands, in Tierra del Fuego and Isla de Chiloe, which are quite different habitats than those occupied by the song thrush in Britain, yet both build similar nests. Darwin had already noted in the field (Barlow 1963, p. 250): "Is said to have its nest smoothly lined with mud. I presume like our thrush." He collected at least two specimens of the austral thrush, one on East Falkland Island (Barlow 1963, p. 233, No. 1900), the other one at Chiloe (p. 250, No. 2125).

When Darwin (1872, p. 373) explained parapatric species he also made use of his own observations on the two forms of Rheidae, the greater rhea Rhea americana and the lesser rhea Pterocnemia pennata, which are, in Darwin's view, two 'successive' populations living in the

plains near the Strait of Magellan, and northwards of the plains of La Plata, with an overlap in North Patagonia. Darwin went on to elaborate that, in modern words, although the Rheidae have built themselves a similar niche to ostriches and emus, they are still their own family, thus evolving in some respects also convergently.

Discussion

Many authentic sources from the "Beagle" voyage and much secondary literature (cf. Kohn 1985) has not been consulted for this work, and it must be borne in mind that the impact on ornithology of the "Beagle" circumnavigation as a whole is slightly greater than Darwin's contribution alone. Benjamin Bynoe (1804–1865) was initially the assistant and from April 1832 the acting surgeon of the "Beagle", also carried out the duties of the ship's naturalist. He kept detailed notes on the natural history of all the places they visited. Even more detailed was the work of Captain Robert FitzRoy, who, as mentioned above, had his own small bird collection. He also entered notes on birds in his log books. Syms Cov-ington wrote a diary, including comments on natural history, and had a small private bird collection of a dozen birds or so. FitzRoy's personal steward, Harry Fuller (dates unknown), collected some birds (Porter 1985), now at CUMZ, and his Galapagos finches were appar-ently used by Darwin for his talk on 10 May 1837 (Barrett 1977, p. 40) and by John Gould for the 1841 publication. A "Beagle" mate named Charles Musters, classified as 1st Class volunteer, died of a fever "caught whilst snipe-shooting" (Barlow 1933, p. XXIII). Last but not least, the ship's clerk Edward Hellyer (?–1833) drowned on the shores of the Falkland Islands attempt-ing to retrieve a bird he had shot (Sulloway 1982a).

The important steps towards Darwin's theory of evolution have already been discussed and summarized by much greater authorities (i.a. Herbert 1974; Sulloway 1979; Mayr 1991, cited 1994; Keynes 1997), and there-fore this paper remains with the simple conclusion that birds were not Darwin's first love, and that the orni-thological knowledge gained on the "Beagle" voyage do not seem to have been indispensable for his evolution theory, although he did use many examples from captive birds⁴⁷ (Fig. 4; cf. Darwin 1872, pp. 37–38, 45–51, 58– 59, 61, 101, 167, 256–258, 288–289, 452–453) and

⁴⁷For example, the observation on hybrids of his distinguished friend Eyton, that: "amongst birds, species originally coming from distant parts of the world, are more likely to breed together, than those from nearer countries" (Burkhardt and Smith 1986, vol. 2, p. 181; Eyton 1837, p. 359: cited ibid. 182, footnote 5) made clear to Darwin that, in general terms, hybrids are less well adapted than the parent species, so that, where the overlapping of species occurs, natural selection avoids hybrids, whereas bird populations at a geographical distance from each other and which do not overlap have no need to avoid hybrids and are thus not able to distinguish between their own and different species when brought together artificially.

[page] 317

cuckoos (pp. 251, 258–261, 282).^{48 49} Ten and a half percent of the index of the 1872 edition of On the Origin of Species was taken up by subjects related to (palaeo-) ornithology. This is probably not representative of the proportion of ornithological knowledge as opposed to other sciences held by Victorian society. For Darwin, then, ornithology, especially of captive breeds, provided essential evidence for his theory of evolution.

Acknowledgements Richard Keynes (Cambridge, UK) initiated this study when he came to the Natural History Museum in 1998 to see the bird skins of his great-grandfather for his forthcoming book Charles Darwin's Zoology Notes & Specimen Lists from H.M.S. Beagle (Keynes 2000). Robert Prys-Jones (Head of Bird

⁴⁸Other ornithological examples in On the Origin of Species (Darwin 1872) are a discussion onbird races versus species with the example of the red grouse of Britain being perhaps a subspecies of the willow grouse (p. 69), survival rates of eggs and nestlings (pp. 82, 84–85), mortality rate of birds in Darwin's garden during the winter of 1854– 55 (p. 86), the range extension of bird species such as mistle thrush and swallow, causing the decline of other species (p. 92), the natural selection of camouflage colours in grouse (p. 100), the hatching of birds (p. 101), sexual selection in birds, especially in birds of paradise, rock-thrushes, peacocks and turkeys (pp. 103–104), a separate sub-species of guillemot (Uria aalge spiloptera) on the Faroe Islands (p. 106), plumage colours (p. 145), constraints of construction in bird morphology explained by the variety of kidney forms in adaptation to different pelvis-bones (p. 154), flightless and rarely flying birds (pp. 146–147, 181, 224), bird flight (p. 182), differences in the behaviour of the great tit (pp. 183, 275), the diving ability of the white-throated dipper, which is seen in relation to hardly any visible adaptation (p. 184), the webbed feet of some birds, such as waterfowl, frigate birds and grebes, compared to the long toes of some waders (p. 184), the green colour of the green woodpecker, discussed in the light of nat-ural versus sexual selection (pp. 200–201), the naked head of the vulture as an adaptation to scavenging versus the naked head of the turkey for display purposes (p. 201), the beauty of plumages and songs (p. 205), beaks of northern shoveler and Egyptian goose (pp. 227–229), bird nests, including those of swiftlets (pp. 254, 275), differences in and reasons for the timidity and tameness of birds (pp. 255, 257–258), the breeding behaviour of ostriches and pheasants (p. 262), the similarities in nest construction between closely related but geographically separated species such as wrens, thrushes and horn-bills (p. 282), fossil birds (pp. 336, 356, 365), birds as vectors for plant seeds and fresh-water shells (pp. 382–384, 402–404), a seed-eating heron (p. 403), birds from certain islands like the Bermudas, Madeira and the Canary Islands as an example of the means of geographical distribution (p. 406), birds replacing large mammals in certain regions shown by the moas on New Zealand (p. 406), near world-wide ranges of some avian genera (p. 416), and the parallel embryological development of different vertebrates, including birds, as well as the similarity of nestlings and juveniles of the same genus, such as those of different thrush species (p. 447). The glossary (pp. 493–525) ex-plains the following ornithological terms: Furcula (bone), Gallina-ceous Birds, Gallus, Grallatores, Primaries, and Scutellae (scales on bird feet). In the concordance of Darwin (1859), 180 entries are for bird(s), 63 entries for mammal(s), 58 for fish(es), 12 for reptile(s), none for amphibian(s), 117 entries for insect(s), 58 for fossil(s), 160 for terms on geology &mineralogy and 364 entries for plant(s), giving birds quite a prominent place (Barrett et al. 1981). ⁴⁹In addition to the "Beagle" birds, the following items of Darwin's collection can be found at the BMNH: 60 domestic pigeon skins, 6 domestic duck skins, 11 skeletons of ducks, 46 skeletons of pigeons and 28 skeletons of chickens, and 26 (25 still at the BMNH) skins of Persian birds from Teheran, which were previously in Sir John Murray's (1841–1914) possession (Steinheimer 2003).

Group, Natural History Museum, Tring) kindly forwarded Richard's interesting request to me, giving me the opportunity to research and data-base all known "Beagle" birds, in close cooperation with Richard Keynes and his notebook transcripts. Many thanks also to the following colleagues for comments and a wide range of additional support, from sending me reprints, books, information on birds and data on specimens to giving me access to the collections and literature in their care: Malgosia Atkinson, Ernst Bauernfeind, W.R.P. Bourne, Les Christidis, Paul Cooper, Ann Datta, Barry Davis, Rene Dekker, Clem Fisher, Alison Harding, Tony Irwin, Les Jessop, Henry McGhie, Bob McGowan, Matthew Jarron, Ingeborg Kilias, Michael Mules, Storrs Olson, Eric Pasquet, Susan Snell, Christine Stein-heimer, Ray Symonds, Juan C. Torres-Mura, Julia Voss, Effie Warr, Mic G. Wells and Robin Woods. Thanks also to Phil Rainbow and the Department of Zoology of the Natural History Museum for funding my research in Paris, Norwich, Liverpool and Leiden. Lucy Cathrow discussed linguistic aspects of the paper and I gratefully acknowledge her invaluable advice. I would like to thank Walter Bock, Armin Geus, Gordon Pater-son, Robert Prys-Jones and Walter Sudhaus for commenting on an earlier draft. Last but not least I would like to thank Franz Bairlein for his patience during the editing process.

References

Anonymous (1996) Lost Darwin specimens found in museum vaults. The Daily Telegraph, 5 January 1996:18

Armstrong P (1992) Darwin's desolate islands: a naturalist in the Falklands, 1833 and 1834. Picton, Chippenham, UK

Azara DF (1802–1805) Apuntamientos para la historia natural de los paxaros del Paraguay y Rio de la Plata, escritos por Don Felix de Azara. Viuda de Ibarra, Madrid

Barlow N (ed) (1933) Charles Darwin's diary of the voyage of H.M.S. Beagle. Cambridge University Press, Cambridge

Barlow N (ed) (1958) The autobiography of Charles Darwin 1809– 1882, with the original omissions restored. Collins, London

Barlow N (ed) (1963) Darwin's ornithological notes. Bull Br Mus Nat Hist, Hist Ser 2:201–278

Barlow N (ed) (1967) Darwin and Henslow: the growth of an idea. Letters 1831–1860. University of California Press, Berkeley

Barrett PH (ed) (1977) The collected papers of Charles Darwin, with a foreword by Theodosius Dobzhansky, vol 1. University of Chicago Press, Chicago

Barrett PH, Weinshank DJ, Gottleber TT (1981) A concordance to Darwin's Origin of Species, first edition. Cornell University Press, Ithaca, N.Y.

Barrett PH, Gautrey PJ, Herbert S, Kohn D, Smith S (eds) (1987) Charles Darwin's notebooks, 1836–1844—geology, transmutation of species, metaphysical enquiries. British Museum (Nat-ural History), London

Beer G de (1963) Charles Darwin—evolution by natural selection. Nelson, London

Blainville HMD de, Blainville MM de, Brongniart A, Savary, Cordier (1834): Rapport sur les resultats scientifiques du voyage de M. Alcide d'Orbigny [...]. Nouv Ann Mus Hist Nat 3:84–115

Bourne WRP (1992) FitzRoy's foxes and Darwin's finches. Arch Nat Hist 19:29–37

Brandon-Jones C (1996) Charles Darwin and the repugnant cura-tors. Ann Sci 53:501–510

Burchell WJ (1822) Travels in the interior of Southern Africa, vol 1. Longman, Hurst, Rees, Orme, and Brown, London

Burchell WJ (1824) Travels in the interior of Southern Africa, vol 2. Longman, Hurst, Rees, Orme, Brown, and Green, London

Burkhardt F, Smith S (eds) (1985) The correspondence of Charles Darwin, vol 1: 1821–1836. Cambridge University Press, Cambridge

Burkhardt F, Smith S (eds) (1986) The correspondence of Charles Darwin, vol 2: 1837–1843. Cambridge University Press, Cambridge

[page] 318

Chancellor G, DiMauro A, Ingle R, King G (1988) Charles Dar-win's Beagle collections in the Oxford University Museum. Arch Nat Hist 15:197–231

Clausen L (1996) How a bit of history got hidden away—fresh from last century, a bird studied by Darwin on the Beagle voyage is rediscovered. The Times, 12 February 1996:27

Curio E (1993) Die Darwinfinken. In: Grzimek B (ed) Grzimeks Tierleben, vol 9 (Vogel 3). Deutscher Taschenbuch Verlag, Munich, pp 344–349

Cuvier G (1817) Le Regne Animal distribue d'apres son organisation pour servir de base a l'histoire naturelle des Animaux et d'introduction a l'Anatomie Comparee, vol 1 (introduction, mammiferes et oiseaux). Deterville, Paris

Cuvier G (1829) Le Regne Animal distribue d'apres son organisation pour servir de base a l'histoire naturelle des Animaux et d'introduction a l'Anatomie Comparee, nouvelle edn, vol 1 (mammiferes et oiseaux). Deterville, Paris

Darwin C (1837a) Notes upon the Rhea Americana, and upon the newly described species. Proc Zool Soc Lond 5:35–36

Darwin C (1837b) Finches from the Galapagos Islands. Proc Zool Soc Lond 5:49

Darwin C (1838) Observations of proofs of recent elevation on the coast of Chili, made during the survey of H.M.S. Beagle, commanded by Capt. FitzRoy. Proc Geol Soc Lond 2:446–449

Darwin C (1839) Journal of researches into the natural history & geology of the countries visited during the voyage round the world of H.M.S. Beagle under the command of Captain Fitz Roy, R.N., 1st edn. Murray, London

Darwin C (1845) Journal of researches into the natural history & geology of the countries visited during the voyage round the world of H.M.S. Beagle under the command of Captain Fitz Roy, R.N., 2nd edn. Murray, London

Darwin C (1851) A monograph on the sub-class Cirripedia, with figures of all the species—the Lepadidae, or, pedunculated Cirripedes. Ray Society, London

Darwin C (1859) On the origin of species by means of natural selection, or, the preservation of favoured races in the struggle for life. Murray, London

Darwin C (1870) Note on the habits of the pampas woodpecker (Colaptes campestris). Proc Zool Soc Lond 38:705–706

Darwin C (1872) The [sic] origin of species by means of natural selection or the preservation of favoured races in the struggle for life, 6th edn. Odhams, London

Darwin C (1881) The parasitic habits of Molothrus. Nature 25:51

Darwin C (1890a) Journal of researches into the natural history & geology of the countries visited during the voyage round the world of H.M.S. Beagle under the command of Captain Fitz Roy, R.N., 11th edn. Murray, London

Darwin C (1890b) On the structures and distribution of coral reefs; also geological observations on the Volcanic Islands and parts of South America visited during the voyage of H.M.S. Beagle. Ward, Lock, London

Desmond A, Moore J (1995) Darwin. 2. Auflage, List, Munich, Leipzig

Dickinson EC (ed) (2003) The Howard & Moore complete checklist of the birds of the world, 3rd edn. Christopher Helm, London

Eyton TC (1869) A synopsis on the Anatidae, or duck tribe. Hobson, Wellington, Salop, UK

Field C (2003) Darwin's finch is icing on cake for Manchester Museum relaunch. 24-hour Museum News 14 October 2003

Forster JR (1778) Observations made during a voyage round the world (in H.M.S. Resolution) on physical geography, natural history [...]. Robinson, London

Fuller E (2001) Extinct birds, rev edn. Cornell University Press, Ithaca, N.Y.

Futuyma DJ (1990) Evolutionsbiologie. Birkhauser, Basel

Gause GF (1934) The struggle for existence. Williams & Witkins, Baltimore, Md.

Gould J (1837a) Remarks on a group of ground finches from Mr. Darwin's collection, with characters of the new species. Proc Zool Soc Lond 5:4–7

Gould J (1837b) Observations on the raptorial birds in Mr.

Darwin's collection, with characters of the new species. Proc

Zool Soc Lond 5:9–11 Gould J (1837c) Exhibition of the fissirostral birds from Mr.

Darwin's collection, and characters of the new species. Proc

Zool Soc Lond 5:22–24 Gould J (1837d): Three species of the genus Orpheus, from the

Galapagos, in the collection of Mr. Darwin. Proc Zool Soc

Lond 5:27 Gould J (1837e) On a new rhea (Rhea Darwinii) from Mr. Darwin's

collection. Proc Zool Soc Lond 5:35 Gould J (1837f) Exhibition of Mr. Darwin's birds, and description

of a new species of wagtail (Motacilla leucopsis) from India.

Proc Zool Soc Lond 5:77–79 Gould J (1838) Exhibition of another portion of the birds collected

by Charles Darwin, Esq. Proc Zool Soc Lond 6:4 Gould J, Darwin C (1838) The zoology of the voyage of H.M.S.

Beagle, under the command of Capt. Fitzroy, R.N., during the

years 1832 to 1836. Part 3: birds, issue 1. Smith, Elder, London,

pp 1–16 Gould J, Darwin C (1839a) The zoology of the voyage of H.M.S.

Beagle, under the command of Capt. Fitzroy, R.N., during the

years 1832 to 1836. Part 3: birds, issue 2. Smith, Elder, London,

pp 17–32 Gould J, Darwin C (1839b) The zoology of the voyage of H.M.S.

Beagle, under the command of Capt. Fitzroy, R.N., during the

years 1832 to 1836. Part 3: birds, issue 3. Smith, Elder, London,

pp 33–56 Gould J, Darwin C (1839c) The zoology of the voyage of H.M.S.

Beagle, under the command of Capt. Fitzroy, R.N., during the

years 1832 to 1836. Part 3: birds, issue 4. Smith, Elder, London,

pp 57–96 Gould J, Darwin C, Eyton TC (1841) The zoology of the voyage of

H.M.S. Beagle, under the command of Capt. Fitzroy, R.N.,

during the years 1832 to 1836. Part 3: birds, issue 5. Smith,

Elder, London, pp 97–164 Grant PR (1986) Ecology and evolution of Darwin's finches.

Princeton University Press, Princeton Grant PR, Abbott I, Schluter D, Curry RL, Abbott LK (1985)

Variation in the size and shape of Darwin's finches. Biol J Linn

Soc 25:1–39 Griffith E, Pidgeon E, Gray JE (1828) The class Aves arranged by

the Baron Cuvier, with specific descriptions. In: Griffith E (ed)

The animal kingdom arranged in conformity with its organi-zation, by the Baron Cuvier, Member of the Institute of France,

&c. [...] with additional descriptions of all the species hitherto

named, and of many not before noticed [...], vol 1 [of Aves] =

vol 6 [of Animal Kingdom]. Whittaker, Treacher, London Grinnell G (1974) The rise and fall of Darwin's first theory of

transmutation. J Hist Biol 7:259–273 Guntert M, Steinheimer FD, Gessner S (2004) The mystery of the

Gouldian birds. Tech Bull Nat (in press) Gunther AE (1975) A century of zoology at the British Museum

through the lives of two keepers 1815–1914. Dawsons, Bristol Gunther AE (1980) The founders of science at the British Museum

1753–1900. Halesworth, Halesworth, Suffolk Gurney JH (1884) A list of the diurnal birds of prey, with references

and annotations: also a record of specimens preserved in the

Norfolk and Norwich Museum. van Voorst, London Gurney JH (1894) Catalogue of the birds of prey (Accipitres and

Striges), with the number of specimens in Norwich Museum.

Porter, London Hazevoet CJ (1995) The birds of the Cape Verde Islands: an

annotated check-list. British Ornithologists' Union checklist

No. 13. British Ornithologists' Union, Tring, Herts, UK Heberer G (1959) Charles Darwin—Sein Leben und sein Werk. Die

Kosmos-Bibliothek, Band 224, Franckh'sche, Stuttgart Herbert S (1974) The place of man in the development of Darwin's

theory of transmutation. Part I. to July 1837. J Hist Biol 7:217–

258 Herbert S (ed) (1980) The red notebook of Charles Darwin. Bull Br

Mus Nat Hist, Hist Ser 7:1–164

[page] 319

Houston DC (1994) Family Cathartidae (New World Vultures). In: Hoyo J del, Elliott A, Sargatal J (eds) Handbook of the birds of the world, vol 2. Lynx, Barcelona, Spain, pp 24–41

Hudson WH (1870a) A third letter on the ornithology of Buenos Ayres. Proc Zool Soc Lond 38:158–160

Hudson WH (1870b) A eighth letter on the ornithology of Buenos Ayres. Proc Zool Soc Lond 38:748–750

Jaksic FM, Lazo I (1994) Darwin's contribution to the knowledge of Chile's terrestrial vertebrates. Rev Chil Hist Nat 67:9–26

Junker T (2001) Charles Darwin (1809–1882). In Jahn I, Schmitt M (eds) Darwin & Co. Eine Geschichte der Biologie in Portraits. Beck, Munich, pp 369–389

Keynes RD (1997) Steps on the path to the origin of species. J Theor Biol 187:461–471

Keynes RD (ed) (2000) Charles Darwin's zoology notebooks & specimen lists from H.M.S. Beagle. Cambridge University Press, Cambridge

Keynes RD (2002) Fossils, finches and fuegians—Charles Darwin's adventures and discoveries on the Beagle, 1832–1836. Harper Collins, London

Keynes RD (2003) From bryozoans to tsunami: Charles Darwin's findings on the Beagle. Proc Am Philos Soc 147:103–127

King PP (1827) Narrative of a survey of the intertropical and western coasts of Australia—performed between the years 1818 and 1822. With an appendix, containing various subjects relating to hydrography and natural history. Murray, London

King PP, Fitzroy R, Darwin C (1839) Narrative of the surveying voyages of H.M.S. Adventure and Beagle between [...] 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the Globe, 3 vols. Colburn, London

Kohn D (ed 1985) The Darwinian heritage. Princeton University Press, Princeton

Kottler MJ (1978) Charles Darwin's biological species concept and theory of geographic speciation: the transmutation notebooks. Ann Sci 35:275–297

Labillardiere JJH de (1799–1800) Relation du voyage a la recherche de La Perouse: fait par ordre de l'Assemblee constituante, pendant les annees 1791, 1792, et pendant la 1ere. et la 2de. annee de la Republique francaise, 2 vols and atlas. Jansen, Paris

Lack D (1961) Darwin's Finches—an essay on the general bio-logical theory of evolution. Harper Torchbooks, New York

Larson EJ (2001) Evolution's workshop—god and science on the Galapagos Islands. Allen Lane, London

Lesson RP (1828) Manuel d'ornithologie, ou, Description des genres et des principales especes d'oiseaux, 2 vols. Roret, Paris

Lichtenstein [M]H[C] (1823) Verzeichniss der Doubletten des zoologischen Museums der Konigl. Universitat zu Berlin nebst Beschreibung vieler bisher unbekannter Arten von Saugethier-en, Vogeln, Amphibien und Fischen. Trautwein, Berlin

Mayr E (1994) ...und Darwin hat doch recht. Piper, Munich

Mayr E, Greenway JC (1960) Check-list of birds of the world—a continuation of the work of James L. Peters, vol 9. Museum of Comparative Zoology, Cambridge, Mass.

Molina GI (1782) Saggio sulla storia naturale del Chili. Nella Stamperia di S. Tommaso d'Aquino, Bologna, Italy

Molina GI (1809) The geographical, natural, and civil history of Chili. Translated from the original Italian: to which are added notes from the Spanish and French versions, and two appendixes, by the English editor; the first an account of the archipelago of Chiloe, from the Description historial of P. F. Pedro Gonzalez de Agueros; the second, an account of the native tribes who inhabit the southern extremity of South America, extracted chiefly from Faulkner's Description of Patagonia, 2 vols. Longman, Hurst, Rees, & Orme, London

Moorehead A (1969) Darwin and the Beagle. Hamish Hamilton, London

Orbigny AD d', Gervais P (1835–1847) Voyage dans l'Amerique meridionale: (le Bresil, la Republique Orientale de l'Uruguay, la Republique Argentine, la Patagonie, la Republique du

Chili, la Republique de Bolivia, la Republique du Perou), execute pendant les annees 1826, 1827, 1828, 1829, 1830, 1831, 1832, et 1833, vol 4, part 3: oiseaux. Bertrand, Paris

Owen R, Darwin C (1840) The zoology of the voyage of H.M.S. Beagle under the Command of Captain Fitzroy, R.N., during the years 1832 to 1836, part 1: fossil mammalia. Smith, Elder, London

Peters JL et al (1934–1987) Check-List of birds of the world, vols 1– 16. Harvard University Press, Cambridge, Mass.

Porter DM (1985) The Beagle collector and his collections. In Kohn D (ed) The Darwinian heritage. Princeton University Press, Princeton, pp 973–1019

Quammen D (1997) The song of the dodo—island biogeography in an age of extinctions. Pimlico, London

Quoy J-R-C, Gaimard JP (1824) Zoologie. In: Freycinet, M. L. de (ed) Voyage autour du monde, entrepris par ordre du roi [...] execute sur les corvettes de S. M. l'Uranie et la Physicienne, pendant les annees 1817, 1818, 1819 et 1820, vol 6. Pillet, Paris

Rice T (1999) Voyages of discovery—three centuries of natural history explorations. Natural History Museum, London

Ridgely RS, Tudor G (1994) The birds of South America, vol 2: the suboscine passerines. Oxford University Press, Oxford

Riedl R (2003) Riedls Kulturgeschichte der Evolutionsbiolo-gie—Die Helden, ihre Irrungen und Einsichten. Springer, Berlin Heidelberg New York

Saint-Vincent B de, Audouin JV, Bourdon JBI, Marcellin JBG (eds 1822–1831) Dictionnaire classique d'histoire naturelle, 17 vols. Rey et Gravier, Paris

Sauer GC (ed) (1998) John Gould the bird man: correspon-dence—with a chronology of his life and works, vol 1: through 1838. Sauer, Kansas City

Schmitz S (1983) Hermes Handlexikon: Charles Darwin—Leben, Werk, Wirkung. Econ Taschenbuch, Dusseldorf

Sharpe RB (1906) Birds. In Lankester ER (ed) The history of the collections contained in the natural history departments of the British Museum, vol. 2(3). Trustees of the British Museum (Natural History), London, pp 79–515

Sobol SL (ed) (1959) Charles Darwin Autobiographie. Urania, Leipzig

Spix JB von, Martius CFP von (1824) Travels in Brazil, in the years 1817–1820: undertaken by command of His Majesty the King of Bavaria. Longman, Hurst, Rees, Orme, Brown, and Green, London

Steadman DW (1982) The origin of Darwin's finches (Fringillidae, Passeriformes). Trans San Diego Soc Nat Hist 19:279–296

Steinheimer FD (1999) The whereabouts of Darwin's bird collections. Bull Br Ornithol Club 119:141

Steinheimer FD (2000) The bird collections of Tring—restoring data to an historical collection. Nature First 18:16

Steinheimer FD (2001) Geschichte und Verbleib von Darwins Beagle Sammlung. J Ornithol 142 [Sonderheft 1]:217

Steinheimer FD (2002) Christian Ludwig Landbeck's drawings for the "Birds of Germany, Alsace and Switzerland" from the Rothschild Library, Tring, with some notes on Landbeck's and Wiebke's bird collections. Okol Vogel 24:1–43

Steinheimer FD (2003) Darwin, Ruppell, Landbeck and Co.—important historical collections at the Natural History Museum, Tring. Bonn Zool Beitr 51:175–188

Steinmuller A, Steinmuller K (1987) Charles Darwin—Vom Kafersammler zum Naturforscher. Neues Leben, Berlin

Strathern P (1999) Darwin and die Evolution. Fischer Taschenbuch, Frankfurt am Main

Stresemann E (1951) Die Entwicklung der Ornithologie von Aristoteles bis zur Gegenwart. Peters, Berlin

Sulloway FJ (1979) Geographic isolation in Darwin's thinking: the vicissitudes of a crucial idea. Stud Hist Biol 3:23–65

Sulloway FJ (1982a) The Beagle collections of Darwin's finches (Geospizinae). Bull Br Mus Nat Hist, Zool Ser 43:49–94

Sulloway FJ (1982b) Darwin and his finches: the evolution of a legend. J Hist Biol 15:1–53

Sulloway FJ (1982c) Darwin's conversion: the Beagle voyage and its aftermath. J Hist Biol 15:325–396

[page] 320

Swarth HS (1929) A new bird family (Geospizidae) from the Galapagos Islands. Proc Calif Acad Sci Ser 4 18:29–43

Swarth HS (1931) The avifauna of the Galapagos Islands. Occ Pap Calif Acad Sci 18:1-299

Taylor PB (1998) Rails—a guide to the rails, crakes, gallinules and coots of the world. Yale University Press, New Haven

Vieillot LJP (1816) L'Histoire generale et particuliere des Oiseaux, leurs moeurs, habitudes, etc. Nouveau Dictionnaire d'Histoire Naturelle, Appliquee aux Arts / A l'Agriculture, a l'Economie rurale et domestique, a la Medecine, etc. / par une Societe de Naturalistes et d'Agriculteurs. Nouvelle Edition presqu'enti-erement refondue et considerablement augmentee; avec des figures tirees des trois regnes de la nature, vol 4. Deterville, Paris

Wagler JG (1827) Systema avium. Part 1. Cotta, Stuttgart

Warren RLM, Harrison CJO (1970) Type-specimens of birds in the British Museum (Natural History)—passerines, vol 2. Trustees of the British Museum (Natural History), London

Wehner R, Gehring W (1990): Zoologie. 22. Auflage. Thieme, Stuttgart

Weiner J (1994) The beak of the finch: evolution in real time—an

extraordinary scientific adventure story about birds, biology

and the origin of species. Cape, London White G (1789) The natural history and antiquities of Selborne in

the county of Southampton: with engravings and an appendix.

White, London Wied-Neuwied M von (1820) Travels in Brazil in the years 1815, 1816, 1817. Colburn, London

Willson MF, Armesto JJ (1996) The natural history of Chiloe: on Darwin's trail. Rev Chil Hist Nat 69:149–161

Winkler H, Christie DA, Nurney D (1995) Woodpeckers—a guide to the woodpeckers, piculets and wrynecks of the world. Pica, East Sussex, UK

Woods R (1988) Guide to birds of the Falkland Islands. Nelson,

Oswestry, Salop, UK Wytsman P (ed) (1905–1914) Genera avium. Verteneuil & Desmet,

Brussels Zimmer JT (1926) Catalogue of the Edward E. Ayer ornithological library, part 1. Field Museum of Natural History, Chicago, Illinois