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History of Ecological Sciences, Part 37: Charles Darwin's Voyage on the Beagle

2010; Ecological Society of America; Volume: 91; Issue: 4 Linguagem: Inglês

10.1890/0012-9623-91.4.398

2327-6096

Frank N. Egerton,

Species Distribution and Climate Change

Click here for all previous articles in the History of the Ecological Sciences series by F. N. Egerton …I was very much interested and surprised at seeing a large black and scarlet Hemipterous insect, many moths (Zygaena) and a Cicindela, which are not found in Shropshire. I almost made up my mind to begin collecting all the insects which I could find dead, for on consulting my sister, I concluded that it was not right to kill insects for the sake of making a collection. Six years later, in October 1825, he went to the University of Edinburgh to study medicine. He stayed only two years and learned as much or more outside the classroom as he did in it. He associated with zoologist Robert Grant, who encouraged his studies on marine life and told him about Lamarck's theory of transformism, and probably discussed Erasmus Darwin's also (Desmond and Moore 1991:40). Darwin made some original discoveries about several marine invertebrates from the Firth of Forth, which he reported to the Plinian Society (Allan 1977:35, Darwin 1977, II:285–291, Desmond and Moore 1991:37–39, Browne 1995:82–87, Stott 2003:35–36). Although Darwin later remembered Professor Robert Jameson's geology lectures as boring (Darwin 1959:52), he probably heard or read Jameson's comprehensive "On the Growth of Coral Islands" (1827 [Sponsel 2009:83]). After two years, Darwin decided against a medical career and left Edinburgh. His four years at Cambridge University (1828–1831) were meant to prepare him to become a Church of England clergyman, but his most important education there came outside clerical preparatory classes (van Wyhe 2009), from friendship with Professor of Botany John Stevens Henslow (1796–1861) (Barlow 1967, Mathew 1972, Walters and Stow 2001:78–104, Walters 2004a, b, Kohn et al. 2005) and his geological field trip with Prof. Adam Sedgwick (1785–1873) in the summer of 1831 (Barrett 1974, Dolan 2004, Herbert 2005:36–46, Secord 2005). "But no pursuit at Cambridge was followed with nearly so much eagerness or gave me so much pleasure as collecting beetles" (1959:62). Darwin had been inspired by reading Humboldt's Personal Narrative of travels in Spanish America to plan his own expedition to Teneriffe, Canary Islands (Darwin 1959:67–68), but it never materialized. Then, in August 1831, Henslow received an appeal to serve as a naturalist on a naval survey ship, Beagle, under Captain Robert FitzRoy (1805–1865); Henslow declined and recommended Darwin instead (Darwin 1985–1988, I:127–129). The purpose of the voyage was to map the coastlines of South America and some oceanic islands. During most of the 1800s the best British education in physical sciences was offered to army engineers and naval officers (Ratcliff 2008:26). FitzRoy was an officer who benefited from this training, and he made important contributions to hydrography and meteorology (Mellersh 1968, Basalla 1972, Nichols 2003, Gribbin and Gribbin 2004, McConnell 2004a, b). His habits of precision and thoroughness set a good example for Darwin. Contrary to the assumption of two historians (Gruber 1969, Burstyn 1975), Darwin went on the voyage as the official naturalist, though his expenses were paid by his father, Dr. Robert Darwin. There was a substantial library aboard the Beagle, including multi-volume reference works on zoology (Stoddart 1962:118–120, Burkhardt and Smith 1985a). As a parting gift, Henslow gave Darwin a copy of Humboldt's Personal Narrative. Like all great scientists, Darwin was a hard worker, and during the almost five years voyage (27 December 1831–1832 to October 1836), he diligently observed, collected specimens, took notes, and preserved superb evidence of his work. Virtually all his evidence still exists, mostly published. All his works that he himself published are now accessible at 〈Darwin-online.org.uk〉. His surviving correspondence from the period of the voyage consists of 152 letters, about half of which he wrote (Darwin 1985–1988, I:192–504). He drew upon his records, collections, and his memories to write one of the most valuable science travel books ever published, Journal of Researches into the Geology and Natural History of the Various Countries Visited by H. M. S. Beagle 1839. Three modern books discuss his voyage (Barlow 1945, Moorehead 1969, Keynes 2003), and numerous briefer accounts add to our understanding and appreciation of what he experienced and achieved (including von Hagen 1945:169–229, Life Editors and Barnett 1960, Hopkins 1969, Armstrong 1991, Armstrong 1992, 2004, Rice 1999:230–259, McCalman 2009:15–81). Darwin made a good collection of fishes, though he said little about them in his Journal of Researches (Pauly 2004:213–240). Although he would have four entomologists describe his new species of insects, he did have interesting discussions of insects in his Journal of Researches (Riley 1882:71–73, Remington and Remington 1961). Duncan Porter points out (1980:515) that Darwin's geological notes from the voyage (1383 pages) are almost four times longer than his biological notes (368 pages), and Darwin's geological observations have now received the attention they deserve (Herbert 2005). Since Darwin had learned little geology before participating in geological field work in the summer of 1831, a few months before the voyage began, why did he so quickly become a geologist? He became deeply influenced by Charles Lyell's Principles of Geology (three volumes, 1830–1833), volume I of which was given to him by FitzRoy, while Henslow sent him volume II (1832) during the voyage (Darwin 1959:77, 101, Barlow 1967:10–11, Gruber 1985:15–18, Herbert 2005:63–70). A major attraction was that Lyell defended the uniformitarian theory that Darwin found more creditable than Georges Cuvier's catastrophist theory. Natural histories of plants and animals contained very limited theories—in contrast to Lyell's theory of geology, which seemed to encompass all geological phenomena (1959:77). Geology and natural history merged in Darwin's studies of fossils and of coral islands. The first of three volumes he published on the geology of the voyage (Freeman 1965:41) was on the structure of coral reefs. The Beagle's first landing was at St. Jago in the Cape Verde Islands on 18 January 1832 (Armstrong 2004:38–45, Chancellor and van Wyhe 2009:3–6). Darwin wanted to see the kind of tropical vegetation that Humboldt had described at Teneriffe in the Canary Islands and was at first disappointed because the Cape Verde Islands were rather arid. However, he eventually found a deep valley that retained moisture and provided the emotional experience he sought (Darwin 1988:23). The commonest bird was a kingfisher (Dacelo jagoensis) that sat on castor-oil plants and darted out to catch grasshoppers and lizards (Darwin 1839:2). He took extensive notes on marine invertebrates (Darwin 2000:9–21), but only the notes on an octopus appeared in his Journal of Researches (1839:6–7). He watched it change color, squirt ink, and when he bent down to look more closely, it squirted water at him. They departed on 8 February, and their second stop, 16–19 February 1832, was at the uninhabited (by people) St. Paul's Rocks, near the equator (Edwards 1985, Campbell 1997:42–48, Armstrong 2004:46–50, Chancellor and van Wyhe 2009:6). Darwin (1839:8) gave the location as 0°58′ North latitude and 29°5″ West longitude, and 540 miles from South America. From a distance, the rocks appeared white, due partly to a glossy white substance in some rocks and partly to accumulated bird guano. In 1813 H.M.S. Rhin visited the rocks and Lt. George Chrichton drew a map and profile chart, but in 1832 FitzRoy did both again (Edwards 1985: Figs. 1a and 2). St. Paul's highest point is only about 60 feet (18.3 m) above sea level, and the islets are only 3/4 of a mile (2 km) in diameter. The route of H. M. S. Beagle in its voyage around the world. De Beer 1967:39. St. Paul's Rocks, from the east. Illustrated London News. By the side of many of these nests a small flying-fish was placed, which I suppose, had been brought by the male bird for its partner…quickly a large and active crab ( Graspus ), which inhabits the crevices of the rock, stole the fish from the side of the nest, as soon as we had disturbed the birds. The following list completes, I believe, the terrestrial fauna: a species of Feronia and an acarus, which must have come here as parasites on the birds; a small brown moth, belonging to a genus that feeds on feathers; a staphylinus ( Quedius ) and a wood louse from beneath the dung; and lastly, numerous spiders, which I suppose prey on these small attendants on, and scavengers of the waterfowl. After reading this account, Rear-Admiral William Symonds told Darwin that he had seen at St. Paul's crabs drag young birds from nests and eat them. Darwin added this information to this account in the second edition of his Journal (1845:10). Because this is the earliest known food web (Egerton 2007:51), Darwin's brief observations now carry a historical significance that they lacked when published, and so more details about the species he observed are desirable. He collected two unknown tick species, now named Amblyomma hirtum (Neumann 1906:201–203) and A. darwini (Hirst and Hirst 1910:239–240), which are only known from two locations: St. Paul's Rocks and the Galapagos Islands. He was first to collect them in both places. His specimens are now in the Natural History Museum, London (Robinson 1926:156–158, 221–222). He also collected the argasid tick, Ornithodoros capensis, and a bird louse, Actornithophilus sp. from the brown booby nests and hippoboscid flies from dead brown boobies. No spiders were in his collections, but Scytodes sp. now lives there and may be what he saw (Edwards and Lubbock 1983:54–55). Darwin's brown moth, Erechthias darwini, was only named in 1983, because no specimen survived in his collections. It resembles the genus Darwin had in mind but is in a different family that cannot digest the keratin of feathers. It eats instead the dried seaweed of the nests (Robinson 1983). As previously mentioned, (Egerton 2009), Darwin was the second known user of what we now call a plankton net (with Lesueur earlier using a dip net). He drew a crude sketch of his, four feet deep, and first discussed its use on 10 January 1832 in his Beagle diary, which he never published (Darwin 1988:21). On 11 January he wrote in his diary (1988:22): "I am quite tired having worked all day at the produce of my net—The number of animals that the net collects is very great & fully explains the manner so many animals of a large size live so far from land." His Zoology Notes (2000:3–7) contain notes and drawings on the animals collected in his net on these two days. In Journal of Researches (1839:14–18), he first discussed phytoplankton observations after his account of St. Paul's Rocks, on 18 March; the account consists mostly of superficial anatomical descriptions. Next, he discussed numerous small crustaceans which sealers called whale-food, but he failed to make explicit here a food chain that was implicit–from minute organisms to crustaceans to whales—and only much later in his Journal of Researches (1839:189) did he even mention using his net. The day has passed delightfully: delight is however a weak term for such transports of pleasure: I have been wandering by myself in a Brazilian forest: amongst the multitude it is hard to say what set of objects is most striking: the general luxuriance of the vegetation bears the victory, the elegance of the grasses, the novelty of the parasitical plants, the beauty of the flowers.—the glossy green of the foliage, all tend to this end.—A most paradoxical mixture of sound & silence pervades the shady parts of the wood.—the noise from the insects is so loud that in the evening it can be heard even in a vessel anchored several hundred yards from the shore. Darwin's South American travels. Von Hagen 1945:228. The next day, he gushed: "I can only add raptures to the former raptures." However, rapture does not necessarily lead to discernment. A commentator on his bird watching in South America points out (Haupt 2006:54–56) that he was on the continent having the greatest diversity of bird species and in the forest that contains most of that diversity, yet he was unable to see much more than the vegetation and insects. This was early into the voyage, and he lacked the binoculars which every birder now takes to any rain forest. Near the Guardia we find the southern limit of two European plants, now become excessively common. The fennel in great profusion covers the ditch banks in the neighbourhood of Buenos Ayres, Monte Video, and other towns. But the cardoon ( Cynara cardunculus ) has a far wider range: it occurs in these latitudes on both sides of the Cordillera, across the continent. I saw it in unfrequented spots in Chile, Entre Bios, and Banda Oriental. In the latter country alone, very many (probably several hundred) square miles are covered by one mass of these prickly plants, and are impenetrable by man or beast. This was "the earliest documented transformation of a landscape by alien plants" (Mack 1989:160). …if what was told me in London is true, viz. that there are no small insects in the collections from the Tropics, I tell entomologists to look out and have their pens ready for describing. I have taken as minute (if not more so) as in England, Hydropori, Hygroti, Hydrobii, Pselaphi, Staphglini, Cuscaliones, Bimbidia, &c. &c. It is exceedingly interesting to observe the difference of genera and species from those which I know…as a specimen how little the insects are known, Noterus, according to Dic[tionnaire] Class[ique d'histoire naturelle, 17 volumes] consists solely of three European species. I, in one haul of my net, took five distinct species… Many of Darwin's biological observations recorded at Rio were not entirely new to science, though new to him and seen in newly discovered species, and so not previously published (Chancellor and van Wyhe 2009:7–49). He had been preceded in biological exploration of South America by Azara (see below), Humboldt (Egerton 2009), and Alcide Charles Victor Dessalines d'Orbigny (1802–1857); d'Orbigny collected zoological specimens for the Muséum d'Histoire Naturelle in Paris in 1826–1834 and published Voyage dans l'Amerique meridionale (10 volumes, 1834–1847; for English translated extracts, see von Hagen 1948:182–200), a notable achievement, which did not attract as wide an audience as did Darwin's Journal of Researches (Goodman 1972:301–303, Tobien 1974, Boulinier 1995, Brygoo 1995, Legre-Zaidline 2002, Moreau and Dory 2005:11–17, 81–89), though his contributions are now well appreciated (Taquet 2002). Near Rio de Janeiro Darwin observed army ants which caused other insects to either flee or be eaten, he watched a wasp paralyze a spider to provide food for its young when they hatched from eggs, and he watched spiders kill and feed on insects (1839:39–42). The insects he collected and his notes about them are now published and his specimens surveyed (Smith 1987). Some of his specimens were described in print by specialists, and Smith (1987:115–123) includes citations to that literature in his bibliography. The Zoology of the Voyage of H.M.S. Beagle (Darwin 1839–1843) was limited to vertebrates. Darwin's remarks about the people he encountered were what one expects in a travel book, but he also followed Humboldt's example of providing figures or estimates of human populations (Egerton 1970). Both travelers frequently commented on environmental conditions in places visited that either favored or inhibited population increase. Henslow had given Darwin the first two volumes of the English translation of Humboldt's Personal Narrative to take on the voyage, and the Beagle diary that he never published has frequent references to Humboldt (Darwin 1988); some of these references later appeared in his Journal of Researches 1839. He reported that Buenos Ayres had 60,000 people, Monte Video 15,000 (1839:140), Coquimbo 6000–8000 (1839:421), Charles Island in the Galapagos 200–300 (1839:456), Sydney 23,000 (1839:516), Port Louis 20,000 (1839:570), Cape Town itself 15,000, and the whole colony 200,000 (1839:575), and St. Helena Island 5000 people and 746 plant species, of which only 52 were native (1839:580). In Brazil Darwin remarked that the country would support a vast population when the land became more extensively cultivated (1839:27). When Darwin expressed shock at the massacre of Indians near Bahia Blanca, the response was, "Why, what can be done, they breed so!" (1839:120). Darwin focused on mammals and birds not in the rain forest where they were abundant though elusive, but where they were more visible on "the grassy plains around Maldonado," east of Montevideo at the mouth of Rio de la Plata. His Diary (Darwin 1988:152–162, Chancellor and van Wyhe 2009:50–58) shows he was there from 28 April to 23 July 1833. The Beagle had visited Montevideo from 26 July to 19 August 1832, and in his Journal of Researches (1839:45) Darwin mentions his arrival there on 26 July, then warns: "To prevent useless repetitions, I will extract those parts of my journal which refer to the same districts, without always attending to the order in which we visited them." Then follow his observations on mammals and birds mostly, if not entirely, from 1833. During that stay in Maldonado he hired the Beagle's cabin boy, Syms Covington, to be his servant to collect biological specimens and also to be his secretary. The number of specimens then increased. Darwin began this part of his Journal of Researches (1839:51–60) by describing a partridge (Tinamus rufescens) that was "very silly" in allowing men to ride up and lazo them. The only large mammal on this plain was a deer, Cervus campestris, which was abundant as far south as Rio Negro (latitude 41°). It could be approached close enough to kill by crawling, but it fled from men on horseback; it was not frightened by gunfire. Rodentia was very numerous in species. Hydrochaerus capybara was common. One weighed 98 pounds, was 3 feet two inches long, with a girth of 3 feet 8 inches. They lived on islands at the mouth of Rio de la Plata, and more abundantly along borders of lakes and rivers, and they ate aquatic plants. They were tame because jaguars had been eliminated and gauchos did not hunt them. On Rio Parana they were the ordinary prey of jaguars. Tucutuco (Ctenomys Braziliensis) was a small rodent with habits of a mole, living in sandy soil with a gentle slope. They were gregarious, nocturnal, and ate plant roots. The name comes from its sound; it was easily tamed. An earlier Spanish military engineer and amateur naturalist, Félix de Azara (1742–1821), had discovered that cowbirds were parasitic in their reproductive behavior (Friedmann 1964:295, Guerra 1970, Egerton 2009:263), as cuckoos were in Europe. Darwin later misremembered that he had discovered Azara's work only after he had published his Journal of Researches (Darwin 1870), probably misled because Azara is not listed in the index. In fact, he three times (1839:60–62) cited Azara's Voyages dans l'Amerique meridionale (1809, III:169–170) on brood parasitism. Darwin was quite interested in this behavior in two unrelated species on different continents. They otherwise had very different behavior, cuckoos being solitary and secretive and cowbirds being gregarious and associating with livestock. Azara had observed this behavior in Shiny Cowbirds Molothrus bonarienses, and probably Darwin observed the same species in the Maldonado area near Monte Video, though there were two other cowbird species in the area, one of which practices brood parasitism while the other does not (Ortega 1998:11, 67–68, 214). Darwin devoted an extensive amount of time and notes (Darwin 1963:233–245) to carrion-feeding hawks and vultures, which frequented the extra-tropical parts of South America. He found them interesting, though at times also disgusting. They were easily observed. Many of his notes on them were incorporated into his Journal of Researches (1839:63–69). He discussed four species of caracara (Polyborus), plus the turkey buzzard, gallinazo, and condor. Besides carrion, carranchas (P. Braziliensis) ate eggs and picked scabs from the backs of horses and mules. Azara reported that they also ate worms, shells, slugs, grasshoppers, and frogs. They also pursued gallinazos and forced them to vomit up carrion recently eaten. P. Braziliensis was larger than P. chimango and lived farther north than the latter. A third species ("allied to d'Orbigny's P. montana, but distinct") was seen in only one Patagonian valley, and a fourth species, P. Novae Zelandiae, lived on the Falklands and other islands but not on the mainland. The Turkey Vulture (Vultur aura) lived in moderately damp country from Cape Horn to North America and was solitary or lived in pairs. Gallinazos (Cathartes atratus) lived near fresh water, were abundant in Brazil and La Plata, and never occurred south of latitude 41°. Since condors were uncommon in Patagonia, he discussed them later. Darwin found pampas woodpeckers Colaptes compestris living on the pampas devoid of trees (1859:184). When Argentine immigrant naturalist William Henry Hudson (1841–1922), who had grown up near Buenos Aires (Shrubsall 2004), challenged this observation in 1870, claiming these woodpeckers only fly over the pampas from trees on one side to trees on the other, Darwin cited Azara (1802, II:311) in support of his observation (Darwin 1870). Darwin developed a persistent interest in the distribution of species and in similar species that are geographically adjacent. Two examples were rheas and foxes. He had a number of chances to observe Rhea americana on the Patagonian plains in September 1833 (Chancellor and van Wyhe 2009:66–67), and he provides a detailed account of its natural history in Journal of Researches (1839:105–109, 1963:268–276). Like ostriches in Africa and emus in Australia, females lay in a communal nest and males incubate eggs and tend the young. Although generally vegetarians, when bodies of water are drying up, they go there and catch small fish. At Rio Negro in northern Patagonia, he heard of a smaller species, and one day when a rhea was being cooked for dinner he realized that this bird was of the smaller species, and he gathered its head, neck, legs, wings, skin, and feathers and sent them to London, where, after his return, John Gould (1804–1881) named it Rhea darwinii (Gould 1837b), though d'Orbigny had already named it Rhea pennata in 1834 (Simkins 1972, Lambourne 1987:45, Tree 1991:55–56, England 2004, Sauer 2004). Haupt (2006:96–97) observed that Darwin began his observations of Rheas at Bahia Blanca in September, when there were eggs in nests, and so he had missed the breeding season, when he might have noticed that Rheas have a claw on their wings and the males have a penis. A third species, R. macrorhyncha, lives in the high Andes (Cutright 1940:171). The Beagle visited East Falkland Island 26 February–April 1833 and in March–April 1834. Darwin found the desolate island of more geological than biological interest (Darwin 1988:144–149, 228–231, Armstrong 1992, 2004:79–106, Chancellor and van Wyhe 2009:91–94). A modern explorer found more interesting wildlife there than Darwin did (Campbell 1997:108–142). Darwin easily observed the foxes, which had no fear of humans (1839:249–250). He agreed with the gauchos that black rabbits in the Falklands were introduced and were the same species as the gray rabbit of the mainland, as they produced piebald offspring when bred together. However, he was equally confident that the Falkland fox (now Dusicyon australis), which he collected, was a distinct species because the gauchos and Indians said there was no such fox on the mainland. Later, he saw the mainland fox, Vulpes Magellanicus, and agreed it was a different species. More recently, zoologists have speculated the Falkland fox may have been a hybrid between a fox and an Indian or Spanish dog (Bourne 1992:30). He knew of no other island so far from a mainland that had so large a native quadruped (land mammal). The Falkland fox stole food when it could, and Darwin predicted correctly that it would soon be exterminated (Armstrong 2004:79–106). On Chiloe, offshore from Chile, he encountered and collected another, related fox (now Darwin's fox Pseudalopex fulcipes), also fearless of humans (Darwin 1839:341). It was not found again until 1922 (Bates 1964:81–82), and now is endangered (Schafer 2010). Although the Falklands seemed desolate, the surrounding ocean did not. Darwin discovered that the kelp growing offshore sheltered a complex group of animals, and he included this important account of them in his Zoology Notes (2000:214–215). Rhea darwinii (now Pterocnemia pennata). Darwin 1838–1843:part III. Falkland fox (Canis antarcticus, now Dusicyon australis). Darwin 1838–1843: Part II. The Zoology of the sea is I believe generally the same here as in Tierra del Fuego. Its main striking feature is the immense quantity & numbers of kinds of organic beings which are intimately connected with the Kelp. This plant (the Fucus giganticus of Solander) is universally attached to rocks. From those which are awash at low water & those being in fathoms water, it even frequently is attached to round stones lying in mud. From the degree in which these Southern lands are intersected by water, & the depth in which Kelp grows, the quantity may well be imagined, but not to a greater degree than it exists. I can only compare these great forests to terrestrial ones in the most teeming part of the Tropics; yet if the latter in any country were to be destroyed I do not believe that nearly the same number of animals would perish in them as would happen in the case of Kelp. All the fishing quadrupeds & birds (& man) haunt the beds, attracted by the infinite number of small fish which live amongst the leaves: (the kinds are not so very numerous, my specimens I believe show nearly all). Amongst the invertebrates I will mention them in order of their importance. Crustaceae of ev[e]ry order swarm, my collection gives no idea of them, especially the minute sorts. Encrusting Corallines & Clytia's [bryozoans] are excessively numerous. Every leaf (excepting those on the surface) is white with such Corallines or Corallinas [coralline algae] & Spirobeae [fan worms] & compound Ascidiae [sea squirts]. Examining these with a strong microscope, infinite numbers of minute Crustaceae will be seen. The number of compound & simple Ascidiae is a very observable fact, as in a lesser degree are the Holuthuriae [sea cucumbers] Asterias [starfish]. On shaking the great entangled roots it is curious to see the heap of fish, shells, crabs, sea-eggs, cuttle fish, star fish, Planariae [flatworms], Nereidae [polychaete worms], which fall out. This latter tribe I have much neglected. Among the Gasteropoda [snails], Pleurobranchus [comb jelly] is common: but Trochus [top snails] & patelliform shells [limpets] abound on all the levels. One single plant form is an immense & most interesting menagerie. If this fucus was to cease living, with it would go many: the seals, the cormorants & certainly the small fish & then sooner or later the Fuegian man must follow. The greater number of the invertebrates would likewise perish, but how many it is hard to conjecture. There are a number of sophisticated ecological concepts expressed and implied in the passage. Darwin is comparing the productivity of the Kelp beds on the margins of the Southern Ocean to that of the tropical rain forest, distinguishing carefully between population size (number of individuals) and species diversity (number of species). He comes close to using the concepts of food-chain, food web, ecological niche and dominant species, although of course these actual terms were not coined until nearly a century later. He appreciates that there are links between microscopic and macroscopic forms, and also, very clearly, that humans are linked to their environment. The passage represents a truly integrated and holistic viewpoint… A modern book on the kelp forest quotes Darwin on it with approval (McPeak et al. 1988:130). Darwin's descendant, Richard Darwin Keynes, a zoologist who edited three books on the Beagle voyage and wrote one on it, claims (2003:218–219) Darwin as a founding father of ecology and as supporting evidence quotes the above paragraph from the Zoology Notes. The majestic Andean Condor attracted Darwin's interest, as it had Humboldt's (Egerton 2009:264–265). Humboldt saw it in the northern limits of its range; Darwin, who traveled around South America's coast from Brazil to Peru, could define its range rather precisely (1839:219–223, 1963:240–245). It inhabited the entire Andes and eastward along the coast as far north as the Rio Negro, latitude 41°, but only lived around steep cliffs. Like Humboldt, Darwin reported gaucho complaints about condors killing young goats and lambs. John James Audubon had reported (1826a, b) experiments in which he demonstrated that Turkey Vultures and crows had a poor sense of smell: he placed meat covered with paper near caged birds and they ignored it until the cover was removed. Darwin tried the same experiment with condors and got the same result (1838–1843, III:3–6). His and Audubon's experiments seemed decisive, but we now know that turkey vultures can smell but reject rotten meat; condors cannot smell (Cutright 1940:182–186, Steinheimer 2004:308, Haupt 2006:141–144). The Patagonian plains south of 41° impressed Darwin with its barrenness (1839:194–197). Some nomadic Indians lived there, but the Spanish attempt to colonize the area had failed. He collected a cactus which Henslow (1837) described and named Opuntia Darwinii. Darwin also collected a few be