DARWIN,sci-s03.bacs.uq.edu.au/biol/biol-news/november09/scott.pdf · The Voyage of the Beagle has...

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DARWIN,WALLACE

THE PRINCESSIN THE SOUTH

Australia and its wildlife had a surprising influenceon the revolutionary theories of the evolutionary titans

Charles Darwin and Alfred Russet Wallace.

STORY B1' TIM LOW

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Copyright Agency Limited (CAL) licenced copy

Australian GeographicSunday 1/11/2009Page: 94Section: General NewsRegion: National Circulation: 142,868Type: Magazines LifestyleSize: 3,990.57 sq.cms.Frequency: Quarterly

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THE POPULAR MISCONCEPTION GOES SOMETHING LIKE THIS: the young Charles

Darwin, on the voyage that shapes his concept of evolution, visits a fledgling Australia

in 1836 to study its unique wildlife. Australia in turn honours the great man and his

world-changing theories by naming its northernmost city after him.

The trouble with such a story is that it wouldn't be entirelytrue. Certainly the NT capital bears his name, but onlybecause the natural harbour around which it sprawls wasnamed Port Darwin in 1839 by a shipmate of Darwin's whowas the captain of the Beagle's subsequent surveying voyage,30 years before any settlement was founded and 20 yearsbefore Darwin spoke publicly about evolution. And Darwinappears to have had mixed emotions about Australia. Hisparting shot in The Voyage of the Beagle, his best-sellingpublished journal of the trip, is often quoted:

"Farewell, Australia! you are a rising child, and doubtlesssome day will reign a great princess in the south: but youare too great and ambitious for affection, yet not great enoughfor respect. I leave your shores without sorrow or regret."

Before we succumb to despair, it's worth noting thatDarwin disliked many of the places he visited during histime aboard HMS Beagle. New Zealand came out far worse,Darwin finding it neither pleasant nor attractive and rank-ing its Englishmen "the very refuse of society". He was toohomesick after four long years abroad to muster up muchenthusiasm for new lands, and looked back more fondly onAustralia in later diary entries and letters, eventually decid-ing that Australia was a "fine country".

The Voyage of the Beagle has more to say about convictsand squatters than parrots or kangaroos, but Australia didinfluence Darwin's thinking about evolution; it rates morethan 20 mentions in On the Origin of Species and as manyagain in its sequel, The Descent of Man. Ultimately,Australia has mattered much more to evolutionary thoughtthan this might imply; the other great man who conceivedof evolution by natural selection, Alfred Russel Wallace,wrote at length about Australasian fauna, and Australianscientists are currently at the forefront of research basedon the most fundamental evolutionary concepts (see"Evolutionary tale: defeating dengue", page 102).

J

WO HUNDRED YEARS have passed since CharlesDarwin's birth, and 15o since the publication of

L his revolutionary book, On the Origin of Species.What should we think about this amazing man who choseto sneer at Australia? The biologist in me salutes Darwinfor providing the core idea around which biology couldgrow into a major scientific discipline. Darwin gave peoplenew ways to think about themselves, their "animal" originsand their place in the world. But our antipodean gratitudeshould also go to Alfred Russel Wallace, whose definingessay on the topic was written during travels in theAustralasia region.

In his autobiography, Darwin (above, in 1878) ascribed

his success to "complex and diversified mental qualities

and conditions", including a love of science and

"a fair share of invention as well as common sense".

Darwin was not, as is often supposed, the first to con-ceive of evolution. His grandfather Erasmus was one ofmany before him to argue for the concept. Darwin's con-tribution was to identify natural selection as the mechanismthat drives evolution, by recognising that many are bornbut only the best suited survive and reproduce. Darwinexplained this in Origin, an accessible and easy-to-readbook at a time when Europe was in the midst of a goldenage of discovery and scientific inquiry.

The Beagle expedition's main task was to provide Britainwith better charts of South America. During the three anda half years of detailed survey work in those waters, Darwin,the ship's self-funded naturalist (he'd had to pay much of hisown way), pondered the forces that had shaped the continent.Geology was Darwin's first passion and the glamour

Tim Low is a Brisbane-based biologist and author with a deep

affection for Darwin, Wallace, and other 19th-century naturalistsHe has written six books, most recently The New Nature.

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12 JANUARY 1836 "The harbour-appears fine and spacious, with diff-fortnedh ores of hol izontafly stratified sanndstotie. The nearly level couIIt a is

(eo1le),uld irith thin scri11)! v tree . 1)us iyaking t 7e c curse of 5teriiity. ,

I .1RIVINON E TrRi'N( SYf.WEY HARBOUI . FROM'I'l II: VOYAGE OF THF. BEAGLE

science of that era. On his return to London, he convincedthe Geological Society that the Andes were slowly rising,thereby ending the long-standing debate between the Cata-strophists, who interpreted landscapes as the work of rarecataclysms, and the victorious Uniformitarianists, whoargued that geological formations are the result of constantslow-moving processes still occurring.

Knowing this, it's no surprise that Darwin was mostenthusiastic about Australia, not when he was admiring afinch or rat-kangaroo, but when he scaled the Blue Moun-tains, west of Sydney. He recorded his impressions in abook devoted to geological observations on his voyage:

"It is not easy to conceive a more magnificent spectacle,than is presented to a person walking on the summit-plains,when without any notice he arrives at the brink of one of these

cliffs, which are so perpendicular, that he can strike with astone (as I have tried) the trees growing, at the depth of betweenwoo and isoo feet [305-457 ml below him; on both handshe sees headland beyond headland of the receding line of cliff,and on the opposite side of the valley, often at the distance ofseveral miles, he beholds another line rising up to the sameheight with that on which he stands, and formed of the samehorizontal strata of pale sandstone."

The Beagle had sailed into Sydney on 12 January 1836, andDarwin crossed the Blue Mountains on a trip to Bathurst toglimpse the country's interior. He slept in a cosy inn at Black-heath and descended on superb convict-built roads beforestaying at Wallerawang homestead by the Coxs River. Herehe was fortunate to see platypuses "diving and playing" in thewater, although a proper examination Continued poge 98

CLOSE BUT NO AKUBRA: PREDICTIONS MISS THE MARKDarwin saw a cloudp future for Australia's mammals.

D ARWIN had the right ideabut the wrong victims in

mind when he predicted

extinction for the emu andkangaroo. After a day spent ona kangaroo hunt in NSW during

which no kangaroo was seen,Darwin predicted in TheVoyage of the Beagle a future

that proved wide of the mark:"A few years since this country

abounded with wild animals;

but now the emu is banished toa long distance and the kanga-

roo is become scarce. To both

the English greyhound has

been destructive. It may belong before these animals are

altogether exterminated, buttheir doom is fixed."

Darwin wasn't alone in imag-ining the kangaroo's doom. In1914 the Field NaturalistsSociety of South Australiasuggested they were alreadyextinct. Kangaroos are thrivingtoday, as beneficiaries ofpastures and dams created for

introduced grazing animals;emus have vanished from

some regions but remainplentiful in others. Darwin'sconcerns about hunting wereunderstandable, given the lossof bears, wolves and boarsfrom Britain. By the time of hisAustralian visit, hunters hadexterminated one, and perhapstwo, species of island emu -not that any naturalist knewthis back then. The KangarooIsland emu and King Island

emu were two dwarf speciesthat were killed for their meat

by sealers, the Kangaroo Islandspecies gone by 1827 and King

Island emus probably earlier.The irony is that Darwin

discovered a mammal in NSW

that did become extinct, but henever knew this. The Beagleleft Sydney with the first everspecimen of what was laternamed Gould's mouse, a smallyellow-brown rodent withdainty white feet. It wasprobably caught near Bathurst.Gould's mice were collectedagain in temperate Australia,but were not seen after WilliamBlandowski's 1856-57expedition to the junction ofthe Darling and the Murray.

It's uncertain why thisspecies became extinct,but feral cats seem likelyculprits. Since Darwin'sday, Australia has lostmore mammals to extinctionthan any other country.

Once common and widespread in Australia, Gould's mouse (above

from John Gould's Mammals of Australia, 1863) is now extinct.

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LIFE ON TFUE BEAGLE

'Shalooksrreoat b*autftul," wrpte Parwln of HMS $eagle (above) before

setting sail. He suffered dreadfully from seasickness throughout the voyage

LiMS EreLgle was firm launched in 1826 as a 1O-gurthrig-51oap, which was typically ve ft 4 iii[lic't {x7.53 nn) longand Zg ft 6 inches (7.47 ml wide, In top conditions, she could

make wavat o knots (16.6 kmfh).

The Beagle,set saii from Devanport on 27 flecemher i83z and dockedat Falmouth on 2 October 1836 - a total of 7740 days away from

England. For 1144 nights, Darwin slept on board in a hammock withx ft (61 cm) of space between his face and the ceiling.

Ship's cninlrany of 74 men included Capta in Robert F'3t'Roy, artiftAugustus Earle, an "r i nsljurnt nt maker" who douhEed as a

librarian, a surgeon ahcl Iris assistant, and Sym4 Cucingtan,who became Dar in's auist.ini.

Fully laden, the fieagkr weighed atlas tons (288.15 tonnes) a[ndcarried enough stores to last three monthr;. M;-.se included

pickles, dricciapples and lcmcxn juice to guard against scurvy;prescr+cd1 meat; medicine; water; gunpowder; coal and wood;

carpenter's stores; and spare masts, booms and canvas (for sails).

The stain tools of navigation in this era were' the sextant,chrgnnmeter and compae , and the Stars. The 1teugiv carried

as Cirronom;'ters. cant snspeitded in ginihals to keep it horixaillal,in a wooden box cushioned with sawdust to protect it from vibra-tions, Captain FitzRoy also insisted utt brass cannon (inswad (if

iron) -it) as not to i_nterfrrc with the ship''; four c:ntnpass rs.

Darwldspocket sextant from the &cugre_

It was presented to the Royal Geographical Society In 1912,

S r`f'ify1S

41

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of an animal was only obtained by shooting one. Darwin'sprivate journal also records a reflective moment whichanticipated his future thinking:

"I had been lying on a sunny bank and was reflecting onthe strange character of the animals of this country as comparedwith the rest of the world. An unbeliever in everything beyondhis own reason might exclaim, `Surely, two distinct Creatorsmust have been at work; their object, however, has been thesame, and certainly the end in each case is complete'."

Twenty-three years later Darwin revisited different"creations" in different places in On the Origin of Species.Australia, he would explain, had both living and fossilmarsupials, and South America had living and fossil slothsand anteaters, showing that animals on each continent wereallied by descent: "We see the full meaning of the wonder-ful fact, which has struck every traveller, namely, that onthe same continent, under the most diverse conditions,under heat and cold, on mountain and lowland, on desertsand marshes, most of the inhabitants within each greatclass are plainly related; for they are the descendants of thesame progenitors and early colonists."

Darwin didn't enjoy his destination, Bathurst - comment-ing in his personal journal about its "hideous little red brickchurch" - and he was soon on his way back to Sydney. TheBeagle sailed on to Hobart, and then to King George Sound inWestern Australia. Darwin disliked the vegetation on main-land Australia, describing it as a product of "sterile" soil, andexpressing a wish in Voyage to never walk again in suchuninviting country. Only in Tasmania was there enoughverdure to please his English eyes. A trek up Mt Wellington,

behind Hobart, turned sour when the guide chose a difficultroute, but the forest was majestic: "In many parts the eucalyptigrew to a great size, and composed a noble forest. In some ofthe dampest ravines, tree-ferns flourished in an extraordinarymanner; I saw one which must have been at least twenty feet[6 m] high to the base of the fronds, and was in girth exactlysix feet [2 m]. The fronds forming the most elegant parasols,produced a gloomy shade."

Tasmania was probably in his thoughts many years laterwhen, in Origin, he explained that similar animals foundon adjoining lands implied common origins: "Britain isseparated by a shallow channel from Europe, and themammals are the same on both sides; and so it is with allthe islands near the shores of Australia."

After King George Sound, the Beagle sailed to the Cocos(Keeling) Islands (at the time ruled by Captain John Clunies-Ross, but they became Australian territory in 1955), wherethe captain, Robert FitzRoy, took soundings, and whereDarwin pondered the formation of coral atolls and obtainedgood evidence to support his theory on their formation (see"Darwin's sinking feeling", below). Despite seeing prettyfish, Darwin grumbled that coral grottoes had been overratedas places of colour and beauty:

"I must confess I think those naturalists who havedescribed, in well-known words, the submarine grottoesdecked with a thousand beauties, have indulged in ratherexuberant language."

The Beagle then sailed for England, and Darwin's nearlyfive years of travel - from 27 December 1831 to 2 October1836 - reached an end.

DARWIN'S SINKING FEELINGEvolution wasn't the only groundbreaking theory that sailed in on the Beagle.

EVEN IF he had never

uttered the 'E' word,Darwin would still havesecured his place in science -as a geologist. He introduced

the word "atoll" to the Englishlanguage by proposing a theoryfor their formation that hasstood the test of time.

On the Cocos (Keeling)Islands, now part of Australia,

the Beagle's captain tooksoundings that contributed toDarwin's theorising. "CaptainFitzRoy found no bottom witha line 7200 feet [2195 m] inlength, at the distance of only

2200 yards [2011 ml from theshore." wrote Darwin in The

Voyage of the Beagle, "hence

this island forms a lofty sub-marine mountain, with sidessteeper even than those of themost abrupt volcanic cone."

Darwin correctly reasoned

that atolls were rings of coralgrowing ever-upwards from thegradually subsiding remains of

volcanic mountains. A fringingreef forms around a volcanic

island, growing most vigor-ously on the outer edge whereconditions for growth are bet-ter. The island eventually sinks

below water level, but the coralcontinues to grow, surviving as

a ring-shaped reef enclosing alagoon. Sand deposited on the

coral can produce a low island

that follows the curvature of

the submerged mountain.The two atolls on the Cocos

(Keeling) Islands are perched

atop a 5000 m tall seamountthat forms part of a substantialunderwater mountain range.The word "atoll" originatedwith the people of theMaldives.

Darwin devoted many pages

of his Beagle journal to adetailed explanation of atollformation. He also wrote TheStructure and Distribution ofCoral Reefs (1842), a book Corals, Darwin determined,

that grew from his Beagle were the foundation of many

observations. tropical islands.

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"Borneo and New Guinea, as alike physically as two distinct countries canbe, are zoologically wide as the poles. ..while Australia... produces birds and

quadrupeds which are closely related to those inhabiting... New Guinea.

FROM THE MALAY ARCHIPELAGO, BYALFRED WALLACE

S A MAN of inherited means, Darwin could devotethe rest of his life to independent scholarship.Wallace was born instead into a family of failing

fortunes and had to leave home at 13 to make a living. Hisfirst attempt to earn income as a naturalist foundered whenin 1852 the ship bearing him home after four years in SouthAmerica caught fire and his notes and thousands ofspecimens he hoped to sell were lost. After writing a bookabout his travels and travails, Wallace turned his sights tothe East Indies, spending eight years on the islands strungbetween Malaya and Australia, from 1854 to 1862. Whilecollecting birds and insects, he observed telling differencesbetween islands, as Darwin in the Galapagos had donedecades before him.

It was an essay sent by Wallace to Darwin that joltedthe older man into action, rushing him into print before hewas ready. Wallace had sent his article to Darwin with theintention of it being shown to the eminent geologist CharlesLyell. To Darwin's dismay, Wallace had grasped the notionof evolution driven by a surplus of individuals, an idea bothhad developed after reading Thomas Robert Malthus's "AnEssay on the Principle of Population". Malthus's radicaltreatise proved amazingly influential; his comments onoverpopulation also convinced Britain that free migrationto Australia and other outposts would strengthen ratherthan weaken English society.

Darwin's colleagues arranged for Wallace's essay to beread in 1858 to the Linnean Society in London, together withextracts from two unpublished documents encapsulatingDarwin's thoughts, written by Darwin in 1844 and 1857.Darwin then set about writing the book that would changehow people thought about their place in nature. On The Originof Species by Means of Natural Selection, or the Preservation ofFavoured Races in the Struggle for Life - to give the first editionits full title - was published the following year.

A book about Australia that was published soon after wasamong the first to lend support to Darwin and Wallace. Itsauthor, Joseph Dalton Hooker, was a botanist who travelledon James Ross's Antarctic expedition from 1839 to 1843 andwho became one of Darwin's closest friends and ardentsupporters. Hooker's volume about Tasmanian plants, whichdrew upon specimens he collected on the expedition, beganwith a spirited endorsement of natural selection, beforesummarising Australia's flora in a volume that remainsrelevant today. Darwin wrote to tell him it was the mostinteresting essay on nature he had ever read.

Wallace returned from Asia in 1862. He wrote The MalayArchipelago (1869), followed the next year by Contributions

Alfred Wallace is often pigeonholed as the man in

Darwin's shadow, when in fact he was widely admired in

his time and remains a popular scientific figure today.

to the Theory of Natural Selection. To illustrate evolution atwork Wallace invoked the kangaroo, whose sandy coatafforded a measure of disguise, and Australian seahorsesthat impersonate seaweed. Wallace proposed that only birdswith hidden nests could afford to have colourful females,nominating Australian treecreepers, sitellas, pardalotesand ground parrots as evidence.

Wallace wrote prolifically over the years about manytopics, including Australia. He edited a geography book,Australasia, expanded from an earlier publication by a Germanhistorian. In one article he proposed an Australian origin forthe parrots of the world, an idea lent support by recent DNAevidence, and in another, written for a gardening magazine,he extolled the "very beautiful" eucalypt in his garden inDorset. Wallace's knowledge of Australia helped make hisname as the world's foremost biogeographer. He definedthe continent largely by what it lacked, as is clear from TheMalay Archipelago:

"It is well known that the natural productions of Australiadiffer from those of Asia more than Continued page 102

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Butterflies 0 and birds, beetles and bugs Q were all collected by Wallace and shipped in their thousands back to England

where they were valued by scientists and collectors alike. One of the more spectacular specimens on display here is the

New Guinea shield-back bush-cricket (below, at centre top) - probably the same specimen illustrated in The MalayArchipelago. Wallace described these giant insects, which have wingspans of up to 27 cm, as "sluggish in their motions,

depending for safety on their resemblance to foliage, their horny shield and wing coverts, and their spiny legs".

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Variation within and between species is all too clear in Wallace's collections of moths 0 and butterflies, which are

prodigious in size and diversity. In the six years he spent in the Malay Archipelago, Wallace procured more than 125,000

specimens. His meticulous attention to detail, which included attaching a "locality ticket" to each specimen, helped him

hone theories about the geographical distribution of animals. One of the more unusual drawers in his collection O includes

the emerald-green wings of an Amazonian parrot and pieces of mammal skin and fur, possibly from New Guinea.

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those of any of the four ancient quarters of the world differfrom each other. Australia, in fact, stands alone: it possessesno apes or monkeys, no cats or tigers, wolves, bears, or hyenas;no deer, or antelopes, sheep or oxen; no elephant, horse, squir-rel, or rabbit; none, in short, of those familiar types of quad-ruped which are met with in every other part of the world."

But Wallace is most famous today for the line that bearshis name. Most Australians have seen it on maps withoutknowing exactly what it means. Passing between Bali andLombok and Borneo and Sulawesi, Wallace's Line separatesthe Asian faunal realm from the transitional zone betweenAsia and Australia, called Wallacea. Sulawesi, the Moluccasand the Nusa Tenggara chain make up this island domain.

Before a visit to Wallacea two years ago, I delved deeplyinto Wallace's Malay Archipelago, and like him I marvelled atthe mix of species in Sulawesi's rainforests. I saw cuscuses,honeyeaters and lorikeets which reminded me of Australia,alongside such typically Asian fare as monkeys, hornbills, anda reticulated python that lunged at my face. I came away fullof respect for Wallace, who spent eight trying years in whatwas then a very remote and unsafe region. The best of the19th-century naturalists learnt biology the hard way, spend-ing far more time in the field than do most biologists today.

Wallace showed far more interest in Australia thanDarwin, but whether or not he actually visited the place

depends on how you define it. New Guinea and the AruIslands, which were two of his collecting grounds, countbiogeographically as Australia in the same way thatTasmania does. But Wallace never set foot on what todayis Australian national territory.

ARWIN'S relationship with Australia was nothing ifnot complicated. His servant on the Beagle voyage,Syms Covington, shared his distaste for the convict-

tainted culture of Sydney. But although he wrote in hisjournal that he was "heartily happy to leave" Sydney,Covington emigrated to NSW just a few years later. Darwinarmed him with fulsome references, which mentioned hisslight deafness - attributed to his employment as Darwin'sshooter, responsible for obtaining specimens. When Darwindecided he needed more credibility as a naturalist beforepublishing on evolution, he embarked on a major study ofbarnacles and asked his ex-servant to send him Australianspecimens, which Darwin scientifically described. Covingtonprospered in Australia, eventually becoming a countrypostmaster at Pambula, on the NSW south coast.

The letters exchanged by the two friends show thatDarwin's thinking about Australia shifted. "Yours is a finecountry," he wrote Covington in 1857, "and your childrenwill see it a very great one." The letter also refers to Darwin's

EVOLUTIONARY TALE: DEFEATING DENGUEToday's cutting-edge science still dips its lid to Darwin s 150-year-old theory.

A GENERATION AGO,

hopes were high thatmosquito-borne diseasescould be banished worldwideby generous applications of

insecticides. Those hopes weredashed when mosquitoesevolved insecticide resistance.

Dreams of eliminating weedsand insect pests from cropsalso foundered when herbicideand pesticide resistance

evolved. With a better grasp ofDarwinian evolution, scientistsin Australia and abroad are

today developing new inte-grated techniques to counterthe pests that blight humanity.

A University of Queenslandteam, led by Professor Scott

O'Neill, has found a way tohalve the lives of the mosqui-toes that transmit denguefever. "Mosquitoes only

become infectious about twoweeks after they have fed on

someone with a disease, by

which time they are quite old,"Scott says. Scott's team hassuccessfully infected mosqui-

toes with a life-shorteningbacterium, Wolbachia, obtained

from fruit flies.In research funded by the

Grand Challenges in GlobalHealth initiative of the Bill &Melinda Gates Foundation,the bacterium was grown in aculture of mosquito cells forthree years until it evolvedsufficient compatibility tosurvive when injected intomosquito eggs. The infectedmosquitoes live long enoughto breed, passing the bacte-rium to their offspring, butnot long enough to pose ahigh risk of spreading disease.

The results in the laboratoryhave proved very promising,

but the real test will come whenScott's group releases theirmosquitoes into the wild, inattempts to infect mosquitopopulations, "hopefully in 2-3years", Scott says. The insects

released will carry a strain of

Wolbachia that does not killmosquitoes before they breed,but which is virulent enough toshorten their lifespans, and

thus halt the spread of dengue.Wolbachia virulence is difficultto predict because it varies in

different environments, butScott hopes that malaria aswell as dengue can be coun-

tered by this technique.

"Evolutionary thinking allowsus to identify where the prob-lems may arise and how to

respond," Scott explains.

Mosquitoes (above) - and the diseases they harbour -are piercing reminders of evolution's continuing relevance

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24 DECEMBE1 i oo. "./ know not how or to n'horn to express full'

v

niv o&hiiir otion of Darwin c book...] really feel thankful that it hasnot been 1c ft t to me to give the thcor v to the public'."

LETTER FI Ol4'f WALLA E TO NATURALIST I !ENl\YB 5! F.

t -It4 I l I k - % 4 5 1 1 1 A I I I I. i1, r.h. I,I1I. II s7 .h tly II

This Tasmanian scene appears in Wallace's 1876 book The Geographical Distribution of Animals

The palms in the background are an artistic invention - Tasmania has none.

dinner in England with Australian sheep-breeder SirWilliam Macarthur, during which he "drank someadmirable" Australian wine. Some years later, feeling moredespondent about his life than usual, Darwin wrote toCovington with a most unusual inquiry: "When I think ofthe future I very often ardently wish I was settled in oneof our Colonies... Tell me how far you think a gentlemanwith capital would get on in New South Wales."

Australia had evolved into a true princess in the south,a place Darwin thought might be better than England.Darwin so disliked sea travel, and was so often ill, that onecan hardly imagine him boarding a ship bound for Sydney,but had he done so On the Origin of Species might have beenan Australian book, and this story may have turned outvery differently.

Darwin's legacy is vast. He changed for all time ourpicture of life on earth and our place in it. Arguing clearlyand powerfully from examples drawn from all over the globe,

he showed that nature and humanity are not opposingcategories but part of the same flourishing of life. He providednature with a past that explains what it is today. He did thesame for us. His influence lives on in disciplines as diverseas medicine, agriculture, philosophy and psychology.

A century and a half after he gave the world his theory,Charles Darwin remains as relevant as ever. M

FURTHER READING

On the Origin of Species, by Charles Darwin, first published 1859 by

John Murray; The Malay Archipelago, by Alfred Wallace, first published

1869 by Macmillan and Company; Darwin: a Life in Science, by Michael

White and John Gribbin, 1995, Simon and Schuster; Where Worlds

Collide: The Wallace Line, by Penny van Oosterzee, 1997, Reed. See

also http://darwin-online.org.uk/ to access Darwin's works online.

AUSTRALIAN GEOGRAPHIC and Tim Low thank all those who

provided assistance with this article.

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