CHARLES DARWIN AND THE EVOLUTION OF THE ATLANTIC OCEAN, THE MACARONESIAN ISLANDS, AND THE AÇORES

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CHARLES DARWIN AND THE EVOLUTION OF THE ATLANTIC OCEAN, THE MACARONESIAN ISLANDS, AND THE AÇORES
AÇOREANA, 2011, 7: 39-71

                           CHARLES DARWIN AND THE EVOLUTION OF THE
                          ATLANTIC OCEAN, THE MACARONESIAN ISLANDS,
                                       AND THE AÇORES

                                                     Brian Morton

                            Department of Zoology, The Natural History Museum, Cromwell Road,
                              London SW7 5BD, U.K. e-mail: prof_bmorton@hotmail.co.uk

                   Darwin wrote in his notes for the Voyage of the Beagle when he stopped off
                        at Terceira on 20 September 1836 on his way back to England:
                        ‘I enjoyed my day’s ride, though I did not find much worth seeing’.

                                                         ABSTRACT
                           The first stop on 16 January 1832 on the voyage of H.M.S. Beagle was the
                       island of Santiago in the Cape Verde Archipelago. While there, Darwin, only
                       22 years old and with a Cambridge degree in theology, examined an upli ed
                       layer of fossiliferous sandy breccia sandwiched between lava flows. These first
                       observations eventually played an historic role in his and our understanding
                       of ocean genesis and the colonization and subsequent evolution of island
                       floras and faunas.
                           The English botanist Joseph Hooker had noted floristic similarities between
                       the Falkland Islands and Iceland, neither having hardly any indigenous
                       species, and South America and Europe, respectively. To explain this, the
                       English malacologist Edward Forbes proposed in 1846 that a great land mass
                       had existed in the Miocene encompassing northern Europe and Spain, and
                       extending out from the Mediterranean far westwards into the Atlantic Ocean
                       virtually to the coast of North America. On his return to England, Darwin
                       became skeptical of Forbes’s lost land and sent seeds of the Western Atlantic
                       fabaceans Entada gigas and Mucuna urens from Açorean beaches to the Royal
                       Botanic Gardens at Kew where they were planted, germinated and produced
                       healthy, mature, vines. Subsequently, Darwin set up his own experiments in
                       the glasshouse at Down House where he immersed the seeds of 87 species
                       of common plants in seawater for a month. He then tried to germinate them
                       and found that over half (64) had survived. By his own calculations, ocean
                       currents could thus have taken such seeds well over half way across the
                       Atlantic Ocean. He undertook similar work on dried muds collected from
                       the feet of migrating birds and concluded that no Forbesian landmass was
                       necessary to explain Hooker’s biogeographic similarities.

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                 The observations of Darwin on Santiago eventually led on Cocos-
             Keeling in the Pacific to his concept of oceanic islands having either
             risen or sunk, or of sea levels falling and rising, respectively, or both.
             Darwin thus set in place an alternative, and more plausible, theory to
             that of Forbes, which suggested that newly-emergent islands could be
             colonized naturally by plants and animals from other locations and
             that, through natural selection, such isolated individuals could evolve
             into distinct species.
                 Thus, although Darwin stayed but a few days in the Açores, the
             islands subsequently played a critical role in his understanding of the
             evolution of islands and life upon them. Terrestrially, the Açorean flora
             and fauna have been much modified by the hand of man. This is also
             true of the few low-lying wetlands, but the islands remain, because
             of their Atlantic isolation, one of the best places to test Darwin’s
             hypotheses about island colonization and to effect Macaronesian
             marine conservation.

                                               RESUMO
                 A primeira paragem em 16 de Janeiro de 1832 na viagem do H.M.S
             Beagle foi a ilha de Santiago no Arquipélago de Cabo Verde. Enquanto
             ali, Darwin, com apenas 22 anos de idade e graduado em teologia
             por Cambridge, examinou uma camada erguida de breccia arenosa
             fossilífera entalada entre derrames de lava. Estas primeiras observações
             eventualmente tiveram um papel histórico no seu e nosso entendimento
             da génese do oceano e da colonização e subsequente evolução das floras
             e faunas das ilhas.
                 O botânico Inglês Joseph Hooker notou semelhanças florísticas entre
             as Ilhas Falkland e a Islândia, nenhuma possuindo praticamente espécies
             indígenas, e a América do Sul e a Europa, respectivamente. Para explicar
             isso, o malacólogo Inglês Edward Forbes propôs em 1846 que uma grande
             massa de terra existiria no Mioceno abrangendo a Europa do Norte e a
             Espanha, e estendendo-se para fora do Mediterrâneo muito para oeste
             Oceano Atlântico adentro virtualmente até à América do Norte. No seu
             regresso à Inglaterra, Darwin tornou-se céptico quanto à terra perdida
             de Forbes e enviou sementes de fabáceas do Atlântico Oeste Entada gigas
             e Mucuna urens das praias Açorianas para os Royal Botanical Gardens
             em Kew onde foram plantadas, germinaram e produziram trepadeiras

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                       saudáveis, maturas. Subsequentemente, Darwin preparou as suas
                       próprias experiências na estufa em Down House onde imergiu em água
                       do mar durante um mês as sementes de 87 espécies de plantas comuns.
                       Experimentou então germiná-las e descobriu que mais de metade (64)
                       haviam sobrevivido. Por seus próprios cálculos, as correntes oceânicas
                       poderiam pois ter trazido tais sementes bem mais do que metade do
                       caminho através do Oceano Atlântico. Ele fez trabalho semelhante
                       com lamas secas recolhidas das patas de aves migradoras e concluiu
                       que não seria necessária a massa de terra Forbesiana para explicar as
                       similaridades biogeográficas de Hooker.
                           As observações de Darwin em Santiago levaram eventualmente, em
                       Cocos-Keeling no Pacífico, ao seu conceito de ilhas oceânicas haverem
                       subido ou afundado, ou de mares descendo e subindo, respectivamente,
                       ou a ambas as situações. Darwin então colocou uma teoria alternativa
                       à de Forbes, e mais plausível, que sugeria que ilhas recém-emersas
                       poderiam ser colonizadas naturalmente por plantas e animais de outras
                       localidades e que, através da selecção natural, esses indivíduos isolados
                       puderam evoluir em espécies distintas.
                           Assim, embora Darwin tenha permanecido nos Açores apenas alguns
                       dias, as ilhas subsequentemente desempenharam um papel crítico na sua
                       compreensão da evolução das ilhas e da vida nelas existente. Em terra, a
                       flora e a fauna Açorianas têm sido muito alteradas pela mão do homem.
                       Tal é verdade acerca das poucas zonas húmidas baixas mas, devido ao
                       seu isolamento no Atlântico, as ilhas permanecem um dos melhores
                       lugares para testar as hipóteses de Darwin acerca da colonização das
                       ilhas e para levar a cabo a conservação marinha da Macaronésia.

                            INTRODUCTION                     Darwin arrived at the University
                                                             of Edinburgh to read medicine.

                  C     harles Darwin was born on
                        12 February 1809. One of the
                   greatest influences in his early
                                                             This lasted for but two years and
                                                             in 1828 (aged 19), Darwin enrolled
                                                             at the University of Cambridge to
                   life was his paternal grandfather         read divinity. Unsuccessfully, as
                   Erasmus Darwin (1731-1802) who            it transpired. Here, however he
                   recognized and described ‘biolo-          came under the influence of two
                   gical evolution’. In 1825 (aged 16),      great men – the geologist Adam

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        Sedgwick (1785-1873), who had       not an outstanding student, in-
        become Woodwardian Professor        deed his father ‘despaired of
        of Geology at Cambridge in 1818,    him’ but, today, there is a statue
        and the botanist John Stevens       of Charles Darwin as a young
        Henslow (1796-1861), who had        man in the gardens of Christ’s
        been appointed Professor of         College, Cambridge (Figure 1).
        Botany at Cambridge contempo-       On the steering wheel of H.M.S.
        raneously with Darwin in 1825.      Beagle, however, were carved
        Darwin left Cambridge in 1931,      the words of Nelson’s exhor-
        not a clergyman, but with a B.A.    tation to his men at Trafalgar:
        in Theology, Euclid and Classics.   ‘England expects every man to do
        He is now, however, with train-
        ing by Sedgwick and Henslow
        an enthusiastic, albeit, amateur
        geologist and botanist. On 24
        August 1831, Henslow informed
        Darwin that there is a space for
        him on H.M.S. Beagle as natural-
        ist companion to Captain Robert
        FitzRoy (1805-1865).      Darwin
        accepted the invitation and on
        27 December 1831 (aged 22), he
        and the Beagle departed on her
        second voyage, principally to
        conduct a hydrographic survey
        of the coast of South America.
        The expedition was not to return
        to England until 2 October 1836,
        almost five years after it had
        set sail. In Cambridge, Darwin
        had become much influenced
        by the writings of Charles Lyell
                                            FIGURE 1. The statue of Charles Darwin
        (1797-1875), author of Principles
                                            as a young man in the gardens of Christ’s
        of Geology and took Volume I        College, Cambridge. ‘I believe that I was
        with him on the voyage and          considered by all… [to be] an ordinary boy
        received Volume II on route.It      well below the common standard of intellect’.
        has to be said that Darwin was      Charles Darwin, 1876.

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                   his duty’. Such a daily reminder      in 1462, they became impor-
                   of his duty must have kept him        tant as a stopping off point for
                   working throughout the long           the trans-Atlantic slave trade.
                   periods of boredom, tedium, sea-      Subsequently, the importance of
                   sickness and homesickness that        the islands declined. Darwin ar-
                   he endured for the 58 months of       rived at the capital, Porto Praya
                   the voyage.                           (Praia), on the island of St Jago
                      The Beagle’s last stop on its      (Santiago) on the 16 January
                   five year voyage was the Açores.      1832 (the Beagle’s first stop)
                   Although Charles Darwin was           and described his surroundings
                   singularly unimpressed by the         as wearing a ‘desolate aspect’
                   islands on his visits to Terceira     (Darwin, 1845).
                   and São Miguel in 1836, the              At the time of Darwin’s vi-
                   archipelago, along with other         sit to Santiago, the Christian
                   macaronesian      ones    would       church still believed in the cos-
                   eventually play an important          mogony of James Ussher (1581-
                   role in his understanding of          1656), Archbishop of Armagh,
                   biogeography and the evolution        Primate of All Ireland, and Vice-
                   of island isolated floras and         Chancellor of Trinity College,
                   faunas. The first stop of the         Dublin, who had identified crea-
                   Beagle in 1832 was to be the          tion as starting on Sunday 23
                   Canaries. The island’s Spanish        October 4004 B.C. Such a view
                   authorities, however, had heard       had the important support of
                   that cholera was rampant in           Sir John Lightfoot (1602-1675),
                   England and so forbade any            Vice Chancellor of Cambridge
                   landing. Instead, therefore, the      University, who even added an
                   first stop of H.M.S. Beagle was       exact time - 9 a.m. - to the pre-
                   the Portuguese colony of the          scribed date. Having established
                   Cape Verde Islands.                   the first day of creation, Ussher
                                                         calculated the dates of other bi-
                       THE CAPE VERDE ISLANDS            blical events, concluding, for
                                                         example, that Adam and Eve
                      The Cape Verde Islands are         were driven from Paradise on
                   located 450 kilometres off the        Monday 10 November 4004 B.C.
                   coast of West Africa (Senegal)           Charles Lyell, however, did
                   and were discovered in 1456.          not believe any of this. Rather, he
                   Colonized by the Portuguese           believed the Earth to be billions

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        of years old. In addition to the           When Darwin arrived on the
        debate regarding the Earth’s age,      Cape Verde Islands – as deso-
        however, the scientific world          late today through over-grazing,
        was similarly divided as to            as they were then, his mind was
        whether the geological record          debating the conflicting views
        was (i), best interpreted as a         in relation to creation, and the
        series of catastrophic events, the     arguments about catastrophe
        most recent being equated with         versus gradualism. Captain
        the deluge of Noah’s flood with        FitzRoy, who was a devout crea-
        the ark touching down on Mount         tionist, not just believing in the
        Ararat, according to Ussher,           literal truth of Genesis but con-
        on Wednesday 5 May 2348 B.C.           demning Lyell as a heretic too,
        (thereby achieving convergence
        between geology and scripture)
        or (ii), as Lyell believed, slowly
        acting processes of formation,
        erosion, deposition and defor-
        mation. The Temple of Serapis,
        at Pozzuoli, Naples, actually
        provided evidence of the latter,
        even then. Here, date mussels,
        Lithophaga lithophaga (dattero di
        mare) have made holes in the 2nd
        century columns, at a height of
        ~5.7 metres showing that they
        were originally uncovered, then
        covered and now, again, unco-
        vered by the sea – attesting, be-
        cause the columns were undis-
        turbed, slow, gradual, changes
        in sea level over historical time.
        The temple and its date mussel         FIGURE 2. The Temple of Serapis, at
                                               Pozzuoli, Naples, where date mussels,
        artifacts were so important to
                                               Lithophaga lithophaga (da ero di mare)
        Charles Lyell that an illustration     have made holes in the 2nd century
        of it (Figure 2) was used as a fron-   columns, at a height (today) of ~5.7
        tispiece to his book ‘Principles of    metres. The frontispiece to Charles Lyell’s
        Geology’ (Lyell, 1830-1833).           Principles of Geology (1830-1833).

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                   would be of little help to him in           chamid (upper) shell valves,
                   this debate.                                crab ‘fingers’ and maerl frag-
                      But the harbour at Praia                 ments, attesting to its origin.
                   on Santiago held a secret that              That is, an uplifted shallow, tro-
                   Darwin discovered. Here, on                 pical, seabed composed of large
                   Quail Island (now Ilhéu de Santa            sand grains, with a surface
                   Maria) he noted the presence of             cover of calcareous maerl nodu-
                   a horizontal white band run-                les (Darwin’s ‘coral’), attesting
                   ning through the rocks, about 30            to gentle surface wavelets, and a
                   feet (~10 metres) above sea-level           mixture of organic fragments of,
                   (Figure 3). It was composed of              mostly, mollusc shells derived
                   a sandy breccia, made of com-               from rocky and sandy inshore
                   pressed shells and ‘corals’ (actu-          and soft offshore habitats. The
                   ally maerl) and continued as far            fossils obviously suggest that
                   as the eye could see. Figure 4A             the whole area had once been
                   shows some of the fossils collec-           under water but, for Darwin, the
                   ted by this author from Darwin’s            question was: why not now?
                   stratum on Quail Island in 2009.               The geology of the Cape Verde
                   The breccia matrix contains pa-             Archipelago has been described
                   tellid and fissurellid limpets,             by Mitchell-Thomé (1972) and
                   other gastropod fragments, ve-              Figure 5 is the only detailed
                   nerid, donacid, glycymerid and              map of the geology of the

                   FIGURE 3. On Quail Island (now Ilhéu de Santa Maria) at Santiago in the Cape Verde
                   Islands, Darwin noted the presence of a horizontal white band (arrow) running through
                   the rocks about 30 feet (~10 metres) above sea-level. (Photo: B. Morton).

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        FIGURE 4. A, Some of the fossils collected by this author from the limestone of Quail
        Island in 2009. The breccia matrix contains patellid and fissurellid limpets, other
        gastropod fragments, venerid, donacid, glycymerid and chamid (upper) shell valves,
        crab ‘fingers’ and maerl fragments. B, The coral Siderastrea radians a ached to a piece
        of Darwin’s fossiliferous limestone collected from the beach at San Tome on Santiago.
        (Photo’s: B. Morton).

        twelve islands and shows that                it dips locally below sea level
        they represent a within-plate                in an asymmetrical fashion. In
        archipelago located some 500                 Figure 6, A represents the sea; B,
        kilometres west of Senegal. The              ancient volcanic rocks; C, upper
        archipelago probably dates back              basaltic lava and arrowed is a
        some 180 Ma and the islands are              thin layer of white limestone
        made up mostly of Tertiary and               between them.
        younger volcanics. The oldest                   Four years later on into the
        rocks on Maio are Mesozoic                   voyage, H.M.S. Beagle stopped
        limestone but most are Cenozoic              at the Cocos-Keeling islands
        (
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                                                                  in the Cape Verde’s. For here,
                                                                  he believed the maerl was ‘coral’
                                                                  and, in fact, corals do occur in
                                                                  the shallow subtidal of the Cape
                                                                  Verde’s, for example Siderastrea
                                                                  radians here attached to a piece
                                                                  of limestone (Figure 4B), so that
                                                                  this must have been the first
                                                                  time he contemplated how such
                                                                  coral islands evolve.
                                                                      Also on Santiago, however,
                                                                  Darwin discovered an African
                                                                  baobab tree (Adansonia). His
                                                                  plant is no longer alive (Pearson
                                                                  & Nicholas, 2007), but there is
                   FIGURE 5. The only geological map of the       still a pair of trees to be found
                   of the Cape Verde Islands showing that         on the island in the valley
                   the 12 islands represent a within-plate        extending north from the
                   archipelago located some 500 kilometres
                                                                  original capital, Cidade Velha,
                   west of Senegal. (Courtesy of Dr Lyall
                   Anderson, University of Cambridge).            on the south coast of Santiago.
                                                                  The male tree illustrated (Figure
                                                                  7, note the man standing beside
                   lost to the depths still holds true            it), and the female, are bigger
                   today, attesting not only to his               than Darwin’s and must have
                   remarkable insight, but also to                been mature when he visited
                   the significance of Quail Island               the island. Darwin also knew

                   FIGURE 6. The woodcut Darwin made of the limestone stratum on Quail Island, Cape
                   Verde, in which he shows how it dips locally below sea level in an asymmetrical fashion.
                   A represents the sea, B, ancient volcanic rocks; C upper basaltic lava with a thin layer of
                   white limestone (arrowed), between them. (A er Darwin, 1844, p. 9.)

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                                                importantly, because here his
                                                first thoughts on oceanic island
                                                formation were born and he
                                                began to question the accepted,
                                                literal, truth of creation.

                                                    FORBES’S LOST LAND

                                                   At the time of Darwin’s visit
        FIGURE 7. An African baobab tree
        (Adansonia) today, on Santiago, Cape
                                                to Cape Verde, the English natu-
        Verde Islands. (Note the man standing   ralist, malacologist, geologist,
        beside it.) (Photo: B. Morton).         palaeontologist and Professor of
                                                Natural Philosophy at Edinbugh
                                                University, Edward Forbes
        that the largest baobab trees           (1815-1854), and therefore an
        could attain a great age of 6,000       eminent person, had proposed
        years. Could this therefore, be         (Forbes, 1846) that a great land
        evidence for a lost landmass that       had existed in the Miocene (7-
        had once connected Africa with          26 Ma) encompassing northern
        these Atlantic islands? If so, his      Europe and Spain, and extending
        baobab must have sprouted in            out from the Mediterranean,
        the first week of creation, but         past the Açores (and thus way
        only if one believed in Bishop          beyond the Canaries and Cape
        Ussher’s account of Biblical            Verde), as far out into the
        chronology. But how, therefore,         Atlantic as the Sargasso Sea.
        could this tree be older than the       That is, virtually to the coast
        Quail Island fossils? The truth         of North America (Figure 8).
        is, however, much more prosaic:         Such a land largely filled the
        the trees were probably taken           Atlantic Ocean. Hence, as far as
        to the Cape Verde Islands, as           Forbes’s theory was concerned,
        seeds, by either slaves or their        Darwin’s baobab would simply
        traders just a few hundred years        be a remnant of a time when the
        previously. But, at the time,           Cape Verde’s had been joined
        Darwin was not to know that.            to Africa by this, now sunken,
        Nevertheless, Darwin’s sojourn          inter-connecting landmass.The
        on Santiago was significant in          English botanist Joseph Dalton
        a number of ways but, most              Hooker (1817-1911), at 22 (the

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                   FIGURE 8. Forbes concept of a land that largely filled the Atlantic Ocean, thereby
                   accounting for Hooker’s similarities in the plant and animal fossils of Europe and North
                   America. (From Forbes, 1846).

                   same age as Darwin when he                   -known example at the time
                   joined H.M.S. Beagle), enlisted              was the beach pea (Lathyrus ja-
                   on Captain James Clark Ross’s                ponicus), illustrated in Figure
                   Expedition to Antarctica (1839-              9, which was then thought to
                   1843).   Subsequently, Hooker                have a North American and rare
                   noted the floristic similarities             British occurrence and thus pro-
                   between the Falkland Islands                 vide evidence of a formerly con-
                   and Iceland, neither of which                tiguous Atlantic distribution.
                   had hardly anything indige-                  Edward Forbes’s lost landmass
                   nous, and South America and                  also explained this, thereby fur-
                   Europe, respectively. One well-              ther accounting for similarities

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                                             on H.M.S. Challenger that, on
                                             its famous, pioneering, voyage
                                             between 1872-1876, found life at
                                             9,000 metres. Challenger, like the
                                             Beagle, put into Porto Praia on
                                             St Iago (Santiago), Cape Verde
                                             Islands, and Henry Nottidge
                                             Moseley (1844-1891), one of the
                                             naturalists on board, was keen
                                             to see the rocks described by
        FIGURE 9. The beach pea, Lathyrus    Darwin - so significant had they
        japonicus. (B. Morton).              become.
                                                Forbes died at an early age of
        in the plant and animal fossils of   39, his theories, though interes-
        Europe and North America. It is      ting, all subsequently shown to
        now known that the beach pea,        be wrong. But the Challenger
        whose seeds can survive many         Expedition did more – its 50
        months in seawater, has a cir-       volumes of research findings,
        cum-polar distribution. Later,       largely edited by John Murray
        Darwin, after looking at seed        (1841-1914), who had been
        dispersal more scientifically,       Wyville-Thompson’s        assistant
        would offer an alternative hypo-     on the voyage and who took
        thesis to that of Forbes.            over the editing and publishing
           Forbes had also dredged           of the expedition’s results (com-
        in the Aegean from 1841-42           pleted in 1896), effectively, laid
        aboard H.M.S. Beacon giving          the foundations for the mo-
        rise to his second, Azoic (or        dern science of o ce a n o g ra p h y.
        Lifeless), theory, that no life      Mu rra y, for example, was the
        existed below a depth of ~500        first person to note the existen-
        metres. This view, too, would        ce of the Mid-Atlantic Ridge and
        later be challenged by Charles       ocean trenches and, indeed, coi-
        Wyvi l l e Th omp s on (1830-        ned the word ‘oceanography’.
        1882) who succeeded Forbes as        And, for continuing and refining
        Professor of Natural Philosophy      this newest of sciences, we have
        at Edi nb urg h Un ivers ity.        to thank the researches of three
        Thompson aimed to disprove           other 20th century physical ocea-
        the Azoic Theory and did so          nographers.

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                   ALFRED WEGENER, EDWARD
                    BULLARD AND FREDERICK
                            VINE

                      Forbes’ view of a sunken
                   land bridge across the Atlantic
                   persisted into the 20th century
                   and it was not until 1911 that the
                   German meteorologist Alfred
                   Wegener (1880-1930), noticing
                   that the fossils of identical plants
                   and animals could be found on
                   opposite sides of the Atlantic,
                   suggested that the continents
                   were ‘drifting’ apart.
                      Orthodox science, that is, those
                   believing Forbes, at the time
                   explained this by postulating
                   that land bridges, now sunk, had
                   once connected far-flung conti-        FIGURE 10. The maps in the 1966
                   nents. Wegener also, however,          edition of A. Wegener’s book illustrating
                                                          continental dri for three epochs.
                   noted the close fit between the
                   coastlines of South America and
                   Africa and wondered whether
                   the continents might not have          (1907-1980), who became profes-
                   been joined at one time. He ar-        sor of Geodesy and Geophysics
                   gued the continents could drift        at Cambridge University, later
                   about, laying down the ground-         produced a computer fit of the
                   work for the 1950’s palaeomag-         continents that Wegener had
                   netism research that reconstruc-       called ‘Pangaea’, showing just
                   ted the world’s historical geolo-      how the continents do actually
                   gy in terms of a unified theory        fit very closely into a jigsaw-like
                   of continental drift. Figure 10        picture.
                   shows the maps in the 1966 edi-            In 1963, the Cambridge geo-
                   tion of Wegener’s book illustra-       physicist Frederick Vine (born
                   ting continental drift for three       1939) proved the concept of sea-
                   epochs. Edward Crisp Bullard           -floor spreading by studying

Paginação_21.indd 51                                                                        19-01-2012 11:51:19
52                           AÇOREANA               2011, Suplemento 7: 39-71

        the reversals in the polarity of     they were planted, germinated
        the Earth’s magnetic field. Vine     and produced healthy mature
        showed that the parallel bands       vines. Clearly, these and, indeed
        of palaeomagnetic anomalies          other seeds of several species of
        on either side of oceanic moun-      sea beans are capable of survi-
        tain ridges resulted from the        ving Gulf Stream transport from
        combined effects of reversals in     the tropical Western Atlantic to
        the polarity of Earth’s magnetic     the Açores, including Mucuna
        fields and the generation of new     urens as well as M. sloanei and
        floor along the axes of the rid-     M. fawcetti. The seeds of these
        ges. Today, through the science      three species of coastal legumi-
        of geomagnetism, we can inter-       nous vines share the characteris-
        pret the evolution of the earth’s    tic of a darkly pigmented hilum
        continents and oceans over time,     encircling five-sixths of the mar-
        explaining why similar fossils       gin of each. The embryo within
        occur on different continents.       the seed is protected by a thick,
        Through the pioneering work of       resistant, exocarp capable of
        Wegener, Bullard and Vine, we        prolonged exposure to seawater
        now know an enormous amount          (Gunn et al., 1976).
        about the Earth’s seabed, inclu-         In fact, the Açores receives a
        ding its age(s) and about the pla-   variety of seeds and vegetative
        tes that form the continents and     dissemules, many of which ori-
        are separated from each other        ginate from the tropical Western
        by the mid-oceanic ridges. The       Atlantic (Table 1). The sea heart,
        Açores Archipelago sits on the       Entada gigas, is the most com-
        Mid-Atlantic Ridge.                  mon disseminule on Açorean be-
                                             aches, delivered by Gulf Stream
             DARWIN’S EXPERIMENTS            currents from Caribbean shores.
                                             Entada gigas is a large, climbing,
           Upon his return to England,       tropical, coastal, vine producing
        Darwin became skeptical of           large chocolate-brown seeds up
        Forbes’s lost land and after re-     to six centimetres in diameter
        ceiving several seeds of the         and some two centimetres thick.
        Western Atlantic Entada gigas        The seeds are buoyant and can
        and Mucuna urens from Açorean        remain afloat at sea for at least
        beaches, sent them to the Royal      two years (Gunn et al., 1976).
        Botanic Gardens at Kew where         Veríssimo (1989) suggested that

Paginação_21.indd 52                                                                    19-01-2012 11:51:19
MORTON: DARWIN, THE ATLANTIC AND THE AÇORES                                     53

                   TABLE 1. Plant disseminules recorded from Açorean beaches (a er Gunn et al., 1976).
                       Disseminule                Common name             Family           Source, habit and habitat
                       Astrocaryum sp.            Starnut palm            Palmaceae        Caribbean; wet lowland forest tree

                       Calocarpum mammosum        Egg fruit               Sapotaceae       Tropical America; tree

                       Crescentia cujete          Calabash tree           Bignoniaceae     New World tropics; tree

                       Dioclea reflexa            Sea purse               Leguminoseae     Asia; vine

                       Entada gigas               Sea heart               Leguminoseae     Caribbean; climbing vine

                       Manicaria saccifera        Sea coconut             Palmaceae        American tropics; coastal tree

                       Merremia discoidosperma    Many’s bean             Convolvulaceae   Central America; woody, wet forest

                       Mucuna sloanei             True sea bean           Leguminoseae     Caribbean vine

                       Sacoglo is amazonica       Cojon de burro          Humiriaceae      South America; lowland forest tree

                       Sapindus sapinaria         Black pearl/Soapberry   Sapindaceae      American tropics; tree

                   Christopher Columbus (1451-                                 germinated after an immersion
                   1506), upon finding E. gigas                                period of 28 days and a few
                   seeds along the shores of either                            survived immersions of 137
                   Porto Santo or Madeira during                               days. Nine legumes were tried
                   his residence on the islands be-                            and, with one exception, all
                   tween ~1479-1482, used the ob-                              died, including the common
                   servation to support his argu-                              pea (Pisum sativum) although, as
                   ment for, an as yet undiscove-                              noted above, it is known that the
                   red (then in ignorance of Viking                            sea pea occurs in North America
                   achievements and the evidence                               and along the southeast coast
                   thereof on Newfoundland), west-                             of England, for example on the
                   ward land – the Americas – or,                              Dungeness shingle banks, on the
                   rather, as he thought, Asia.                                southeast coast of England, and
                      As a consequence of the                                  has a circum-polar distribution.
                   results of the Kew experiments,                                Darwin also noted that
                   Darwin set up his own in the                                many of the seeds, without
                   Old Laboratory and Greenhouse                               either the capsule or fruit, sank
                   in the grounds of Down House                                within a few days and could
                   where he immersed the seeds of                              not, therefore, have survived a
                   87 species of common plants in                              long sea voyage. Later, he tried
                   seawater for a month. He then                               larger fruits and capsules and
                   tried to germinate them and, to                             eventually to dry and then try the
                   his surprise, 64 subsequently                               stems and branches of 94 plants

Paginação_21.indd 53                                                                                                        19-01-2012 11:51:19
54                            AÇOREANA                2011, Suplemento 7: 39-71

        with ripe fruits. The majority        to evolve into distinct species.
        sank quickly, including fresh         Thus, Darwin’s interpretation
        cuttings whereas dried ones           of the stratum on Quail Island
        floated for longer. For example,      initially stimulated him to think
        ripe hazelnuts sank immediately       about how oceanic islands are
        whereas dried ones floated            formed, but his experiments on
        for 90 days and subsequently          seed dispersal also demonstrated
        germinated, as did the ripe           to him how such new entities
        seeds of Helosciadium (water          might be colonized. This latter
        parsnip) and an asparagus-plant       observation ultimately led to his
        (Asparagus officinalis) with ripe     great theory about the evolution
        berries (85 days). Of the 94 dried    of life itself.
        plants, 18 floated for >18 days.
        Darwin consulted ‘Johnston’s                    THE AÇORES
        Physical Atlas’ wherein it was
        stated that the average rate of          The Açores archipelago is
        several Atlantic currents is 33       located on the Mid-Atlantic
        miles·day-1, with some running        Ridge on its only triple junction.
        at 60 miles·day-1, and concluded      The discovery of the Açores,
        that 14 out of 100 (14%) plants       ac co rd in g to 1 4 t h ce n tu ry
        might be floated across 924 miles     Genoese maps, the Portulanos,
        of sea and if blown to a favourable   seems to have taken place
        spot, would germinate. By his         between 1317 and 1339 (Bento,
        own calculations, therefore,          199 4 ) bu t th e Po rtu g u e s e
        ocean currents could thus have        navigator Diogo de Silves is
        taken such seeds well over half       credited with the discovery of
        way across the Atlantic, that is,     the islands in 1427. Portuguese
        easily, to the Açores.                settlers first colonized the islands
            Darwin’s experiments thus         permanently beginning with
        set in place an alternative, more     Santa Maria in 1432, São Miguel
        plausible, theory to that of          in 1439 and the other seven
        Forbes and which suggested that       subsequently. Darwin and the
        newly emergent islands could be       Beagle stopped off in the Açores
        colonized by plants and animals       (the last stop) for six days on the
        from other locations and that         way back to England and visited
        through natural selection such        Terceira on 20 September 1836
        isolated individuals could come       and São Miguel subsequently,

Paginação_21.indd 54                                                                      19-01-2012 11:51:20
MORTON: DARWIN, THE ATLANTIC AND THE AÇORES                         55

                   although he probably did not                the nine Açorean islands to have
                   disembark the ship at the latter.           a sedimentary cover and has a
                   His comment about Terceira                  rich fossil record dating from
                   (Darwin, 1845) was:                         the Late Miocene-Early Pliocene
                                                               to the Pleistocene (Madeira et
                   ‘I enjoyed my day’sride, though I           al., 2007). As on Santiago in the
                      did not find much worth seeing’.         Cape Verde Islands, there is on
                                                               Santa Maria an uplifted Early
                      We h ave to remembe r,                   Pliocene fossiliferous stratum
                   however, that Darwin was                    and Janssen et al. (2008, figure
                   terribly homesick, indeed had               3) illustrate an accumulation
                   been for many months.          He           of Cavolinia marginata in a
                   even dismissed Australia as                 crustacean burrow from the
                   uninteresting! There, he did not            Lower Pliocene of Cré, on this
                   even see a kangaroo! As we now              island. Ages of fossil molluscs
                   know, however, the Açorean                  recovered from Santa Maria
                   islands are extremely interesting,          range from 10.03 to 2.24 Ma
                   especially geologically. Santa              (Kirby et al., 2007).
                   Maria is the oldest island of the                Similarly, to the rear of the
                   Açores and arose from the sea               long expanse of sandy beach
                   in the Late Miocene ~7 million              on the southern shore of Porto
                   years ago. It is the only one of            Santo, one of the Madeiran
                                                               islands, there is a raised beach
                                                               of offshore sand (Figure 11) not
                                                               unlike that seen on the coast at
                                                               Tarrafal on Santiago (Cape Verde
                                                               I s la n d s ). H e n ce , th ro u g h o u t
                                                               Macaronesia, there is ample
                                                               evidence of uplifted marine
                                                               sediments attesting not only to
                                                               active vulcanism over relatively
                                                               recent time but to the insight of
                                                               Darwin in recognizing this very
                   FIGURE 11. A raised beach towards the       early on during his voyage with
                   rear of the long sandy beach on the south   the Beagle.
                   coast of Porto Santo, Madeira. (Photo: B.        We also know that sometime
                   Morton).                                    between 3.1-3.6 million years

Paginação_21.indd 55                                                                                19-01-2012 11:51:20
56                               AÇOREANA                  2011, Suplemento 7: 39-71

        FIGURE 12. The map of the Gulf Stream created by Benjamin Franklin on 2 May 1775.
        He thought of it as a river.

        ago, after the Middle Pliocene            by Benjamin F ra n k lin (1 7 0 6 -
        upli of the Central American              17 9 0 ), w h o was postmaster
        Seaway, the connection between            general of the American colony
        the Pacific and Atlantic Oceans           at the time, during a voyage from
        was closed at the approximate             England to the soon to become
        position of modern Panama                 United States in 1775 (Figure 12).
        (Vermeij & Rosenberg, 1993), iso-          He recorded that the current was
        lating coral reefs in the Caribbean       some 3.5 °C warmer than the
        and creating the modern pa ern            surrounding sea and by taking
        of circulation of both oceans, but        regular temperature readings
        importantly for the Atlantic – the        could trace its course. It is this
        Gulf Stream –, first mapped               current which allows the Azores

Paginação_21.indd 56                                                                           19-01-2012 11:51:20
MORTON: DARWIN, THE ATLANTIC AND THE AÇORES              57

                   to be colonized sporadically by       colonization times and routes
                   species with long-lived larvae        but also marine biogeography.
                   from the Caribbean, such as              Today, the Açorean islands
                   the taenioglossan tonnoidean          are influenced by the North
                   gastropod Charonia variegata          Atlantic Current at the surface
                   (Lamarck, 1816) recorded only         and by water arriving from
                   once from Faial at 15 metres          the Mediterranean at mid-
                   depth (Gofas & Beu (2002). Also       water depths. The variety of
                   in the early Pliocene, the Bering     currents influencing the islands
                   Strait between Alaska and Siberia     at different depths thus has the
                   opened and cool-temperate and         potential therefore to create
                   polar marine species were able        in the Açores a meeting place
                   to move between the North             of marine plants and animals
                   Pacific and the Arctic-Atlantic       from different locations (Morton
                   basins (Vermeij, 1991).               et al., 1997). It should therefore
                      Such relatively recent histori-    be possible to monitor past
                   cal events have been responsible      and present colonization (and
                   for not just the Atlantic circula-    spreading) events to not just test
                   tion pattern we see today, but        Darwin’s theory of island and
                   also for the species of marine        biological evolution but to expand
                   plants and animals found on           it into a programme that tests
                   the shores and shallow subtidal       current ideas about the human-
                   waters of the Macaronesian, in-       assisted introduction of exotic
                   cluding Açorean, islands as they      species (Morton & Bri on, 2000b).
                   arose successively either prior or
                   subsequent to these major events         The above-described variety
                   (Morton & Britton, 2000a). We,        of ocean currents is predomi-
                   therefore, know much about the        nantly responsible for the histo-
                   circulation of the Atlantic Ocean     rical import onto the Açorean is-
                   and how it has evolved and in         lands’ shores of its characteristic
                   broad terms how and when the          marine flora and fauna (Morton
                   marine biota of the Açorean is-       & Britton, 2000a). Just as Darwin
                   lands arrived. That the islands       showed with plant seeds,
                   emerged either before or after        Dinesen (pers. comm.) has sug-
                   these events, however, make           gested that the planktonic lar-
                   them a chronometer for the tes-       vae of many marine invertebra-
                   ting of theories related to island    tes may spend up to four weeks

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58                           AÇOREANA               2011, Suplemento 7: 39-71

        in the upper water mass where        been colonized since the early
        current speeds of ~1.8 km h-1 are    part of the 15th century, the ori-
        common. Such larvae, again as        gins and routes of such intro-
        with Darwin’s seeds, could thus      ductions have been lost in time.
        be transported ~1,200 kilome-        Morton & Britton (2000b) have
        tres from their spawning site and    reviewed this subject.
        hence easily reach Açorean waters.
        Subsequently, such larvae may              AÇOREAN BIOTIC
        travel for up to three weeks in              ENDEMICITY
        near-bottom waters before set-
        tling is required for post-meta-        The terrestrial environment
        morphic survival. During this        of the Açores has been much
        time, at a common speed for near     modified by man since the initial
        bottom advection of 0.18 kilome-     colonization of the islands in the
        tres hour-1 (50 mm second-1),        early part of the 15th century.
        the larvae could be transported      Thus, of the ~3,000 species of
        horizontally for a further 90 ki-    vascular plants found on the
        lometres, during which time          Açores, only 72 [68 according
        settlement-inducing cues could       to Borges & Gabriel, 2009] are
        be encountered. Hence, it is         endemic (the Açorean enigma;
        not surprising and despite their     Carine & Schaefer, 2009) and,
        isolation, that the marine envi-     even so, some 52 of these
        ronment of the Açorean islands       are considered threatened.
        comprises communities, which         Borges & Gabriel (2009) have
        contain species that are largely     catalogued the terrestrial taxa
        encountered elsewhere, notably       of the Açores and identified
        in the Mediterranean (Morton         a total of 4,515 species. Of
        & Britton, 2000a) and that are       these, 420 are considered to be
        transported to them in surface       endemic: that is, 267 arthropods,
        ocean currents.                      68 spermatophytes, 12 lichens,
            Thus, both algae and animals     9 bryophytes, 49 gastropods, 3
        may have arrived at the Açores       nematodes and 13 chordates.
        with ocean currents, or through      Some endemic terrestrial bio-
        rafting on mats of floating vege-    diversity may have been lost
        tation or wood and via human         because it is estimated that,
        interventions although, in the       overall, but 2% of natural forest
        latter case, as the islands have     remains on the islands.

Paginação_21.indd 58                                                                    19-01-2012 11:51:20
MORTON: DARWIN, THE ATLANTIC AND THE AÇORES               59

                      This has had readily identi-       the degree of isolation, that is,
                   fiable consequences for some          the distance from a mainland
                   species and in the past, the sta-     and its biota. In the case of the
                   tus of the endemic Açorean bull-      Açores, some 1,200 kilometres
                   finch – the priolo – (Pyrrhula mu-    from mainland Europe, combi-
                   rina) was considered threatened       ned with the relative youth of
                   although it is possibly making        the islands, the low level of en-
                   a recovery in some locations.         demicity is hardly surprising.
                   Interestingly, this species is to-    Further, even though they are
                   day restricted to the eastern end,    volcanic, oceanic islands are ty-
                   Nordeste, of the island of São        pically unproductive. This is be-
                   Miguel. In Darwin’s time, howe-       cause such young, steep, islands
                   ver, it would have had a much         cannot hold water nor retain nu-
                   wider distribution on this island     trients and this helps us under-
                   and thus if he had landed on São      stand why the carrying capacity
                   Miguel, he may well have seen it,     of islands is low. That is, they
                   easily recognized it as subtly        cannot sustain the energy-ex-
                   different from the European           pensive lives of large animals,
                   bullfinch (Pyrrhula pyrrhula) and,    either herbivorous or carnivo-
                   thereby, not only considered the      rous. Similarly, there are few en-
                   Açores to be more interesting         demic species of marine plants
                   than he did, but obtaining a clear    and animals on such young is-
                   example of island endemicity. We      lands (Morton & Britton, 2000a,
                   have to remember, moreover, that      b). Some algae, a couple of
                   Darwin did not discover the signi-    fishes, a chiton, an ophiuroid,
                   ficance of the Galapagos finches’     a few sponges, barnacles and,
                   endemicity until a er his return      possibly, some of the 20 rissoid
                   to England – the priolo, however,     gastropods recorded from the
                   had he seen it, might well have       Açores, including Alvania medio-
                   revealed it to him whilst on the      littoralis (Ávila et al., 2008), are
                   voyage.                               considered to be endemic. As
                      Of the 107 species of Açorean      more research is undertaken,
                   terrestrial snails, some 50% (49)     however, so either conspecifics
                   are endemic (Borges & Gabriel,        or very closely related species
                   2009). Island endemicity is re-       are found nearby, such as in the
                   lated to age, size, elevation,        Mediterranean, the Canaries or,
                   climate, niche availability and       even, on seamounts elsewhere.

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60                            AÇOREANA               2011, Suplemento 7: 39-71

        Frias Martins (1995) recognized       northern shore of the island of
        the Açorean rocky shore elobiid       São Jorge. The sea grass Ruppia
        snail Ovatella vulcani as endemic,    maritima is known only from Fajã
        but as this is a pulmonate, like      dos Cubres (Morton et al., 1995)
        its endemic terrestrial leptaxine     whilst within the sediments of
        relatives, this may well be true.     Fajã de Santo Cristo, also on São
        The intertidal limpet Patella can-    Jorge, occurs the only known,
        dei gomesii is considered to be       probably introduced, popula-
        an Açorean endemic subspecies         tion of Venerupis decussata in the
        (Cúrdia et al., 2005).                Macaronesian islands (Morton &
           With such a wide variety of        Tristão da Cunha, 1993). Both of
        degrees of exposure to wave ac-       these lagoons have been descri-
        tion in the Açorean littoral, there   bed in general terms by Morton
        is a propensity for great mor-        & Tristão da Cunha (1993) and
        phological variation as has been      Morton et al. (1995, 1998), and
        demonstrated for the only com-        there is a marsh at Lajes on the
        mon intertidal littorine, Littorina   island of Pico and which has
        striata (Backeljau et al., 1995).     been described, also in general
        Without care, such ecopheno-          terms, by Morton et al. (1996).
        typic variation may lead to the       All these isolated miniature la-
        unwarranted description of en-        goonal wetlands are threatened
        demicity.                             an d a re in u rg e n t n e e d o f
                                              greater study. For example, it
         MARINE CONSERVATION IN               has been shown by Jordaens et
              THE AÇORES                      al. (2000) that there has been a
                                              loss of genetic variation in the
           The only places where ter-         strongly isolated population of
        restrially derived nutrients are      Tapes (=Venerupis) decussata in
        retained on steep, typically re-      the Fajã de Santo Cristo such that
        cently formed, volcanic, off-         it is highly vulnerable to extirpa-
        shore islands are lagoons and         tion.
        this makes such habitats extre-           In some locations on the
        mely important. By their very         Açores we can demonstrate that
        nature, the young Açorean is-         marine colonization has been not
        lands arise near vertically from      just fast, but dramatic, occurring
        the seabed and two lagoons oc-        within just a few years. On the
        cur on the steep (precipitous)        island of Terceira, there was a

Paginação_21.indd 60                                                                     19-01-2012 11:51:21
MORTON: DARWIN, THE ATLANTIC AND THE AÇORES                        61

                   FIGURE 13. A, The quarry at Cabo da Praia on Terceira in 1995 and B, in 2006. (Photo’s
                   courtesy of J.A. Paulus Bruno).

                   marsh at Paul, Praia da Vitória             now, 26 species identified. It is
                   (Morton et al., 1997).       This,          today considered to be the most
                   however, has now been drained.              significant coastal wader site in
                   There was, however, a coastal               the archipelago. The question is,
                   quarry at Cabo da Praia, also               however, since, the quarry was
                   on Terceira, and which when                 only constructed in ~1983, how
                   studied for the book ‘Coastal               did such a collection of species
                   Ecology of the Açores’ (Morton et           come to colonize it?
                   al., 1998) was found to contain
                   tidally inundated pools (Figure
                   13A). It was also shown to be
                   home to numerous species of
                   coastal plants, two species of
                   Assiminea (Gastropoda), the
                   amphipod Orchestia mediterranea
                   and, a near-unique (for the
                   Açores) collection of migrating
                   coastal birds including some
                   American vagrants, for instance,            FIGURE 14. The floor of the quarry at Cabo
                   the Hudsonian godwit (Limosa                da Praia on Terceira in March 2010 a er a
                   haemastica). Since first described          cleanup by the Environmental Division of
                   by Morton et al. (1997), growing            Bensaúde, one of the companies that has
                                                               a concession for the fuel storage facilities
                   numbers of bird species have
                                                               at Pedreira do Cabo da Praia, adjacent
                   been recorded from the quarry               to the wetland. (Photo: Marco Lopes,
                   at Cabo da Praia (Table 2) with,            Bensaúde).

Paginação_21.indd 61                                                                                19-01-2012 11:51:21
62                                                      AÇOREANA                                 2011, Suplemento 7: 39-71

        TABLE 2. Birds recorded from the quarry at Cabo da Praia, Terceira.
                                                      July 1997        October 1997                                      July/August
                                                                                         February 2006    March 2006
                          Species                   (Morton et al.,   (Morton et al.,                                         2007
                                                                                        (S. Rodebrand)   (B. Carlsson)
                                                        1998)             1998)                                          (B. Carlsson)
         Charadrius hiaticula (Ringed plover)                                                 3               1
         Charadrius dubius (Li le-ringed plover)        Pairs             20-30
                                                                                                                         (96-126, x =
         Charadrius alexandrinus (Kentish plover)   30 (Resident)           30               39               30         >112 in May
                                                                                                                          and June)
         Charadrius semipalmatus (Semipalmated
                                                                                              1                -
           plover)
                                                       1 (First         6 (Winter
         Pluvialis squatarola (Grey plover)
                                                      summer)           plumage)
         Calidris canutus (Red knot)                                        7                 3               3
         Calidris alba (Sanderling)                       1                >100              41               60
         Calidris minuta (Li le stint)                                                        3               3
         Calidris minutilla (Least sandpiper)                                                 1                -
         Calidris fuscicollis (White-rumped
                                                                                              1                -
           sandpiper)
         Calidris ferruginea (Curlew sandpiper)                                               2               2
                                                                        5 (Winter
         Calidris alpina (Dunlin)                                                             4
                                                                        plumage)
         Tringa ochropus (Green sandpiper)                1
                                                     3 (breeding
         Limosa limosa (Black-tailed godwit)                                                  1               1
                                                      plumage)
         Limosa lapponica (Bar-tailed godwit)                               8                                 1
         Limosa haemastica (Hudsonian godwit)                                                                                  1
         Numenius phaeopus (Whimbrel)                     2                 1                 6               4
                                                     12 (Adults +
         Arenaria interpres (Turnstone)                                     20               29               50         (29-68, x = 54)
                                                    first summer)
         Philomachus pugnax (Ruff )                                                            1
         Sterna hirundo (Common tern)                  Present              40
         Sterna dougallii (Roseate tern                Present              2
         Larus cachinnans (Yellow-legged gull)                           Present
                                                                        3 (Winter
         Larus ridibundus (Black-headed gull)
                                                                        plumage)
         Ardea cinerea (Grey heron)                       3                 4
         Egre a garze a (Li le egret)                     1                 2
         Anas platyrhynchos (Mallard)                                   2 females

           Charles Darwin also pondered                                            earth adhering to it. … the earth
        such a question and in the The                                             had been kept for three years, but
        Origins of Species by means of                                             when broken, watered and placed
        Natural Selection, he considered                                           under a bell glass, no less than
        that birds might be responsible                                            82 plants sprung from it…’ and
        for such rapid colonizations. He                                           ‘can we doubt that the many birds
        reported that:                                                             which are annually blown by gales
              ‘..the leg of a red-legged partridge                                 across great spaces of ocean and
           (Caccabis rufa) had a ball of hard                                      which annually migrate….must

Paginação_21.indd 62                                                                                                                       19-01-2012 11:51:22
MORTON: DARWIN, THE ATLANTIC AND THE AÇORES                 63

                      occasionally transport a few seeds   upon whose shoulders Darwin
                      embedded in dirt adhering to their   stood. John Stevens Henslow
                      feet or beaks’ (Darwin, 1878).       was Darwin’s early botanical
                      Regrettably, along with the          mentor. The botanist Joseph
                   destruction of the marsh at             Dalton Hooker inspired Darwin
                   Paul, the entrance to the unique,       to study biogeography and how
                   man-created, quarry habitat at          plants are transported across
                   Cabo da Praia, has been largely         the oceans. Darwin’s greatest
                   developed (Figure 13B). But             mentor, the geologist Charles
                   what a wonderful project it             Lyell, inspired him to think first
                   would have constituted to test          about geology (on Cape Verde)
                   Darwin’s theories of how coastal        and then to marry his thoughts
                   plants and maybe animals could          on this subject with his others
                   have colonized Açorean shores           on biology to come up with the
                   – both by transport across the          great unifying theory of the evo-
                   sea with ocean currents and by          lution of life on Earth, but also
                   birds.                                  the evolution of the Earth itself.
                      A postscript to this develop-        And, eventually, to discover how
                   ment, however, is that one of the       island floras and faunas are ob-
                   companies which has a conces-           tained and, themselves, evolve
                   sion for the fuel storage facilities    in wonderful isolation. Robert
                   at Pedreira do Cabo da Praia on         FitzRoy, Captain of H.M.S.
                   Terceira, adjacent to the quarry,
                   has, in March 2010, undertaken
                   a cleanup of the floor of Cabo
                   da Praia (Figure 14), perhaps
                   signaling a change in environ-
                   mental awareness in relation to
                   this highly important Açorean
                   wetland.

                      Many great men both taught
                   and influenced Charles Darwin,
                   as a young, 22 year-old, adven-         FIGURE 15. The eruption of Sabrina
                   turer on H.M.S. Beagle. These           Island on 19 June 1811, as drawn by
                   included a number of distin-            Lieutenant John William Miles of H.M.S.
                   guished 19th century scholars           Sabrina. (A er Tillard, 1812).

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64                           AÇOREANA               2011, Suplemento 7: 39-71

        Beagle, became the foil against      third son so that, in many ways,
        whom, as his companion natu-         the cultural and, especially, the
        ralist, Darwin debated the ideas     maritime histories of Portugal
        that were slowly fomenting in        and England are inextricably
        his mind. FitzRoy eventually         linked, even today. I had the
        became a Vice-Admiral, famous        privilege of meeting Colonel
        in his own right for creating the    Agostinho in 1965 in Angra do
        modern science of weather fore-      Heroísmo.
        casting. As a devoted Christian,        Although the Açorean islands
        however, he would eventually         were of little interest to Darwin
        divorce himself from Darwin’s        when he visited them in 1836,
        heresy and, as he saw it, unjus-     his views changed subsequently.
        tly awarded fame, and commit-        Firstly, his attention was drawn
        ted suicide by cutting his own       to the islet of Sabrina that arose
        throat on 30 April 1865.             from the sea during June and
           Interestingly, an eminent         July 1811 (Tillard, 1812) just
        Açorean, Colonel José Agostinho      opposite Ponta da Ferraria
        ( 1888- 1978), bec ame c h ief       on São Miguel (Figure 15).
        scientist with the Portuguese        Coincidentally, H.M.S. Sabrina
        National Weather Service and,        was in the vicinity and onto the
        for providing the British with       shores of the newly formed islet
        meteorological and sea-state         stepped Captain James Tillard to
        data during the Second World         claim it for Great Britain. This
        War, was made an Honorary            caused a diplomatic rift until the
        Commander in the Order of the        islet disappeared back into the
        British Empire by King George        sea from which it had arisen in
        VI. This act and its recognition     August 1811. Nevertheless, for
        is in remarkable recognition of      Darwin, this was incontrovertible
        the Anglo-Portuguese Alliance        proof that volcanic islands
        of 1373. This came about when        do arise and may sink over
        John I (1358-1433), the tenth King   the course of their lifetimes,
        of Portugal and the Algarve,         albeit with great differences in
        now famously, married, in 1387,      rates. A similar event occurred
        Philippa of Lancaster, daughter      in the 20th century when the
        of John of Gaunt (1340-1399)         Capelinhos volcano erupted
        of England. Prince Henry the         on the west coast of Faial in
        Navigator (1394-1460) was their      1957. This eruption was studied

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MORTON: DARWIN, THE ATLANTIC AND THE AÇORES                 65

                   by the Açorean vulcanologist           However, more interestingly,
                   Frederico de Menezes Avelino           in response to a request from
                   Machado (1918-2000) and which          Furtado for research advice, in
                   when I visited it in 1965 was still    a reply letter dated 3 July 1881,
                   volcanically active. Machado           Darwin suggested to him that:
                   was assisted in this research by       (1), If possible, the most distant
                   Victor Hugo Lecoq de Lacerda           outlying islands and their plants
                   Forjaz (born 1940), who became         and animals should be com-
                   Professor of Vulcanology at            pared with those of the other is-
                   the University of the Açores           lands; (2), all the plants and ani-
                   and is today Director of the           mals from the highest mountain
                   Observatório Vulcanológico e           summits on all the islands ought to
                   Geotérmico dos Açores.                 be collected; (3), searches made
                      Secondly, and more impor-           for glacial deposits and for the
                   tantly, however, Darwin en-            presence of fossil remains, in
                   tered into correspondence with         such stones and (4), any light-
                   Francisco d’Arruda Furtado             -houses should be inspected for
                   (1854-1887), a native Açorean,         any land-birds that might have
                   much interested in natural his-        flown into the glass and killed.
                   tory, a devotee of Darwin, and         In such cases, their feet and beaks
                   a believer in evolution through        should be examined not only for
                   natural selection. Darwin was          earth, but the whole contents of
                   perplexed as to how the lizard         their alimentary canals dried out
                   (Lacerta dugesi), earthworms and       and placed on damp pure sand
                   snails had arrived in the Açores.      under a small bell-glass to see
                   He asked Furtado specifically          if any seeds are present which
                   to determine if the lizard’s eggs      would germinate. If so, grow
                   could survive in seawater. The         any plants and name them.
                   truth is probably much more               Thirdly, as described above,
                   prosaic, however: all could have       Darwin knew that seeds from the
                   arrived, except for the endemic        New World occurred regularly
                   leptaxine snails (van Riel et al.,     on Açorean beaches and he used
                   2005) via human interventions.         this fact as the raison d’etre for his
                   Indeed, Chaves (1949), provides        experiments on the survival of
                   an introduction date of 1860 for       seeds and other plant pieces in
                   the Macaronesian lizard although       seawater. Thus, when asking his
                   this seems highly improbable.          friend Joseph Hooker for seeds

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66                                 AÇOREANA                   2011, Suplemento 7: 39-71

        to be collected from Hitcham in             ding of the forces that created
        Suffolk, England, he specifically           the archipelago and how they
        requested that they include                 became colonized show us what
        those from plants also known to             is important with regard to mari-
        grow in the Açores.                         ne conservation in the Açores.
            Hence, although Darwin’s vi-                The predominant rocky shores,
        sit to the Açores was not initially         washed by oceanic currents are
        inspiring, the islands subsequen-           probably going to provide us
        tly became so for him. Similarly,           with only a few, if any, examples
        he laid down the initial basis for          of endemic species. There may
        our understanding of how the                similarly be few endemic spe-
        Açorean flora and fauna was                 cies making up the communities
        obtained. Finally, however, and             of other intertidal habitats. In
        although the word conservation              one or two places in the Açores,
        was not in Darwin’s vocabulary              however, there are intertidal,
        (in the natural history sense), his         lagoonal, wetlands. In particu-
        researches upon and understan-              lar, Fajã dos Cubres and Fajã de

        FIGURE 16. An illustration showing the origins of mankind from Paradise – Lemuria
        – located in the Western Indian Ocean. The frontispiece to Ernst Haeckel’s History of
        Creation (1876).

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MORTON: DARWIN, THE ATLANTIC AND THE AÇORES              67

                   Santo Cristo, both on São Jorge,       nued to argue that human evo-
                   the marsh at Lajes on the island       lution consisted of exactly 22
                   of Pico and, especially, the quar-     phases, the 21st (back in time),
                   ry at Cabo da Praia on Terceira,       being the ‘missing link’ and re-
                   have much to reveal about how          presenting a halfway step betwe-
                   such isolated communities have         en apes and humans. But, that
                   become established – a question        our origins were still in Paradise
                   that Darwin was deeply concer-         (Figure 16). Clearly, even this
                   ned with, but who never saw            enlightened contemporary of
                   these little habitats in the Açores.   Darwin, still felt the need to ack-
                   In particular, the modern quarry       nowledge the Christian orthodox
                   at Cabo da Praia, on Terceira,         view of Genesis. But the illustra-
                   because it is isolated from sea,       tion represents more, because its
                   except via the surrounding ba-         image of human evolution still
                   saltic rock wall filter could tell     resonates today in the minds of
                   us much about the origins of the       some, generations later, and high-
                   Açorean biota and, more interes-       lights the continuing struggle
                   tingly, how its representatives        between science and faith for the
                   have obtained purchase on the          soul of humanity and haunts the
                   islands.                               progress of humankind towards
                       In his understanding of the        a more tolerant world.
                   evolution of life, Darwin also             In the context of island
                   discovered important facts con-        evolution, however, and espe-
                   cerned with the evolution of           cially in the comparatively new
                   the Earth, thereby achieving the       geological context of the Açores,
                   marrying of biology and geo-           the researches of Charles Darwin
                   logy, that is, the unification of      remain unchallenged and are
                   the history of life on Earth and,      critical to our understanding
                   finally, the rejection of the bi-      of island biogeography. And it
                   blical account of creation. It         is his seminal discoveries that
                   i s i nteres tin g th ough , th a t    have, in turn, led to a better
                   despite meeting, believing in          understanding of the need for
                   evolution through natural se-          conservation not only on land,
                   lection and supporting Darwin,         but also on the riparian edge
                   Ernst Haeckel (1834-1919), the         of the sea, especially on island
                   great German anatomist, em-            archipelagos where new found
                   bryologist and illustrator, conti-     life clings precariously.

Paginação_21.indd 67                                                                   19-01-2012 11:51:24
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