Palabras clave antrópicos. que impactos naturales procesos más respondan observados biodiversidad de que de patrones los incertidumbre la considerar para hookeriana, la carencia, nueva proponemos una trabajo este En biodiversidad. de estudios los que impactan conocimiento, de o faltas carencias, no menos ocho de postulado han se momento el Hasta natural». «laboratorio como comprometer utilidad su humano impacto el puede y cómo Cuándo humano. impacto el concretamente, insular, biodiversidad la de patrón del configuración la elemento en otro de clave existencia la subrayó Hooker también de conferencia yevolución. La ecología de estudio el para sistémicos modelos como estas que contempla natural», laboratorio de «paradigma como islas las de concepto del desarrollo que al contribuyó decirse puede evolutivos». tal, Como estudios para islas las de portancia im la de sistemática declaración primera «la como sido catalogada ha conferencia Esta Floras insulares titulada Nottingham, en Ciencia la de Avance el para Británica Sociedad Hooker (1817-1911) Joseph Dalton 1866En botánico el ala conferencia una impartió Resumen herbarium. biogeography, impact, evolution,: Island human Keywords discussed. is shortfall this to address help collections history natural for potential The processes. anthropogenic than rather natural reflect patterns biodiversity to which extent the in for uncertainty account to shortfall– Hookerian order –a in is shortfall a further that paper, we suggest this In described. been have of biodiversity– on studies impact that gaps –knowledge “shortfalls” eight evolution? To than for studying no fewer laboratories’ date, ‘natural as of islands lity uti the compromise impact human does when How and impact. human namely patterns, diversity island shaping in driver key another highlighted evolution. Hooker’s also lecture and ecology for studying systems model as islands views that paradigm” laboratory “natural development to of the contributed the to have seen be it such, can studies”. As evolutionary for islands of importance the statement of systematic “the first as described been has lecture Insular floras U.K. entitled Nottingham, in of Science Advancement for the Association Hooker (1817-1911) Joseph Dalton botanist 1866,In the British to the alecture delivered Abstract INSULAR FLORAS ON JOSEPH INSULAR SIR HOOKER INSULAR FLORAS INSULAR CONFERENCIA LA AND THE NATURAL LABORATORY PARADIGM LABORATORY NATURAL THE AND Revista Scientia Insularum, 3; diciembre 2020, pp. 73-88; ISSN: e-2659-6644 ISSN: pp. 73-88; 2020, diciembre 3; Insularum, Scientia Revista Mark Carine* & Miguel Menezes de Menezes Sequeira** &Miguel Carine* Mark biogeografía insular, evolución, impacto humano, herbario. humano, evolución, impacto insular, : biogeografía Y EL PARADIGMA DEL LABORATORIO NATURAL LABORATORIO DEL PARADIGMA Y EL DOI: https://doi.org/10.25145/j.SI.2020.03.05 DOI: HUMANO IMPACTO EL HOOKER: JOSEPH , SIR DE : HUMAN IMPACT : HUMAN . The . The - - .
REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 73 REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 74 and Evolutionary Biology, CIBIO-Azores, Portugal. Biology, CIBIO-Azores, Evolutionary and Biodiversity in Network Research InBio, 9020-105 Funchal-Portugal; Penteada, da Universitário Museum, Cromwell Road, London, SW7 5BD, United Kingdom. Email: [email protected] Email: 5BD, Kingdom. United SW7 London, Road, Cromwell Museum, Insular Floras Insular Hooker’s(2009), reviewing in contribution biology, to island considered 1866 the (1984).1896, 2). by Williamson Figure acommentary reprinted It with was Berry (Hooker, apamphlet Gardeners as the Chronicle in its entirety later in and parts in subsequently people was published (Berry, 2009).of 2000 the lecture of The text Lecture on U.K. His for of Advancement Nottingham, the Science in Association British to the London, in U.K., Kew delivered alecture Gardens, Botanic Director of Royal the Figure 1. A photograph of Sir Joseph Dalton Hooker (1817-1911), Hooker Dalton Joseph of Sir 1. Aphotograph Figure after years 1868, two in taken lecture. © The Trustees of the Natural History Museum, London. Museum, History the Natural of ©The Trustees lecture. Floras Insular his ** On August 27 August On lgae, Fungi and Plants Division, Department of Life Sciences, The Natural History History Natural The Sciences, of Life Department Division, Plants and Fungi lgae, *
A Insular Floras Insular (hereafter Floras Insular M adeira Botanical Group, Faculty of Life Sciences, University of Madeira, Campus Campus of Madeira, University Sciences, of Life Group, Faculty Botanical adeira lecture to be particularly significant describing it as a describing it in significant ’...landmark to particularly be lecture th , 1866, Joseph Dalton Hooker (1817-1911; 1), Figure then 1. INTRODUCTION )
was given to an estimated audience audience estimated given to an was . 1.1. Hooker islands 1.1. and be reconciled.can condition on other– the disturbed evolution highly ying their on one and the hand laboratories’ for as stud role –their ‘natural discussed he floras of island aspects two of Hooker’s overview we paper an give before how considering these lecture this In its significance. and on islands impact considered of extent the human also and studies.’ for evolutionary of islands importance of the ment Origin the for support scientific personal gift of the British sovereign. Hooker maintained an extensive and interna and extensive an sovereign. of British Hooker the maintained gift personal amember made of Order the the in of was award Merit, and work India an his in dom’s premieracademy, scientific from 1873 to for knighthood 1878. a He received of London, President United Society the as King of Royal He the served years. Kew, he apost held Gardens, for 20 Botanic second Director of Royal the the as Hooker Jackson William father his succeeded he had lecture Floras Insular vered his Lecture on Insular Floras Insular on of Hooker’s Lecture page Title 2. Figure booklet by L. Reeve & Co. © The Trustees of the Natural History Museum, London. Museum, History the Natural of ©The &Co. Trustees Reeve byL. booklet Hooker was an influential figure infigure 19 influential an Hooker was evolution and Hooker’s relationships floras the of island addressed lecture ... 1859]’ [i.e Darwin, ‘...the firstsystematic and state th century botany. he By century deli time the , published in its entirety as a 36 page page a36 as entirety its in , published - - - - -
REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 75 REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 76 Hooker and Darwin ( Hooker Darwin and for example, correspondence between, evident from surviving is archipelago the but interest in his Azores except on all Hooker botanized had Islands. Kerguelen St sub-Antarctic Helena the and and of Ascension islands south Atlantic the des, Ver Cape the and Islands Canary Madeira, of Azores, the archipelagos ronesian theMaca groups. discussed island He or islands oceanic seven of floras and ristics floras insular Explaining 1.2. men. two the between long friendship alife established aproject It that was archipelago. fromtions the on Darwin’s collec based was and Darwin invitation of at the made Charles was example, 1847) for (Hooker, Archipelago Galapagos of the Plants of the Enumeration his visited thethose beyondhad extended voyage; on he floras island in interests Tasmania (Hooker, and islands 1844-1860) Antarctic the but Zealand, New his outcomes of Erebus the botanical up the ting wri In plants. to seed from algae everything extensively; to collect opportunity islands). the He Campbell used and Cockburn Kerguelen, Crozet, (Auckland, tic Antarc and sub-Antarctic the and Islands) Falkland Helena, the Saint Ascension, Verdes), Cape the (the Trindade, Atlantic South Pedro São Paulo the São and rocks, in Santiago and Islands Tenerife (Madeira, Canary Macaronesia the in in islands Pole. South magnetic to the Thetook expedition voyage Hooker1843 to Antarctic on 1839- the Ross Clarke James under Navy, Erebus onRoyal HMS Hooker served the British in Surgeon Assistant As a young floras. island of oceanic experience (Endersby,them &b). 2008a explaining but of diversity also patterns concerned not documenting only with was and botanist’ to a‘philosophical be Hooker considered himself Darwin. Charles friend 1883). developed by his theory of evolutionary advocate the an also He was tish India editor seven-volume the of- monumental including works of taxonomic Bri Flora neobotany,and theexplorerandandand author collector aprolific Hooker was org/p/jdh of network http://jdhooker.kew. correspondents (Endersby,tional also see 2008a; in insular floras ‘ floras insular in similarities analogous analogy.highlighted those He of and affinity those of floras: ‘in all cases the flora is quite manifestly closely allied to some some to allied closely manifestly quite is flora the cases all He floras. noted ‘in insular arboreous’ or shrubby are that species allied by represented are or on islets, shrubby ‘ that writing ness, the andphenomenon woodi floras of island insular of form spectrum growth tive distinc the example, for etc...’ continents, from distance and noting, area limited sure, Hooker’s Hooker’s Hooker’s Hooker went on to distinguish two types of relationship of these insular insular of of relationship these types two Hooker went on to distinguish His ‘relationships of affinity’ concerned therelationships of concerned biogeographic ‘relationshipsHis of affinity’ (Hooker, 1872-1897) Genera and Plantarum ) and his botanical interests were wide ranging. Eminent in both palaeo- palaeo- both Eminent in were ranging. wide interests botanical his ) and due to physical conditions common to them all –to their climate, expo climate, their –to all them to common conditions physical to due lecture provided an outline of the physical characte of physical outline provided the an lecture floras Insular lecture reflected his particular interest and wideand interest particular his reflected lecture Floras Insular Plants which are herbs on continents, often either themselves become become themselves either often on continents, herbs are which Plants https://www.darwinproject.ac.uk/ voyage, Hooker produced Floras of of Floras produced Hooker voyage, (Bentham &Hooker, (Bentham 1862- ). . ------and evolution. He also addressed the impact of humans on island floras. on island of humans impact the evolution.and addressed He also impact on human Hooker 1.3. 2017, 2017). al., et Whittaker 2015; biology al., (Warren in et concepts of key 2016; al., et Santos Patiño al., et continue to play amajor islands role development that the in it clear and is 2007) Fernández-Palacios, and of evolution suited for in-situ studies (Whittaker ideally laboratories’; ‘natural as of systems value islands the role highlighting in important today. played an studies lecture evolutionary His biogeographic and to island tral 2015). al., et (e.g.tion processes Mairal - extinc of continental result the are that palaeoendemics also and recent dispersal from (relatively) resulting both neo-endemics, contain floras their complex that and been has ‘stocking’ their that islands of Macaronesian the case the evident in less is it neverthe those Macaronesia, of as such floras island of oceanic origins the lain no longer are considered bridges to a plausible exp land hypothesis Whilst of each. support in evidence the assessed carefully and hypotheses bridge land and dispersal continent’. from the plants with both island He oceanic discussed of an ‘stocking prolific...’less or hardy less the displacing prolific more or hardy more the in terminate must which for existence struggle that It accelerates 2. plant. habits of the the to of suitable spots 1. number the ways. It in reduces various operates island of the sinking This experienced. all have they subsidence the is islands on oceanic species of old ‘old species’,cular extinction or rarity of the he cause aprincipal noted that ‘I believe on parti of in levels diversity. 2007) al., et Discussing (Whittaker cycle life island 2007). al., et poorten Vander see situation more is the (although complex cryptogams; the stand in still Hooker’s number and in few are but on flowering flora the plant they observations represented, Lactuca in for example are taxa sister American comm.). with pers. TUM, Endemic taxa (H. Schaefer, its affinities in European fact in is azorica Sanicula that andindeed suggests flora plantseed the of endemic affinity European thelargely confirmed worknetic has phyloge Recent Sanicula’. genus Umbelliferous of the in aspecies is can substantiate, West, Hooker noted ‘ from the winds but prevailing with from America 2400 km from and Europe 2004). et al., plant floras seed (e.g.Carine island of oceanic phylogenetic analyses temporary conin reflected apattern continent’, other of such vegetation of the faint traces than more presents it never continent, another to approximates much so it by however and continent, mother the from be it may distant however that and Flora, continental one Hooker’s to apresentation biogeography not of island restricted was lecture cen remain lecture his in Hooker addressed of that questions the Many for to account the of hypotheses adiscussion with He concluded essay his consideredthebe impact thewhat would of be Hookertoday to on reflected Writing on the Azores in this context, an archipelago located 1700 km 1700 km located archipelago an context, this in Writing on Azores the The only trace of American influence on the Azorean Flora that I that Flora Azorean the on influence of American trace only The (Dias et al., 2018) al., et (Dias Solidago and (Schaefer, 2015), ------
REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 77 REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 78 Banks and Daniel Solander visited the island in 1771 in island Cook’s first the visited James Solander during Daniel and Banks 3). (Figure of Nesiotawas elliptica Theextinction recentmost Joseph Roxb. When wild the in extinct or extinct consideredglobally endemic flowering species plant world of 35 the the one in with quarter most threatened recognised the among is undoubtedly, St Helenathe florahas, impacted by been humans. hugely Today, it over-estimate but an be may Hooker’s extinction of species of scale the estimate (Cronk, colonisation of and island 2000). the discovery the following vegetation native to the changes brought forof about timber forests rapid catastrophic and theharvesting and introductionThe herbivores andinvasive species large plant of mid-1600s from the Company India East for onwards. British the major of port call a became and colonised was location, island the important of itsresult strategically 1500s a early of the asource meat as in were and, as century. introduced Goats sixteenth before the island.’ of on goats the uninhabited St Helenaintroduction was first since the Naturae Systema the from disappeared thus have plants 100 Helena St collected by Joseph Banks and Daniel Daniel and Banks byJoseph of Trochetiopsisspecimen collected type melanoxylon 3. The Figure is Extinct in in T. Extinct is St Helena: to of Trochetiopsis endemic erythroxylon species two other are There Solander on St Helena in 1771. The species is endemic to St Helena. It is globally extinct and extinct globally is It to St Helena. endemic 1771.is in on St Helena species The Solander is Critically Endangered. Image from the Natural History Museum, Museum, History Natural the from Image Endangered. Critically T. is and ebenus Wild the the Banks and Solander type collection made in 1771 was the last collection of the species. species. of the collection last 1771 the in made was collection type Solander and Banks the ‘Probably Writing he ‘Probably commented on that St south Atlantic Helena the in London data portal (data.nhm.ac.uk), , used under a CC-BY 4.0 licence. licence. 4.0 aCC-BY under ,used (data.nhm.ac.uk), portal data London © The Trustees of the Natural History Museum, London. Museum, History the Natural of © The Trustees the 15 the from to produce for sugar fuel forest cutting and plantations cane sugar of massive Frutuoso,tuoso; see 1998). Moore (2009, 2010) development the that estimated those made today if the history of the forests is not taken into account. not is taken of forests the history the if today made those to different 1800s very have may been early the in would made have been tion that its composition evolu and regarding ferent of avisitor inferences to that and today Erebus the visit aboard his during of laurisilva the view Hooker’s recent relatively past. the in it was to much be more than likely extensive thus is today laurisilva century. Madeiran 20th the in took mostly place that cess 2018).thereforeand recentlate a pro was Madeira in thelaurisilva of The recovery to about back 150 dating (Patiño al., et ago years harvesting ofshow historical signs 4). or (Figure houses to villages linked were usually occur, they did seldom where could found be they and, trees where tall grazed heavily and culture for agri used largely 2010) al., et landscape (Pupo-Correia abarren reveal images Nineteenth century for intensively grazing. used landscapes and plants cultivated of on distribution the based were largely that propose for belts vegetation Madeira Humboldt (1825), Bowdish led authors as such (1826) Khul (1857) Lowe and to 2010). al., et de Sequeira Sloane (Menezes von Alexander Later, of influence the tems in the in tems of forest ecosys rapid document the destruction also reports 2007). Early Dias, (Frutuoso, probably 1998; are and extinct unknown region are but that others present the in some comprising still forests species refer to large Azores) the also (and descriptions of Madeira Early data. historical consistent with it certainly is forest (e.g. http://whc.unesco.org/en/list/934 primary largely are laurisilva ran gical Chronometer’. Geolo the by of can measured onlybe Nature operations ordinary the under as years, 400 last the within a revolution such undergone has island lovely of that history ral natu The overlooked. be not must of man agency He the commented in Madeira ‘But of loss. biodiversity scale true the understand to we are if extinction 2017al., census than rather of context the Macaronesia) in debt Triantis (see for 2010, example, al., et also, extinction on et estimating Otto to need focus the event to highlight and extinction-causing an following to occur Nesiota 2016). Ellick, and (Lambdon 2003 in Cronk (2016) infections the used to fungal succumbed last the until survived individual from this propagated acutting and in died wild plant 1994 last The butseedlings much stature. shorter in all remained, later only 12-15 acentury up to 9 m were there high; trees circumnavigation, trees can also be found in the writings of 17 writings the found be in also can the early 16 early the by destroyed of laurisilva area of total 40% the and one quarter ponds to between That corres close to 25,000 ha. would figure put that estimate optimistic a less th century onwards resulted in close to cut close by of 16,000 ha 1509; in forest being resulted onwards century example to illustrate the sometimes long period of time taken for extinction for extinction long taken of sometimes period the time to illustrate example Recent research has revealed that even the oldest forest stands in Madeira Madeira in oldest forest stands the even that revealed has research Recent This statement is at odds with the idea that the extensive areas Madei of areas extensive the that the idea with statement is odds at This flora. on Madeiran the attention of humans impact to drew the Hooker also Evidence for the lack of forest or for a landscape converted to agriculture converted to agriculture Evidence of forest or for lack for the alandscape
th fifteenth Fru (e.g.fifteenth centuries (Gaspar Fernandes, and1940) sixteenth century. th and 18 and th century authors such as Hans Hans authors as such century would have been very dif would very have been ) but but ) ------
REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 79 REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 80 difficult to separate. to difficult extremely be actually may human-mediated is what and sequently, natural is what Con changes. to responds those vegetation the and time through change Pressures complex.is plants island on activity of human but biodiversity effects of the island have aboutbrought major a aprofound floras. Humans had onloss island impact clearings in a complex mosaic of forest and scrub. (b) Photograph taken in 2007 showing recovery recovery showing 2007 in taken (b) Photograph scrub. and of forest mosaic acomplex in clearings of native vegetation in areas that were not covered in (a). The transition is to high-scrub or young young or high-scrub to is (a). in transition not covered were The that areas in vegetation of native vegetation through time. (a) and (b): Ribeira da Janela Valley. (a) Photograph taken by unknown byunknown Valley. (a) taken Janela da Photograph (a) (b): Ribeira and time. through vegetation in less accessible areas. The trees are exotic species (e.g. Sycamore ( (e.g. Sycamore species exotic are trees The areas. accessible less in photographer between 1876 and 1878 showing the lack of native forest; some native vegetation, vegetation, native some forest; of native 1876 lack the 1878 and between showing photographer photographer in the early twenty century showing large areas without vegetation cover forming forming cover vegetation without areas large showing century twenty early the in photographer mainly heather ( heather mainly ( forest with early secondary, shade intolerant species such as Myrica faya as such species intolerant shade secondary, early with forest (c) and (d): Old Curral dos Romeiros or Curral pequeno. (c) Photograph taken by unknown byunknown (c) pequeno. taken Photograph or Curral Romeiros dos (c) (d): and Old Curral Pinus pinaster Pinus Figure 4. Repeat photography of Madeiran landscapes reveals the different fates of native native of fates different the reveals landscapes of Madeiran photography Repeat 4. Figure unlike in (b), it is exclusively by exotic trees: Tasmanian Blue Gum ( Eucalyptus globulus Blue Gum Tasmanian trees: byexotic (b), in exclusively it is unlike Black wattle ( wattle Black As the examples of St Helena and Madeira serve to highlight, humans have humans to highlight, of serve St Helena Madeira examples and the As )). (d). Photograph taken in 2011 showing the increase of tree cover.However, of tree )). 2011 increase in the (d). showing taken Photograph subsp platycodon Erica Acacia mearnsii Images courtesy of Aida Pupo-Correia. of Aida courtesy Images ) and Sweet Pittosporum ( Pittosporum Sweet ) and . maderincola ), is scattered in the landscape particularly particularly landscape the in ), scattered is Pittosporum undulatum Pittosporum Piatanus xhispanica Clethra arborea Clethra and ) and pine pine ) and ).
),
. - our interpretation of those patterns. As Whittaker and Fernández-Palacios (2007) (2007) Fernández-Palacios and Whittaker As interpretationour patterns. of those how and it impact may scales evolutionary across on today islands observed terns pat diversity the shaped has on islands impact human how past highlight examples These impact. human pre-European and of European aresult as on Zealand New (2018)kakapo the in diversity geneticloss of attention significant to the have drawn reductionbrought its morphological diversity. in about amarked Seerholm al. et (2017) Anolis in showed ferreus how extinction human-mediated Anolis Caribbean in patterns diversity species modified (2014) al. Helmus et archipelagos. introductions have how demonstrated human between comparisons biogeographical to make patterns diversity of contemporary use the 2018). al. et (Boeiro challenges arthropods certainly However, history this for observed been has as Azores, the in than Madeira in of level conservation higher a in resulting areas, inaccessible and small in forest survived not pristine, although forest Conceivably, patches. of all a dynamic mosaic of mature, neous disappearance but have may prevented orographic complexity Madeira, simulta forest in the tine any evidence Thereis of pris no of island. orographic to the complexity the linked post-human be may colonization Madeira in forest persistence have proposed that questions? of evolutionary entire spectrum the across assumption areasonable this is on islands, impact Given of breadth human interpretation the patterns. of those the for not significant of humans impact the with processes of natural result the are to explain seek they that document and they that of diversity patterns the that assume lopment generally biogeographic theory, and of studies such evolutionary the deve in systems as role model have played asignificant islands Whilst tems? loci control (e.g. 2016; al., et Almén 2015). Paun al., et those and what of loci of differences, genetic involved the speciation, nature in the number the to determine of genomic to speciation events investigated, be the basis (e.g. were intractable previously that 2015) al., Mort et lineages allowing but also is island to relationships elucidated evolving rapidly be in evolutionary only allowing (Sousa & Hey 2013; of 2014). lineages to history Ellegren uncover the ties It not is unprecedented for providing opportuni is non-model and rate, creasing organisms, ever-inthe Theacquisitionevents. genomican at of data sequenceat DNA level diversification of island basis genetic to the relating questions fundamental pass to encom studies evolutionary of scope island the extend to greatly potential the Generation Sequencingnext (NGS) approaches and have of speciation processes ses analy to fine-scale or biogeographic analyses macro-evolutionary from broad-scale lost.’ irrecoverably is evidence which but extinct, and living both species, other of affinities of the evidence [extinct species] contextthe In of St floras. Helenacommented ‘ that he island paradigm laboratory natural the and impact Human 1.4. Jardim & Menezes de Sequeira (2008) and Menezes de Sequeira et al. (2013) al. et de Sequeira (2008) Menezes de Sequeira and &Menezes Jardim sys island oceanic in inferences to evolutionary our limits But there are range on islands biogeographic analyses and evolutionary Contemporary understanding for impact ofhuman theHooker significance on reflected was a link in the chain of created beings, which contained within itself itself within contained which beings, chain of alink in created the was lizards whilst Bochaton et al. Bochaton whilst lizards in Guadeloupe has has Guadeloupe in Every one of those of one those Every ------
REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 81 REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 82 the ‘Hookerian shortfall’. The world’s herbaria contain an estimated estimated million 350an The world’s ‘Hookerian shortfall’. the contain herbaria by resolution. poor taxonomic limited often are they although informative be to it who also may were by those witnesses tten wri impact of testimony human direct bear that accounts Historical shortfall’. rian ‘Hooke the address and studies evolutionary help island could to inform baselines Paleoecological baselines. establishing in studies of palaeoecological importance the it. articulate to problem arguably, first the of he this was, and aware from his clear is As ‘Hookerian shortfall’. med the ter be might that ashortfall processes– natural than rather by anthropogenic ped sha of are biodiversity patterns of to extent which the knowledge our in shortfall of biodiversity the unexplored environments. (2018) of of for knowledge impediment lack our proposed Racovitzan recently the and fitness). al. et Ficetola on survival effects their and interactions of species dge to of abiotic species tolerances of conditions) knowle and (lack Eltonian the and about responses of the knowledge functions), (lack their Hutchinsonian and traits of species of knowledge (lack Raunkierian the shortfalls: further addition three in (2015) proposed and al. et Hortal of concept life. the of shortfalls extended further of tree the of for to knowledge account lack (2013) shortfall proposed a Darwinian Diniz-Filho whilst dynamics population and abundance species of knowledge in (2011) al. et Cardoso for deficiency to account the shortfall proposed aPrestonian contextthebiology. conservation in of wereproposed first distributions– incomplete of taxon –the knowledge (Lomolino, 2004) shortfall cean the Walla with together shortfall, Linnean The Azores. florathe the in of diversity of patterns explaining and understanding in of diversity– taxonomic knowledge Lomolino,and (Brown incomplete1998)shortfall –the Linnean of a significance (2011) by a number al. et of authors. Forcussed Schaefer example, the discussed shortfall? hookerian A 1.5. evolution? laboratories’ for ‘natural studying as of islands use simply are not to the possible? alimit there Is processes evolutionary and ecological natural, concerning inferences that robust areso significant effects anthropogenic point belowthere which critical a Is we theobserve. patterns explaining in significant become increasingly processes natural ciation genomics), than rather anthropogenic (e.g. biota) of biogeographic relationships island to the finer-scale questions (e.g. spe tapestries. merly different rather of for fragments more the just resistant are that patterns evolutionary terpreting of misin adanger is there of extinction, impact the noted, without understanding Herbarium collections provide another source of data to help understand to provide help of source data understand collections another Herbarium (2017) al. et Nogue conservation, of context the island In have highlighted here – discussed shortfall the is list Conspicuous by from this its absence have now identified. been knowledge biodiversity in shortfalls Further dis have been floras of island of diversity the knowledge our in Shortfalls moves from broad-scale of studies whether, focus is the not is clear as What lecture, Hooker was Hooker was lecture, floras Insular
t he ------tfall in island biogeography. island in tfall shor the of Hookerian significance the help and to establish time through patterns of biodiversity to robustness help the used test be can provide that of asource data they doing, so In time. at aparticular place present at aparticular addressed) being or (depending scale genes on evolutionary or the genotypes or lineages species the However, time. and do space provide on both of in they asource data biased heavily evolution the of flora. the regarding 3) on (Figure inferences our impact chetiopsis Tro as such or endemic lineages species how within of loss the diversity investigate floraperm from to used (Cronk,2000).St be could specimens Helena Herbarium angios the in of extinction pace 1986) and have providedfor scale and the evidence (Cronk, species tree reduction dominant in range provided for large-scale evidence brought activity. have about To by human island from the collections early date, flora island’s the in profound the changes for understanding aunique resource and colonisation its after soon island provide arecord of endemicoceanic the an flora of early collections These record of florain the herbaria. historical arich is there and 1958); 19 before the were collected endemic plant taxa of known 60% the London in (Dandy, Museum History by 1698 Stonestreet in Natural at the are and weremade collections colonisation. The surviving oldest soon after collected and of collecting. along been history has there that powers means European for importance strategic their since to amenable this particularly be may islands tic Atlan the and of period profound change– –a anthropogenic years 400 last the through patterns from those inferred processes the and patterns of diversity ness samples. archival using but tractable be may analyses textual or, historical indeed palaeoecological using to tractable be unlikely are ges quickly. Such chan surprisingly establish can diversity novel of molecular patterns how (2013) doerfer setting, acontinental to demonstrate in specimens museum used For Mende Hunds example, and scales. time or centennial over decadal change to examine and population studies genetic and evolutionary in plant samples val (2018) of archi use Bieker Martin and contain. increasing the reviewed recently of DNA they the nature sequencing due degraded to the Sanger traditional with or impossible wereutilise to difficult that or samples wood specimens herbarium to biodiversity include archival of for island example, studies in used be may that material of therange expanded also eventshave they but diversification of island basis genetic to the relating questions fundamental to encompass studies lutionary evo of scope island potential the extended haveNGS not techniques only greatly 2016) (Thiers, years. four specimens hundred overcollected than more botanical Herbarium collections are a far from perfect sample of diversity as they are are they as sample of diversity from perfect afar are collections Herbarium In the case of St Helena, for example, its flora was studied, documented studied, of St its Helena, florawas for example, case the In robust the to test of source data apotential thus are specimens Herbarium th century ------
REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 83 REVISTA SCIENTIA INSULARUM, 3; 2020, PP. 73-88 84 and and Ross’s Clarke on Surgeon James Erebus Assistant an as time his through gained ras flo island experience of hand first rootedhis It in was islands. of oceanic diversity tfall’ in island biogeographic and evolutionary studies. studies. evolutionary biogeographic and island in tfall’ of ‘Hookerian shor the scale the to help of source data understand apotential vide today pro collections diversity. to document island hehistorical used Those that of others collections the and on made islands he himself collections the tions: both ‘Hookerian shortfall’. the as diversity of island understanding our in shortfall this poorly We understood. remains propose we should here recognise scales that tionary evolu across varies how and this on islands observed patterns diversity the explain processes natural than rather impact extent The which to human impact. of human evolutionand assemblyextentthe the and biota island of regarding questions ficant Both authors contributed to the writing of this paper. authors of contributed this Both writing to the paper. comments on this constructive for referees their anonymous azorica of relationships Sanicula the in for insights Schaefer 4, Hanno Terror Terror reflected his his theprofoundHooker’s interestin plant reflected floras on Insular lecture Hooker’s interest in islands was rooted in his work with botanical collec work botanical Hooker’s rootedhis with in was islands interest in We are grateful to Aida Pupo-Correia for the photographs used in Figure Figure in for photographs the used Pupo-Correia to Aida We grateful are expedition and his subsequent signi his work It and Floras. expedition on addressed island 4. AUTHORS CONTRIBUTION AUTHORS 4. 3. ACKNOWLEDGMENTS 2. CONCLUSIONS 2. Recibido : junio de 2019; de : junio aceptado : enero de 2020 de : enero and to two to two and ------Diniz-Filho Dias Dias Darwin Dandy Almén Cronk Cronk Bentham Cronk Carine , V.C. &Martin Bieker Berry Cardoso Brown Bowdish Boieiro Bochaton , E. 2007. A Chegada dos portugueses às ilhas –o antes e o depois– Açores. In: Silva, J. (ed). Silva, In: Açores. eodepois– antes –o ilhas às portugueses dos 2007., E. AChegada , E.F., Kilian Biol. J. Linn. Soc Linn. J. Biol. islands. and Hooker 2009. , R.J. , J.E. 1958. J.E. , , M.S., Lamichhaney, M.S., , Q.C.B. 2016. Plant extinctions take time. Science time. take , Q.C.B. 2016. extinctions Plant 119 Nelson A. pp. Oswestry: Helena. St of flora endemic The , Q.C.B. 2000. , Q.C.B. 1986. The decline of the St Helena ebony Trochetiopsisebony the St Helena of , Q.C.B. 1986. melanoxylon decline The , J.H. &Lomolino , J.H. , M.A., Francisco-Ortega , M.A., , M., Matthews , M., On the origin of species by means of natural selection natural of means by 1859., C. species of origin the On Trends in Ecology and Evolution and Trends Ecology in conservation. biodiversity in falls a Protecção da Natureza: 135-164. Natureza: da a Protecção para e Liga o Desenvolvimento para Americana Luso Fundação SA, Social Comunicação Público, Ilhas. das - A Floresta e Madeira Volume Açores 6, Portugal. de e Florestas Árvores ical accounts of the principal contributors niana & sequencing. BioEssays sequencing. Andersson 159-172. 91: 1070-1085. 91: Convolvulus in continent of the back-colonisation subsequent and aronesia Mac into colonisations for multiple evidence molecular floras: continental and of island Biol. Cons them. how overcome to and conservation Bot. 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