Moss-inhabiting diatoms from Campbell Island (sub-Antarctic)
Algen in Antarctica: hoe een reis naar het verleden zes nieuwe soorten oplevert
Haal eens diep adem. Een vierde van de lucht die zonet door jouw longen passeerde, werd geproduceerd door microscopische algjes die diatomeeën worden genoemd. In deze studie van Plantentuin Meise en de Universiteit van Antwerpen werden de diatomeeën, verborgen in een historische mos-collectie, grondig bestudeerd. De collectie werd een halve eeuw geleden verzameld nabij het zuidpoolgebied. Zes nieuwe soorten werden ontdekt. De studie hoopt bovendien om taxonomie weer sexy te maken bij jonge biologen.
Ze produceren meer zuurstof dan alle regenwouden gecombineerd, dienen als filter tijdens het produceren van wijn, vormen een hulpbron in het forensisch onderzoek naar verdrinkingsdood, zijn een veelbelovende toekomstige biobrandstof en zelfs een potentieel kankermedicijn. Kiezelwieren of diatomeeën zijn ware manusjes-van-alles die ondanks hun microscopische grootte in staat zijn tot grootse dingen.
Deze minuscule algen zijn te vinden in eender welke vochtige omgeving, gaande van de aardbodem, rivieren en verschillende soorten mos over oceanen tot zelfs de glazen wanden van een aquarium. Elk kiezelwier leeft in zijn zelfgebouwde glazen huisje, dat versierd is met talrijke prachtige patronen waarmee biologen de soort kunnen achterhalen. Pas in de 18e eeuw werden ze ontdekt door een onbekende Britse man die door zijn microscoop tuurde en melding maakte van ‘vreemde rechthoekige wezens’. Hun evolutionaire geschiedenis begon zo’n 200 miljoen jaar geleden en resulteerde in naar schatting 100.000 soorten. Ter vergelijking, er zijn vandaag 50 dolfijnsoorten en 20.000 vlindersoorten.
Taxonomie: what’s in a name?
Diatomeeën zijn extreem gevoelig aan verandering in hun omgeving. Een kleine wijziging in temperatuur of zoutgehalte kan ervoor zorgen dat sommige soorten het voor bekeken houden en plaats ruimen voor andere soorten die zo’n verandering wel kunnen smaken. Die gevoeligheid maakt van kiezelwieren uitstekende bio-indicatoren. Zo worden ze in Vlaanderen gebruikt om de kwaliteit van de waterlopen te schatten, maar ook in klimaatonderzoek spelen ze een hoofdrol. Via fossiele kiezelwieren kunnen bijvoorbeeld reconstructies worden gemaakt van het klimaat dat miljoenen jaren geleden heerste, maar ook de impact van de huidige klimaatverandering kan ermee worden beoordeeld. Zeker in de poolgebieden is dit belangrijk, aangezien zowel de noord- als zuidpool de eerste tekenen van klimaatverandering weerspiegelen. Al deze toepassingen zijn echter alleen maar mogelijk dankzij nauwkeurig soortonderzoek, maar daar loopt het vaak mis.
“Hoe is het mogelijk om de biodiversiteit te beschermen wanneer we niet eens weten hoeveel en welke soorten er rondvliegen, -wandelen of -zwemmen op de planeet?”
De immense taak om alle dieren, planten, diatomeeën en andere wezens op aarde te beschrijven en te benoemen behoort toe aan de wetenschap die taxonomie wordt genoemd. Taxonomie is een essentiële discipline, want hoe is het mogelijk om de biodiversiteit te beschermen wanneer we niet eens weten hoeveel en welke soorten er rondvliegen, -wandelen of -zwemmen op de planeet? Ondanks dat taxonomisch onderzoek hét fundament is van de biologie, is het vandaag een ondergesneeuwde wetenschap, mede door de opkomst van moleculaire analyses.
In Europa en Noord-Amerika is er relatief veel geweten over de diversiteit aan kiezelwiersoorten, maar eenmaal buiten deze gebieden staat het taxonomisch onderzoek nog in kinderschoenen. Vooral in de poolgebieden is er nog werk. Dat komt omdat biologen vroeger dachten dat de diatomeeënsoorten die ze in Europa en Noord-Amerika zagen, ook elders voorkwamen. Vergelijk het met vogelkundigen die vogels gaan spotten in Maleisië met een Europese vogelgids onder de arm, waarop vervolgens alle waargenomen Maleisische vogels een Europese naam krijgen: een biologisch rampscenario. Om deze vroegere praktijken weer te corrigeren, is er een heuse taxonomische herziening nodig waarbij alle soorten opnieuw kritisch worden onderzocht. Een werk van (zeer) lange adem.
Een reisje door de geschiedenis: historische mossen uit het zuidpoolgebied
In deze studie werden de diatomeeënsoorten van Campbell Island onder de microscopische loep genomen. Campbell Island is een afgelegen eiland nabij het zuidpoolgebied en ligt in het sub-Antarctische gedeelte. Het eiland bevindt zich in de zuidelijke Grote Oceaan, zo’n 700 km ten zuiden van Nieuw-Zeeland en ongeveer 3.500 km ten noorden van Antarctica. Verschillende mossen en struiken sieren het landschap. De mossen, waarin talrijke soorten kiezelwieren leven, werden tijdens de winter van 1969-1970 verzameld door Professor Dale Vitt en bewaard in het mos-herbarium in Cambridge, het Verenigd Koninkrijk. De mossen werden pas een halve eeuw later weer van onder het stof gehaald, speciaal voor deze studie, om te achterhalen welke soortenrijkdom aan diatomeeën er in de oude plantjes schuilt.
“Dure expedities naar de andere kant van de wereld zijn vaak overbodig voor degelijk taxonomisch onderzoek”
De kiezelwieren werden uit de historische mossen gefilterd en op microscopische preparaten gemonteerd. Door middel van zowel de licht- als elektronenmicroscoop werd de verscheidenheid aan wiertjes duidelijk. Meer dan 140 soorten werden er gevonden, waarvan meer dan de helft niet kon worden thuisgebracht in bestaande namen en waarschijnlijk volledig nieuw is. Zes soorten werden tijdens dit onderzoek beschreven en kregen een gloednieuwe benaming. Een van de nieuw ontdekte soorten – Angusticopula cosmica – lijkt op een kosmos gevuld met talloze sterren, waarnaar de naam verwijst. Een passende metafoor voor de kosmos aan kiezelwieren die nog wachten op hun ontdekking.
Werk aan de winkel
Niet enkel Campbell Island onderging een grondige analyse, de mos-bewonende wiertjes van alle eilanden in het gebied werden met elkaar vergeleken. Wat blijkt? Campbell Island is een unieke biotoop waar bepaalde diatomeeën zelfs sneller evolueren in vergelijking met de andere eilanden. De reden hierachter is niet duidelijk, al wordt vermoed dat kiezelwieren van andere eilanden in de zuidelijke Indische Oceaan en zelfs Zuid-Amerika meeliften met de wind en vervolgens op Campbell Island belanden, waar ze ongestoord en naar hartenlust kunnen evolueren.
Het gigantisch aantal onbekende soorten dat werd gevonden tijdens dit onderzoek maakt pijnlijk duidelijk dat toekomstig taxonomisch onderzoek broodnodig is, zeker als we diatomeeën willen blijven gebruiken in allerhande toepassingen, klimaatonderzoek voorop. Deze studie bewijst ook dat dure expedities naar de andere kant van de wereld vaak overbodig zijn voor degelijk taxonomisch onderzoek. In historische collecties van over de hele wereld zitten ongetwijfeld nog talloze onontdekte soorten verborgen, geduldig wachtend op een nieuwe generatie gepassioneerde taxonomen.
[1]
F. E. Round, R. M. Crawford and D. G. Mann, The Diatoms, Cambridge: Cambridge University Press, 1990.
[2]
B. M. Knudson and T. H. Kipling, "Ecology of the Epiphytic Diatom Tabellaria Flocculosa (Roth) Kutz. Var. Flocculosa in Three English Lakes," Journal of Ecology, Vols. Vol. 45, No. 1, pp. 93-112, 1957.
[3]
J. Seckbach and J. P. Kociolek, The Diatom World, Dordrecht Heidelberg London New York: Springer, 2011.
[4]
E. De Tommasi, J. Gielis and A. Rogato, "Diatom Frustule Morphogenesis and Function: A Multidisciplinary Survey," Marine Genomics, vol. 35, p. 10.1016, 2017.
[5]
J. P. Smol and E. F. Stoermer, The diatoms: Applications for the Environmental and Earth Sciences, Cambridge: Cambridge University Press, 2010.
[6]
H. Van Heurck, A Treatise on the Diatomaceae, London: William Wesley & Son, 1896.
[7]
O. P. Sharma, Textbook of Algae, New York : Tata McGraw-Hill Education, 1986.
[8]
P. Kuczynska, M. Jemiola-Rzeminska and K. Strzalka, "Photosynthetic Pigments in Diatoms," Marine drugs, vol. 13, no. 9, p. 5847–5881, 2015.
[9]
J. John, A Beginners Guide to Diatoms, Oberreifenberg: Koeltz Scientific Books, 2015.
[10]
S. M. Adl, A. G. B. Simpson, C. E. Lane, J. Lukes, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, L. Le Gall, D. H. Lynn, M. H., E. A. D. Mitchell, S. E. Mozley-Stanridge and Parfr, "The Revised Classification of Eukaryotes," The Journal of Eukaryotic Microbiology, vol. 59, no. 5, pp. 429-493, 2012.
[11]
F. E. Round and R. M. Crawford, "The Lines of Evolution of the Bacillariophyta. I. Origin," Proceedings of the Royal Society of London. Series B, Biological Sciences, vol. Vol. 211, no. No. 1183, pp. 237-260, 1981.
[12]
W. H. Kooistra and L. K. Medlin, "Evolution of the diatoms (Bacillariophyta). IV. A reconstruction of their age from small subunit rRNA coding regions and the fossil record," Molecular Phylogenetic Evolution, vol. 6, no. 3, pp. 391-407, 1996.
[13]
J. Schieber, D. Krinsley and L. Riciputi, "Diagenetic origin quartz silt in mudstones and implications for silica cycling," Nature, vol. 406, no. 6799, pp. 981-985, 2000.
[14]
J. B. H. Martiny, B. J. M. Bohannan, J. H. Brown, R. K. Colwell, J. A. Fuhrman, J. L. Green, M. C. Horner-Devine, M. Kane, J. A. Krumins, C. R. Kuske, P. J. Morin, S. Naeem, L. Ovreas, A.-L. Reysenbach, V. H. Smith and J. T. Staley, " Microbial biogeography: putting microorganisms on the map," Nature Reviews Microbiology , vol. 4, pp. 102-112, 2006.
[15]
D. R. Nemergut, E. K. Costello, M. Hamady, C. Lozupone, L. Jiang, S. K. Schmidt, N. Fierer, A. R. Townsend, C. C. Cleveland, L. Stanish and R. Knight, "Global patterns in the biogeography of bacterial taxa," Environmental Microbiology, vol. 13, no. 1, pp. 135-144, 2018.
[16]
L. G. M. Baas-Becking, Geobiologie of inleiding tot de milieukunde, The Hague: Van Stockum and Zoon, 1934.
[17]
D. Fontaneto and J. Hortal, "At least some protist species are not ubiquitous," Molecular Ecology, vol. 22, no. 20, pp. 5053-5055, 2013.
[18]
P. Vanormelingen, E. Verleyen and W. Vyverman, "The diversity and distribution of diatoms: from cosmopolitanism to narrow endemism," Biodiversity Conservation, vol. 17, pp. 393-405, 2008.
[19]
W. Vyverman, E. Verleyen, K. Sabbe, K. Vanhoutte, M. Sterken, D. A. Hodgson, D. G. Mann, S. Juggins, B. Van de Vijver, V. Jones, R. Flower, D. Roberts, V. A. Chepurnov, C. Kilroy, P. Vanormelingen and A. De Wever, "Historical processes contrain patterns in global diatom diversity," Ecology, vol. 88, no. 8, pp. 1924-1931, 2007.
[20]
C. Souffreau, P. Vanormelingen, E. Verleyen, K. Sabbe and W. Vyverman, "Tolerance of benthic diatoms from temperate aquatic and terrestrial habitats to experimental desiccation and temperature stress," Phycologia, vol. 49, no. 4, pp. 309-324, 2010.
[21]
B. Van de Vijver, P. Ledeganck and L. Beyens, "Three new species of Diadesmis from soils of Ile de la Possession (Crozet Archipelago, sub-Antarctic)," Cryptogamie Algologie, vol. 23, pp. 333-341, 2002.
[22]
B. Van de Vijver and G. Mataloni, "New and interesting species in the genus Luticola D.G. Mann (Bacillariophyta) from Deception Island (South Shetland Islands)," Phycologia, vol. 47, p. 451–467, 2008.
[23]
S. A. Spaulding, J. P. Kociolek and D. Wong, "A taxonomic and systematic revision of the genus Muelleria (Bacillariophyta)," Phycologia, vol. 38, pp. 314-341, 1999.
[24]
B. Van de Vijver, N. J. M. Gremmen and L. Beyens, "The genus Stauroneis (Bacillariophyceae) in the Antarctic region," Journal of Biogeography, vol. 32, p. 1791–1798, 2005.
[25]
E. J. Cox, "Morphogenetic information and the selection of taxonomic characters for raphid diatom systematics," Plant Ecology and Evolution, vol. 143, no. 3, pp. 271-277, 2010.
[26]
B. Van de Vijver, A. Jarlman, M. de Haan and L. Ector, "New and interesting diatom species (Bacillariophyceae) from Swedish rivers," Nova Hedwigia, vol. 141, p. 237–254, 2012.
[27]
D. G. Mann, "The species concept in diatoms," Phycologia, vol. 38, pp. 437-495, 1999.
[28]
J. P. Kociolek and S. A. Spaulding, "Freshwater diatom biogeography," Nova Hedwigia, vol. 71, pp. 223-241, 2000.
[29]
D. G. Mann and S. J. M. Droop, "Biodiversity, biogeography and conservation of diatoms," Hydrobiologia, vol. 336, no. 1-3, pp. 19-32, 1996.
[30]
P. H. Raven, R. F. Evert and S. E. Eichhorn, Biology of Plants, New York City: W. H. Freeman, 2005.
[31]
W. Vyverman, K. Sabbe, V. Chepurnov, K. Muylaert, C. Cocquyt, K. Vanhoutte and E. Verleyen, "Conservation of diatom biodiversity: issues and prospects," Brussels, 2001.
[32]
P. Falkowski, R. J. Scholes, E. Boyle, J. Canadell, D. Canfield, J. Elser, N. Gruber, K. Hibbard, P. Högberg, S. Linder, F. T. Mackenzie, B. Moore III, T. Pedersen, Y. Rosenthal, S. Seitzinger, V. Smetacek and W. Steffen, "The Global Carbon Cycle: A Test of Our Knowledge of Earth as a System," Science, vol. 290, pp. 291-296, 2000.
[33]
J. D. Hooker, The Botany of the Antarctic Voyage of H.M. Discovery Ships Erebus and Terror in the Years 1839-1843: Under the Command of Captain Sir James Clark Ross, London: Reeve Brothers, 1844.
[34]
C. G. Ehrenberg, Über neue Anschauungen des kleinsten nördlichen Polarlebens. Berichteüber die zur Bekanntmachung geeigneten Verhandlung der Königlich-Preussischen Akademieder Wissenschaften zu Berlin, Berlin: Monatsberichte, 1853.
[35]
P. T. Cleve, "Diatomaceer frän Spetsbergen," Öfversigt af Kongl. Vetenskaps-Akademiens Förhandlingar, vol. 24, pp. 661-669, 1867.
[36]
P. T. Cleve, "On diatoms from the Arctic Sea," Bihang till Kongl. Svenska Vetenskaps-Akademiens Handlingar, vol. 1, pp. 1-28, 1873.
[37]
P. T. Cleve, "Diatoms, collected during the expedition of the Vega," Ur Vega-Expeditionens Vettenskapliga Lakttagelser, vol. 3, pp. 457-517, 1883.
[38]
P. T. Cleve, "Diatoms from Baffin Bay and Davis Strait," Bihang till Kongl. Svenska Vetenskaps-Akademiens Handlingar, vol. 22, pp. 1-22, 1896.
[39]
P. T. Cleve, "Diatoms from Franz Josef Land," Bihang till Kongl. Svenska Vetenskaps-Akademiens Handlingar, vol. 24, pp. 1-26, 1898.
[40]
P. T. Cleve, "Notes on some Atlantic plankton organisms," Kongl. Svenska Vetenskaps-Akademiens Handlingar, vol. 34, p. 20, 1900.
[41]
P. T. Cleve and A. Grunow, "Beiträge zur Kenntniss der Arctischen Diatomeen," Kongl. Svenska Vetenskaps-Akademiens Handlingar, vol. 17, no. 2, pp. 1-121, 1880.
[42]
G. Dickie, "On the algae found during the Arctic Expedition," Journal of the Linnean Society, vol. 17, pp. 6-12, 1880.
[43]
A. Grunow, "Die Diatomeen von Franz-Josefs-Land," Denkschriften der Mathematisch-Naturwissenschaftlichen Klasse der Kaiserlichen Akademie der Wissenschaften, vol. 48, pp. 53-112, 1884.
[44]
R. Pienitz, M. S. V. Douglas and J. P. Smol, Long-term environmental change in Arctic and Antarctic lakes, Dordrecht: Springer, 2004.
[45]
B. Van de Vijver and L. Beyens, "Biogeography and Ecology of Freshwater Diatoms in Subantarctica: A Review," Journal of Biogeography , vol. 26, no. 5, pp. 993-1000, 1999.
[46]
J. D. Holmboe, "IV. Navicula mutica Kütz. Aus dem antarktischen Festlande," Nyt Magazin for Naturvidenskaberne, Christiana, vol. 40, no. 3, pp. 221-222, 1902.
[47]
N. Wille, "Antarktische Algen. Mitteilungen über einige von Borchgrevnik auf den antarktischen Pflanzen III," Nyt Magazine for Naturvidenskaberna, vol. 40, pp. 209-222, 1902.
[48]
L. Gain, "Note sur la Flore algologique d'eau douce de l'Antarctique sud-Américaine," Bulletin du Museum national d'histoire naturelle Paris, vol. 5, p. 371, 1911.
[49]
W. West and G. S. West, "Freshwater algae," British Antarctic Expedition 1907-1909, vol. 1, pp. 263-298, 1911.
[50]
G. W. F. Carlson, "Süsswasser-Algen aus der Antarktis, Süd-Georgien und den Falkland Inseln," in Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition 1901-1903, vol. 4, Stockholm, Lithographisches Institut des Generalstabs, 1913, pp. 1-94.
[51]
P. Petit, Diatomacées. Resultats de la Premiere Expédition Antarctique Française 1903-1905, commandée par le Dr. Charcot, Parijs: Botanique, 1908.
[52]
H. Van Heurck, Expedition Antarctique Belge. Résultats du voyage du S. Y. "Belgica" en 1897-1898-1899, Anvers: J.-E. Buschmann, 1909.
[53]
M. Peragallo, "Diatomées d'eau douce et diatomées d'eau salée," in Deuxiéme Expedition Antarctique Française 1908-1910, Parijs, Masson, 1921, pp. 1-98.
[54]
H. Heiden and R. W. Kolbe, "Die marinen Diatomeen der Deutschen südpolar Expedition 1901-1903," in Deutsche Südpolar Expedition 1901-1903, vol. 8, Berling, W. de Gruyter & Co., 1928, pp. 450-715.
[55]
P. F. Reinsch, "Die Süsswasseralgenflora von Süd-Georgien," in Die Deutschen Expeditionen und ihre Ergebnisse 1882-01883, Berlijn, A. Asher, 1890, pp. 329-365.
[56]
R. Le Cohu and R. Maillard, "Les diatomées monoraphidées des Îles Kerguelen," Annales de Limnologie, vol. 19, pp. 143-167, 1983.
[57]
R. Le Cohu and R. Maillard, "Diatomées d'eau douce des Îles Kerguelen (à l'exclusion des Monoraphidées)," Annales de Limnologie, vol. 22, pp. 99-118, 1986.
[58]
P. Bourrelly and E. Manguin, "Contribution à la Flore Algale d'eau Douce des Iles Kerguelen.," Mémoires de L'Institut Scientifique de Madagascar, séries B, vol. 5, pp. 5-58, 1954.
[59]
B. Van de Vijver, Y. Frenot and L. Beyens, "Freshwater diatoms from Ile de la Possession (Crozet Archipelago, Subantarctica)," Bibl. Diatomol., vol. 46, pp. 1-412, 2002.
[60]
R. Schmidt, R. Mäusbacher and J. Müller, "Holocene diatom flora and stratigraphy from sediment cores oftwo Antarctic lakes (King George Islands)," Journal ofPaleolimnology, vol. 3, pp. 55-74, 1990.
[61]
K. Sabbe, E. Verleyen, D. A. Hodgson, K. Vanhoutte and W. Vyverman, "Benthic diatom flora of freshwater and saline lakes in the Larsemann Hills and Rauer Islands, East Antarctica," Antarctic Science , vol. 15, no. 2, p. 227–248 , 2003.
[62]
B. Van de Vijver and N. Gremmen, "Three New Moss-inhabiting Diatom Species from sub-Antarctic Marion Island," Diatom Research, vol. 21, no. 2, pp. 427-439, 2006.
[63]
B. Van de Vijver and R. Zidarova, "Five new taxa in the genus Pinnularia sectio Distantes (Bacillariophyta) from Livingston Island (South Shetland Islands).," Phytotaxa, vol. 24, pp. 39-50, 2011.
[64]
B. Van de Vijver, "Revision of the Psammothidium manguinii complex (Bacillariophyta) in the sub–Antarctic Region with the description of four new taxa," Fottea, vol. 19, no. 1, pp. 90-106, 2019.
[65]
B. Van de Vijver and L. Beyens, "The genus Stauroneis (Bacillariophyceae) in the Antarctic Region," Journal of Biogeography, vol. 32, pp. 1791-1798, 2005.
[66]
B. Van de Vijver, M. de Haan and H. Lange-Bertalot, "Revision of the genus Eunotia (Bacillariophyceae) in the Antarctic Region," Plant Ecology and Evolution, vol. 147, pp. 256-284, 2014.
[67]
J. P. Kociolek, K. Kopalová, S. E. Hamsher, T. J. Kohler, B. Van de Vijver, P. Convey and D. M. McKnight, "Freshwater diatom biogeography and the genus Luticola: an extreme case of endemism in Antarctica," Polar Biology, vol. 40, pp. 1185-1196, 2017.
[68]
J. Frenguelli, "Diatomeas de Tierra del Fuego.," Anales de la Sociedad Cientijica Argentina, vol. 97, pp. 87-118, 231-66, 1924.
[69]
P. Bourrelly and E. Manguin, "Contribution ala flore algale d'eau douce des lies Kerguelen," Memoires de l'lnstitut Scientijique de Madagascar, vol. 5, pp. 7-58, 1954.
[70]
V. J. Jones, S. Juggins and J. C. Ellis-Evans, "The relation ship between water chemistry and surface sediment diatom 1; assemblages in maritime Antarctic lakes," Antarctic Scienc, vol. 5, pp. 339-348, 1993.
[71]
J. S. Bunt, "Soil-inhabiting nematodes from Macquarie Island," Australian Journal of Zoology 2, pp. 264-274, 1954.
[72]
D. H. Vitt, "A key and synopsis of the mosses of Campbell Island, New Zealand," New Zealand Journal of Botany, vol. 12, no. 2, pp. 185-210, 1974.
[73]
N. J. M. Gremmen, B. Van de Vijver, Y. Frenot and M. Lebouvier, "Distribution of moss-inhabiting diatoms along an altitudinal gradient at sub-Antarctic Îles Kerguelen," Antarctic Science, vol. 19, pp. 17-24, 2007.
[74]
B. Van de Vijver and L. Beyens, "Moss diatom communities from Ile de la Possession (Crozet, sub-Antarctica) and their relationship with moisture," Polar Biology, vol. 22, pp. 232-240, 1999.
[75]
N. Chipev and D. Temniskova-Topalova, "Diversity dynamics and distribution of diatom assemblages in land habitats on Livingston Island (Antarctica)," Bulgarian Antarctic Research, vol. 2, pp. 32-42, 1999.
[76]
A. Suarez and N. D. Tsutsui, "The Value of Museum Collections for Research and Society," BioScience, vol. 54, no. 1, pp. 66-74, 2004.
[77]
E. K. Meineke, C. C. Davis and T. J. Davies, "The unrealized potential of herbaria for global change biology," Ecological Monographs, vol. 88, pp. 505-525, 2018.
[78]
D. S. Park and D. Potter, "Why close relatives make bad neighbours: phylogenetic 460 conservatism in niche preferences and dispersal disproves Darwin's naturalization 461 hypothesis in the thistle tribe," Molecular Ecology, vol. 24, pp. 3181-3193, 2015.
[79]
G. H. Pyke and P. R. Ehrlich, "Biological collections and ecological/environmental research: A review, some observations and a look to the future.," Biological Reviews, vol. 85, pp. 247-266, 2010.
[80]
M. Poulin, "Craspedopleura (Bacillariophyta), a new diatom genus of arctic sea ice assemblages," Phycologia, vol. 32, no. 3, pp. 223-233, 1992.
[81]
J. P. Kociolek, K. Kulikovskiy and C. N. Solak, "The diatom genus Gomphoneis Cleve (Bacillariophyceae) from Lake Baikal, Russia," Phytotaxa, vol. 154, no. 1, pp. 1-37, 2013.
[82]
R. J. Stevenson and E. F. Stoermer, "Diatoms from the Great Lakes. II. Some Rare or Poorly Known Species of the Genus Navicula," Journal of Great Lakes Research, vol. 4, no. 2, pp. 178-185, 1978.
[83]
E. F. Stoermer and J. J. Yang, "Contributions to the diatom flora of the Laurentian Great Lakes. I. New and little-known species of Amphora (Bacillariophyta Pennatibacillariophyceae)," Phycologia, vol. 10, no. 4, pp. 397-409, 1971.
[84]
C. Cocquyt, W.-H. Kusber and R. Jahn, "Otto Müller’s Surirella taxa (Bacillariophyta) from East Africa,based on a historical collection kept at the Botanic Garden and Botanical Museum Berlin-Dahlem (B)," Afrika Focus, vol. 21, pp. 65-75, 2008.
[85]
A. Tuji, "Examination of Type Material and Typification of Seven Diatoms Described by C. G. Ehrenberg," in National Museum of Nature and Science Monographs No. 40, Tokyo, National Science Museum, 2009.
[86]
D. M. Williams and J. P. Kociolek, "Historical Biogeography of Diatoms in Australasia: A Preliminary Assessment," in Handbook of Australasian Biogeography, Boca Raton, Florida, CRC Press, 2017.
[87]
L. Denys, "Historical distribution of'Red List diatoms'(Bacillariophyceae) in Flanders (Belgium)," Systematics and Geography of Plants, vol. 40, pp. 409-420, 2000.
[88]
W. R. Harding, C. G. M. Archibald and J. C. Taylor, "The relevance of diatoms for water quality assessment in South Africa: A position paper," Water SA, vol. 31, no. 1, pp. 41-46, 2005.
[89]
J. C. Taylor, W. R. Harding, C. G. Archibald and L. van Rensburg, "Diatoms as indicators of water quality in the Jukskei-Crocodile river system in 1956 and 1957, a re-analysis of diatom count data generated by BJ Cholnoky," Water SA, vol. 31, no. 2, pp. 237-246, 2005.
[90]
S. L. Chown and P. Convey, "Spatial and temporal variability across life's hierarchies in the terrestrial Antarctic," Philosophical Transactions of the Royal Society, vol. 362, no. 1488, pp. 2307-2331, 2007.
[91]
E. J. Godley, "The botany of Southern Chile in relation to New Zealand and the subantarctic," Proceedings of the Royal Society B: Biological Sciences, vol. 152, pp. 457-474, 1960.
[92]
F. G. Alberts, Geographic Names of the Antarctic, Arlington: National Science Foundation, 1995.
[93]
J. D. Hanson and J. E. Gordon, Antarctic environments and resources: a geographical perspective, New York: Longman Ltd., 1998.
[94]
B. Van de Vijver, H. Lange-Bertalot, C. E. Wetzel and L. Ector, "Michelcostea, a new diatom genus (Bacillariophyta) from the sub-Antarctic Region," Nova Hedwigia, vol. 146, pp. 125-136, 2017.
[95]
E. Verleyen, B. Van de Vijver, B. Tytgat, E. Pinseel, D. A. Hodgson, K. Kopalová, S. L. Chown, E. Van Ranst, S. Imura, S. Kudoh, W. Van Nieuwenhuyze, A. Consortium, K. Sabbe and W. Vyverman, "Geographic Structuring and Endemism in Freshwater Diatom Floras in the Antarctic Realm," in Diversity, Biogeography and Evolutionary History of Diatoms: Pinnularia borealis as a Case Study, Ghent, Ghent University, 2019, pp. 55-91.
[96]
E. Rainsley, C. S. M. Turney, N. R. Golledge, J. M. Wilmshurst, M. Mcglone, A. G. Hogg, B. Li, Z. A. Thomas, R. Roberts, R. Jones, J. G. Palmer, V. Flett, G. A. De Wet, D. K. Hutchinson, M. Lipson, P. Fenwick, B. R. Hines, U. Binetti and Fo, "Pleistocene glacial history of the New Zealand subantarctic islands," Climate of the Past, vol. 15, no. 2, pp. 423-448, 2019.
[97]
S. L. Chown, N. J. M. Gremmen and K. J. Gaston, "Ecological Biogeography of Southern Ocean Islands: Species-Area Relationships, Human Impacts, and Conservation," The American Naturalis, vol. 152, no. 4, pp. 562-575, 1998.
[98]
M. R. Clark and P. R. Dingwall, Conservation of islands in the southern ocean: a review of the protected areas of Insulantarctica, Cambridge: IUCN, 1985.
[99]
P. G. Quilty, "Origin and evolution of the sub-antarctic islands: the foundation," Papers and Proceedings of the Royal Society of Tas- mania, vol. 141, pp. 35-58, 2007.
[100]
V. C. Thorn, "New Zealand sub-Antarctic phytoliths and their potential for past vegetation reconstruction," Antarctic Science, vol. 20(1), pp. 21-32, 2008.
[101]
C. D. Meurk, M. N. Foggo and J. B. Wilson, "The Vegetation of Subantarctic Campbell Island," New Zealand Journal of Ecology, vol. 18(2), pp. 123-168, 1994.
[102]
S. J. Reid, "Surface wind frequencies in the southwest Pacific estimated from radar-wind data," New Zealand Journal ofScience., vol. 25, pp. 303-311, 1982.
[103]
C. D. Meurk and D. R. Given, Vegetation map of Cambell Island 1:25000, Christchurch, New Zealand: DSIR Land Resources, 1990.
[104]
M. S. McGlone, N. T. Moar, P. Wardle and C. D. Meurk, "Late-glacial and Holocene vegetation and environment of Campbell Island, far southern New Zealand," The Holocene, vol. 7(1), pp. 1-12, 1997.
[105]
C. D. Meurk, M. N. Foggo, B. M. Thomson, E. T. John Bathurst and M. B. Crompton, "Ion-rich Precipitation and Vegetation Pattern on Subantarctic Campbell Island," Arctic and Alpine Research, vol. 26, no. 3, pp. 281-289, 1994.
[106]
P. J. Dilks and P. R. Wilson, "Feral sheep and cattle and royal albatrosses on Cambell Island; population trends and habitat changes," New Zealand Journal of Zoology, vol. 6, pp. 127-139, 1979.
[107]
K. M. Saunders, D. A. Hodgson, S. McMurtrie and M. Grosjean, "A diatom-conductivity transfer function for reconstructing past changes in the Southern Hemisphere westerly winds over the Southern Ocean," Journal of Quaternary Science, vol. 30, no. 5, pp. 464-477, 2015.
[108]
M. Hickman and D. Vitt, "The aerial epiphytic flora of moss species from Subantarctic Campbell Island," Nova Hedwigia , vol. 24, pp. 443-458, 1974.
[109]
A. van der Werff, "A new method of concentrating and cleaning diatoms and other organisms," SIL Proceedings, vol. 12, no. 1, pp. 276-277, 1955.
[110]
B. Van de Vijver and L. Beyens, "The epiphytic diatom flora of mosses from Strømness Bay area, South Georgia," Polar Biology, vol. 17, no. 6, pp. 492-501, 1997.
[111]
B. Van de Vijver, L. Beyens, S. Vincke and N. J. M. Gremmen, "Moss-inhabiting diatom communities from Heard Island, sub-Antarctic," Polar Biology, vol. 27, no. 9, p. 532–543, 2004.
[112]
B. Van de Vijver, N. Gremmen and V. Smith, "Diatom communities from the sub-Antarctic Prince Edward Islands: diversity and distribution patterns," Polar Biology, vol. 31, no. 7, p. 795–808, 2008.
[113]
K. Kopalová, R. Ochyra, L. Nedbalová and B. Van de Vijver, "Moss-inhabiting diatoms from two contrasting Maritime Antarctic islands," Plant Ecology and Evolution, vol. 147, no. 1, pp. 67-84, 2014.
[114]
W. L. Kovach, MVSP-A multivariate statistical package for Windows, Version 3.13, Anglesey: Kovak Computing Services, 2002.
[115]
A. Chao, "Non-parametric estimation of the number of classes in a population.," Scandinavian journal of statistics, vol. 11, pp. 265-270, 1984.
[116]
R. K. Colwell, EstimateS: statistical estimation of species richness and shared species from samples. Version 9, User's guide and Applications published at http://viceroy.eeb.uconn.edu/estimates/.
[117]
T. Sørensen, "A Method of Establishing Groups of Equal Amplitudes in Plant Sociology Based on Similarity of Species Content and its Application to Analyses of the Vegetation on Danish Commons," Kongelige Danske Videnskabernes Selskab, Biologiske Skrifter, vol. 5, pp. 1-34, 1948.
[118]
C. J. F. ter Braak and P. Smilauer, CANOCO Reference Manual and Users Guide to Canoco for Windows: Software for Canonical Community Ordination (Version 4.5), Ithaca, NY: USA: Microcomputer Power, 2002.
[119]
R. H. Jongman, C. J. F. ter Braak and O. F. R. Van Tongeren, Data analysis in community and landscape ecology, Cambridge: Cambridge University Press, 1995.
[120]
B. Van de Vijver, P. Ballings and C. Goeyers, "Frankophila dalevittii, a new freshwater diatom (Bacillariophyta) from Campbell Island," Phytotaxa, vol. 429, no. 1, pp. 57-64, 2020.
[121]
M. S. M. Sosef, J. Degreef, H. Engledow and P. Meerts, Botanical classification and nomenclature, an introduction, Meise: Meise Botanic Garden, 2020.
[122]
C. Souffreau, P. Vanormelingen, E. Verleyen, K. Sabbe and W. Vyverman, "Tolerance of benthic diatoms from temperate aquatic and terrestrial habitats to experimental desiccation and temperature stress," Phycologia, vol. 49, no. 4, pp. 309-324, 2010.
[123]
B. Van de Vijver and L. Beyens, "Freshwater diatoms from some islands in the maritime Antarctic region," Antarctic Science, vol. 9, no. 4, pp. 418-425, 1997.
[124]
C. Goeyers and B. Van de Vijver, "Revision of the non-marine centric diatom flora (Bacillariophyta) of the sub-Antarctic Campbell Island (southern Pacific Ocean) with the descriptions of five new species," European Journal of Taxonomy, vol. 694, pp. 1-30, 2020.
[125]
B. Van de Vijver, P. Scholberg and M. Lebouvier, "Pinnularia schoelynckiana, a new limnoterrestrial diatom species (Bacillariophyta) from the sub- Antarctic Iles Kerguelen (southern Indian Ocean)," Botany Letters, vol. 167, no. 1, pp. 42-49 , 2019.
[126]
B. Van de Vijver, R. Zidarova, M. Sterken, E. Verleyen, M. de Haan, W. Vyverman, F. Hinz and K. Sabbe, "Revision of the genus Navicula s.s. (Bacillariophyceae) in inland waters of the Sub-Antarctic and Antarctic with the description of five new species," Phycologia, vol. 50, no. 3, pp. 281-297, 2011.
[127]
T. B. Kellogg and D. E. Kellogg, Non-marine and Littoral Diatoms from Antarctic and Subantarctic Regions: Distribution and Updated Taxonom, Rugell: A.R.G. Gantner Verlag K.G., 2002.
[128]
B. Van de Vijver, L. Ector and D. Williams, "Observations on and typification of Gomphonema auritum A.Braun ex Kützing (Gomphonemataceae, Bacillariophyta)," Notulae algarum, vol. 148, pp. 1-6, 2020.
[129]
J. A. E. Gibson, D. Roberts and B. Van de Vijver, "Salinity control of the distribution of diatoms in lakes of the Bunger Hills, East Antarctica," Polar Biology, vol. 29, no. 8, pp. 694-704, 2006.
[130]
V. J. Jones, "The diversity, distribution and ecology of diatoms from Antarctic inland waters," Biodiversity and Conservation, vol. 5, p. 1433–1449, 1996.
[131]
G. Moser, H. Lange-Bertalot and D. Metzeltin, "Insel der Endemiten Geobotanisches Phänomen Neukaledonien (Island of endemics New Caledonia - a geobotanical phenomenon).," in Bibliotheca Diatomologica, Berlin, J. Cramer, 1998.
[132]
H. Lange-Bertalot, M. Bak and A. Witkowski, "Eunotia and some related genera," in Diatoms of Europe. Diatoms of the European inland water and comparable habitats. Volume 6, Rugell, A.R.G. Gantner Verlag K.G., 2011.
[133]
M. Hirano, "Freshwater algae in the Antarctic Regions," Biogeography and Ecology in Antarctica, vol. 15, pp. 127-193, 1965.
[134]
H. Lange-Bertalot, "Iconographia Diatomologica," in Annotated Diatom Micrographs, Köningstein, Koeltz Scientific Books, 1996.
[135]
B. Van de Vijver, M. de Haan and C. Goeyers, "Analysis of the type material of Navicula tenuis Krasske (Bacillariophyta) and its transfer to the genus Adlafia," Notulae algarum, vol. 115, pp. 1-3, 2019.
[136]
N. Foged, "Diatoms in New Zealand, the North Island," Bibliotheca Phycologica, vol. 47, pp. 1-224, 1979.
[137]
J. John, Diatoms from Tasmania. Taxonomy and Biogeography. The Diatom Flora of Australia. Volume 2, Königstein: Koeltz Scientific Books, 2018.
[138]
R. Zidarova, K. Kopalová, B. Van de Vijver and H. Lange-Bertalot, "Diatoms from the Antarctic Region, Maritime Antarctica," in Iconographia Diatomologica Volume 24, Köningstein, Koeltz Scientific Books, 2016.
[139]
R. Jahn, W.-H. Kusber and O. E. Romero, "Cocconeis pediculus Ehrenberg and C. placentula Ehrenberg var. placentula (Bacillariophyta): Typification and taxonomy," Fottea, vol. 9, no. 2, pp. 275-288, 2009.
[140]
J. C. Kingston, "Araphid and Monoraphid Diatoms," in Freshwater Algae of North America. Ecology and Classification, San Fransisco, Academic Press, 2003.
[141]
M. Kulikovskiy, S. A. Andreeva, E. S. Gusev, I. V. Kuznetsova and N. V. Annenkova, "Molecular Phylogeny of Monoraphid Diatoms and Raphe Significance in Evolution and Taxonomy," Biology Bulletin, vol. 43, pp. 398-407, 2016.
[142]
I. Jüttner, C. E. Wetzel, D. M. Williams and L. Ector, "Investigations of the type materials of Achnanthes parallela J.R.Carter and Achnanthes petersenii Hustedt (Bacillariophyceae) with comments on the genus Rossithidium Round & Bukhtiyarova," Botany Letters, vol. 167, no. 1, pp. 57-69, 2019.
[143]
R. L. Lowe, P. Kociolek, J. R. Johansen, B. Van de Vijver, H. Lange-Bertalot and K. Kopalová, "Humidophila gen. nov., a new genus for a group of diatoms (Bacillariophyta) formerly within the genus Diadesmis: species from Hawai'i, including one new species," Diatom Research Publication details, including , vol. 29, no. 4, pp. 351-360, 2014.
[144]
B. Van de Vijver, L. Ector and E. J. Cox, "Ultrastructure of Diatomella balfouriana with a discussion of septum-like structures in diatom genera," Diatom Research, vol. 27, no. 4, pp. 213-221, 2012.
[145]
E. Pinseel, B. Van de Vijver, J. Kavan, E. Verleyen and K. Kopalová, "Diversity, ecology and community structure of the freshwater littoral diatom flora from Petuniabukta (Spitsbergen)," Polar Biology, vol. 40, p. 533–551, 2017.
[146]
S. Elias and C. Mock, Encyclopedia of Quaternary Science, Amsterdam: Elsevier, 2013.
[147]
J. Seckbach and P. Kociolek, The Diatom World, Berlin: Springer Science & Business Media, 2011.
[148]
D. G. Mann and P. Vanormelingen, "An Inordinate Fondness? The Number, Distributions, and Origins of Diatom Species," Journal of Eukaryotic Microbiology, pp. 414-420, 2013.
[149]
O. Shields, "World Numbers of Butterflies," Journal of the Lepidopterists' Society, vol. 43, pp. 178-183, 1989.
[150]
L. M. Mukhametov, "Unihemispheric slow-wave sleep in the Amazonian dolphin, Inia geoffrensis," Neuroscience Letters, vol. 79, pp. 128-132, 1987.
[151]
F. Schütt, "Bacillariales (Diatomeae)," in Die natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten. I. Teil. 1. Abteilung b: Gymnodiniaceae, Prorocentricaceae, Peridiniaceae, Bacillariaceae, Leipzig, Wilhelm Engelmann, 1896, p. 31–150.
[152]
J. S. L. Gilmour and J. Heslop-Harrison, "The deme terminology and the units of micro-evolutionary change," Genetica, vol. 27, pp. 147-161, 1954.
[153]
E. F. Keller and E. A. Lloyd, Keywords in Evolutionary Biology, Harvard: Harvard University Press, 1994.
[154]
D. J. Campbell and M. R. Rudge, "Vegetation changes induced over ten years by goats and pigs at Port Ross, Auckland Islands (Subantarctic)," New Zealand Journal of Ecology, vol. 7, pp. 103-118, 1984.
[155]
M. S. McGlone and N. T. Moar, "Pollen-vegetation relationships on the subantarctic Auckland Islands, New Zealand," Review of Palaeobotany and Palynology, vol. 96, pp. 317-338, 1997.
[156]
N. J. M. Gremmen, B. Van de Vijver, Y. Frenot and M. Lebouvier, "Distribution of moss-inhabiting diatoms along an altitudinal gradient at sub-Antarctic Îles Kerguelen," Antarctic Science, vol. 19, no. 1, pp. 17-24, 2007.
[157]
J. S. Bunt, "A comparative account of the terrestrial diatoms of Macquarie Island," Proceedings of the Linnean Society of New South Wales, vol. LXXIX, no. parts 1-2, pp. 34-57, 1954.
[158]
E. W. Sanderson, M. Jaiteh, M. A. Levy, K. H. Redford, A. V. Wannebo and G. Woolmer, "The human footprint and the last of the wild," BioScience, vol. 52, pp. 891-904, 2002.
[159]
A. Terauds, S. L. Chown, F. Morgan, H. J. Peat, D. J. Watts, H. Keys, P. Convey and D. M. Bergstrom, "Conservation biogeography of the Antarctic," Biodiversity Research, vol. 18, pp. 726-741, 2012.
[160]
J. P. Smol, A. P. Wolfe, H. J. B. Birks, M. S. V. Douglas, V. J. Jones, A. Korhola, R. Pienitz, K. Rühland, S. Sorvari, D. Antoniades, S. J. Brooks, M. Fallu, M. Hughes, B. E. Keatley, T. E. Laing, N. Michelutti, L. Nazarova and M. Nyman, "Climate-driven regime shifts in the biological communities of arctic lakes," Proceedings of the National Academy of Sciences, vol. 102, no. 12, pp. 4397-4402, 2005.
[161]
H. J. B. Birks, "The importance of pollen and diatom taxonomic precision in quantitative palaeoenvironmental reconstructions," Review of Palaeobotany and Palynology, vol. 83, pp. 107-117, 1994.
[162]
G. W. Prescott, A contribution to a bibliography of Antarctic and Subantarctic algae, Stuttgart: J. Cramer, 1979.
[163]
K. Sabbe, W. Vyverman, L. Ector, C. E. Wetzel, J. John, D. A. Hodgson, E. Verleyen and B. Van de Vijver, "On the identity of Navicula gottlandica (Bacillariophyta), with the description of two new species Navicula eileencoxiana and Navicula bergstromiana from the Australo-Pacific region," Plant Ecology and Evolution , vol. 152, no. 2, p. 313–326, 2019.
[164]
American Water Works Association, Algae Source to Treatment: M57, Denver: American Water Works Association, 2010.
[165]
S. Ekman, Tiergeography des Meeres, Leipzig: Akademie Verlagsgesellschaft, 1939.
[166]
S. Ekman, Zoogeography of the sea, London: Sidgwich and Jackson Ltd., 1953.
[167]
A. E. Ortmann, Grundzüge der Marinen Tiergeographie, Jena: Gustav Fischer, 1896.
[168]
C. De Broyer, P. Koubbi, H. J. Griffiths and et al., Biogeographic Atlas of the Southern Ocean, Cambridge, UK: Scientific Committee on Antarctic Research, 2014.
[169]
E. J. Cox, "Diatom identification in the face of changing species concepts and evidence of phenotypic plasticity," Journal of Micropalaeontology, vol. 33, pp. 111-120, 2014.
[170]
B. Abell-Seddon, Introduction to biogeography, London, UK: Duckworth, 1971.
[171]
I. Sanmartín, "Historical Biogeography: Evolution in Time and Space," Evolution: Education & Outreach, vol. 5, pp. 555-568, 2012.
[172]
B. C. Cox and P. D. Moore , Biogeography: an ecological and evolutionary approach, Hoboken, NJ: Wiley, 2010.
[173]
K. Jürgens and R. Massana, "Protistan grazing on marine bacterioplankton," in Microbial Ecology of the Oceans, Hoboken, New Jersey, John Wiley & Sons, 2008.
[174]
B. Van de Vijver, P. Ledeganck and L. Beyens, "Habitat preferences in freshwater diatom communities from sub-Antarctic iles Kerguelen," Antarctic Science, vol. 13, no. 1, pp. 28-36, 2001.
[175]
T. L. Starks, L. E. Shubert and F. R. Trainor, "Ecology of soil algae: a review," Phycologia, vol. 20, pp. 65-80, 1981.
[176]
Z. Gao, R. Horton, L. Wang and J. Wen, "An improved forcerestore method for soil temperature prediction," European Journal of Soil Science, vol. 59, p. 972–981, 2008.
[177]
H. Håkansson and V. J. Jones, "The compiled freshwater diatom taxa list for the maritime Antarctic region of the South Shetland and South Orkney Islands," in Proceedings of the Fourth Arctic–Antarctic Diatom Symposium (workshop), Canadian Museum of Nature, Ottawa, Ontario. Canadian Technical Report of Fisheries and Aquatic Sciences No. 1957, 1994, pp. 77-83.
[178]
R. Le Cohu and B. Van de Vijver, "Le genre Diadesmis (Bacillariophyta) dans les archipels de Crozet et de Kerguelen avec la description de cinq ensèces nouvelles," Annuales de Limnologie, vol. 38, pp. 119-132, 2002.
[179]
K. Sabbe and W. Vyverman, "Taxonomy, morphology and ecology of some widespread representatives of the diatom genus Opephora," European Journal of Phycology, vol. 30, pp. 235-249, 1995.
[180]
M. Kahlert, M. G. Kelly, D. G. Mann, F. Rimet, S. Sato, A. Bouchez and F. Keck, "Connecting the morphological and molecular species concepts to facilitate species identification within the genus Fragilaria (Bacillariophyta)," Journal of Phycology, vol. 55, no. 4, pp. 948-970, 2019.