Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint

The article provides an overview of the problem of origin of the only native vascular plants of Antarctica, Deschampsia antartica (Poaceae) and Colobanthus quitensis (Caryophyllaceae), from the viewpoint of modern historical phytogeography and related fields of science.

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Datum:	2007
Hauptverfasser:	Mosyakin, S.L., Bezusko, L.G., Mosyakin, A.S.
Format:	Artikel
Sprache:	English
Veröffentlicht:	Інститут клітинної біології та генетичної інженерії НАН України 2007
Schriftenreihe:	Цитология и генетика
Schlagworte:	Обзорные статьи
Online Zugang:	http://dspace.nbuv.gov.ua/handle/123456789/66595
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spelling	irk-123456789-665952014-07-19T03:01:41Z Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint Mosyakin, S.L. Bezusko, L.G. Mosyakin, A.S. Обзорные статьи The article provides an overview of the problem of origin of the only native vascular plants of Antarctica, Deschampsia antartica (Poaceae) and Colobanthus quitensis (Caryophyllaceae), from the viewpoint of modern historical phytogeography and related fields of science. Дан обзор проблемы происхождения аборигенных сосудистых растений Антарктики Deschampsia antartica (Poaceae) и Colobanthus quitensis (Caryophyllaceae) с точки зрения исторической фитогеографии и родственных направлений науки. Подається огляд проблеми походження єдиних аборигенних судинних рослин Антарктики Deschampsia antarctica (Poaceae) та Colobanthus quitensis (Caryophyllaceae) з точки зору історичної фітогеографії та споріднених напрямків науки. 2007 Article Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint / S.L. Mosyakin, L.G. Bezusko, A.S. Mosyakin // Цитология и генетика. — 2007. — Т. 41, № 5. — С. 54-63. — Бібліогр.: 65 назв. — англ. 0564-3783 http://dspace.nbuv.gov.ua/handle/123456789/66595 74.9 : 574.91 (292.3) en Цитология и генетика Інститут клітинної біології та генетичної інженерії НАН України
institution	Digital Library of Periodicals of National Academy of Sciences of Ukraine
collection	DSpace DC
language	English
topic	Обзорные статьи Обзорные статьи
spellingShingle	Обзорные статьи Обзорные статьи Mosyakin, S.L. Bezusko, L.G. Mosyakin, A.S. Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint Цитология и генетика
description	The article provides an overview of the problem of origin of the only native vascular plants of Antarctica, Deschampsia antartica (Poaceae) and Colobanthus quitensis (Caryophyllaceae), from the viewpoint of modern historical phytogeography and related fields of science.
format	Article
author	Mosyakin, S.L. Bezusko, L.G. Mosyakin, A.S.
author_facet	Mosyakin, S.L. Bezusko, L.G. Mosyakin, A.S.
author_sort	Mosyakin, S.L.
title	Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint
title_short	Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint
title_full	Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint
title_fullStr	Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint
title_full_unstemmed	Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint
title_sort	origins of native vascular plants of antarctica: comments from a historical phytogeography viewpoint
publisher	Інститут клітинної біології та генетичної інженерії НАН України
publishDate	2007
topic_facet	Обзорные статьи
url	http://dspace.nbuv.gov.ua/handle/123456789/66595
citation_txt	Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint / S.L. Mosyakin, L.G. Bezusko, A.S. Mosyakin // Цитология и генетика. — 2007. — Т. 41, № 5. — С. 54-63. — Бібліогр.: 65 назв. — англ.
series	Цитология и генетика
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first_indexed	2025-07-05T16:49:02Z
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fulltext	The article provides an overview of the problem of origin of the only native vascular plants of Antarctica, Deschampsia antartica (Poaceae) and Colobanthus quitensis (Caryophy� llaceae), from the viewpoint of modern historical phytogeog� raphy and related fields of science. Some authors suggested the Tertiary relict status of these plants in Antarctica, while others favour their recent Holocene immigration. Direct data (fossil or molecular genetic ones) for solving this controversy is still lacking. However, there is no convincing evidence sup� porting the Tertiary relict status of these plants in Antarctica. Most probably D. antarctica and C. quitensis migrated to Antarctica in the Holocene or Late Pleistocene (last inter� glacial?) through bird�aided long�distance dispersal. It should be critically tested by (1) appropriate methods of molecular phylogeography, (2) molecular clock methods, if feasible, (3) direct paleobotanical studies, (4) paleoclimatic reconstructions, and (5) comparison with cases of taxa with similar distribution/dispersal patterns. The problem of the ori� gin of Antarctic vascular plants is a perfect model for integra� tion of modern methods of molecular phylogeography and phylogenetics, population biology, paleobiology and paleo� geography for solving a long�standing enigma of historical plant geography and evolution. Introduction This article was provoked by the article by I.Yu. Parnikoza, D.N. Maidanuk, and I.A. Koze� retska [1] recently published in «Cytology and Ge� netics» (Kiev). We cannot completely agree with one of the main conclusions of the authors, who postulated the Tertiary (Oligocene�Pliocene) relict status for both angiosperm taxa in Antarctica, and because of that we provide here some additional information and comments on the topics consid� ered in the article by Parnikoza et al. Due to the tremendous progress in and growing availability of molecular methods, the science of historical biogeography is now undergoing rapid and dramatic transformation which can be regarded as a true «molecular revolution» [2–9]. In fact, instead of vague and non�testable hypotheses and assumptions, we have now in many cases a solid evidence indicating possible centers of origin, migra� tion pathways and timing of evolutionary radiations for many previously enigmatic biogeographical cases concerning many taxa of plants, animals, fungi, and even protists. It means that we see the transformation of a previously empirical field of science into a combination of real experimental and historical science, conclusions of which are falsifiable in the true scientific, Popperian sense. Molecular phylogeographic, phylogenetic, and populational�genetic approaches proved to be the most productive ones in reconstructing the history and development of geographical patterns in plants. The most interesting examples bearing concep� tual implications to our topic are, in our opinion, recent results obtained in biogeography in the fields of studying of classical vicariance and/or dis� persal models, island biogeography (especially cases of dispersal to and evolution on oceanic islands), nothal biogeographical links (especially various ex � planations of nothal disjunctions), Late Pleistoce� ne and Holocene history of the Arctic and boreal biota (especially studies using combined methods of phylogeography and paleobiology), and some others, which we cannot discuss here in detail be� cause of space and time limitations. Of course, it is impossible to cover sufficiently in the present article the vast scientific areas men� tioned above. However, we will provide here some general ideas, striking examples, references to sev� eral most important and useful review articles, and, finally, our comments regarding the current concepts of the origins of Antarctic vascular plants, including the interesting concept proposed by 54 ISSN 0564–3783. Цитология и генетика. 2007. № 5 Обзорные статьи УДК 74.9 : 574.91 (292.3) S.L. MOSYAKIN 1, L.G. BEZUSKO 1, A.S. MOSYAKIN 1,2 1 M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, 2 Tereshchenkivska Street, Kyiv, 01601 Ukraine syst@botany.kiev.ua 2 National University «Kyiv�Mohyla Academy», 2 H. Skovorody Street, Kyiv, 04070 Ukraine ORIGINS OF NATIVE VASCULAR PLANTS OF ANTARCTICA: COMMENTS FROM A HISTORICAL PHYTOGEOGRAPHY VIEWPOINT © S.L. MOSYAKIN, L.G. BEZUSKO, A.S. MOSYAKIN, 2007 Parnikoza et al. [1]. Our brief review is by necessity a broad�stroke picture in which we cannot go into too much detail of the discussed extensive issues. However, we believe that interested readers will consult the references we cite here and get a wider and deeper vision of the important biogeographical and evolutionary problems, which are being suc� cessfully solved with the aid of methods of molecu� lar ecology, phylogeography, population genetics, and other modern approaches. Results and discussion Dispersal and vicariance models in phytogeogra� phy. First of all, it should be emphasized that mod� ern historical biogeography is, by definition, a his� torical and evolutionary science [3, 5]. It means that reconstruction of historical traits in dispersal and distribution of organisms is impossible without ta� king into consideration the evolutionary processes occurring in space and time. Non�evolutionary his� torical biogeography is thus a conceptual nonsense. Vicariance models imply gradual migrations from the centers of origin and further changes of ancestral ranges, with their subsequent splitting in most cases of disjunction. Classical examples of vicariance models are explanations of present�day distribution patterns by past continental movements (mobilism, or modern plate tectonics), orogenesis, transgressions and regressions of seas, shifts of physi� ographic, climatic and biotic zones, etc. On the other hand, dispersal models imply gra� dual or geologically momentary dispersal events, during which organisms or their dispersal units migrate over some physical barriers and become successfully established in a new territory, and such migrations are naturally accompanied and followed by evolutionary transformations. Vicariance�based models were especially fashio� nable after the triumph of the Wegenerian mobilis� tic theory and further development of the modern theory of global plate tectonics. However, vicarian� ce was also a respected concept even long before that. If we consider enigmatic distribution patterns of many plants in the Southern Hemisphere, this concept can be traced back to works of J.D. Hoo� ker. In more detail these models and explanations are discussed in several review articles [4, 5, 10–14] and references therein, which are recom� mended to the reader for further acquaintance with the problem. However, recently the vicariance models, espe� cially those appealing and referring to Gondwanan biotic interactions, fell out of fashion for several reasons, which will be briefly discussed below using several case topics. Insular endemics and long�distance dispersal. A flow of recent molecular phylogenetic and phylo� geographic studies of insular floras and faunas indi� cated in many cases long�distance migration path� ways and showed that suggestions of the relict sta� tus and ancient age of many oceanic endemics are often far from being justified [2, 15–20]. It is espe� cially true for remote oceanic islands and archipel� agos that have never been part of any continent and, consequently, are not suitable for vicariance biogeographical models. The Hawaii is probably the best studied archi� pelago in that respect [21–25]. Some evolutionary links of endemic Hawaiian plants explained by long�distance dispersal are truly amazing. For exa� mple, the endemic Hawaiian woody species of vio� lets (Viola sect. Nosphinium, family Violaceae) which were considered evolutionary «primitive» in fact evolved quite recently (probably in the Middle Pliocene) from the subarctic amphi�Beringian ancestors probably related to the modern herba� ceous species of the polyploid V. langsdorffii Ledeb. aggregate through a long�distance dispersal by birds from Alaska or East Siberia and subsequent explo� sive radiation [21]. We can mention also the African links of Hawaiian Hesperomannia A. Gray (Astera� ceae) [23] and the origin of a morphologically diverse group of several Hawaiian endemic genera of Lamiaceae (Haplostachys Hillebr., Phyllostegia Benth. and Stenogyne Benth.) from North Ameri� can taxa of Stachys L. sensu lato [24]. Other striking examples indicating North American, South Paci� fic, African and Asian sources of recent colonization of the Hawaii are extensively discussed in recent literature [17, 22, 25]. It is especially important to stress that the Hawaiian Islands are in fact a volcanic «conveyor belt» in the Pacific, with a chain of volcanic islands emerging as the crust plates move over the mag� matic «hot spots» in the mantle [22]. It means that the islands themselves are comparatively young, and they are arranged linearly according to their age, from the oldest northwestern islands (e.g., Kauai) having ca. 5.1 million years of history to the youngest southeastern islands (like the island Origins of native vascular plants of Antarctica: comments from a historical ... 55ISSN 0564–3783. Цитология и генетика. 2007. № 5 of Hawaii itself) just ca. 430 thousand years old [22, 25]. Yet this geologically young archipelago houses a tremendous biotic diversity, with many unique endemic taxa of high taxonomic ranks that evolved during just a few millions of years. It is commonly agreed that the Hawaiian biota is in fact a long�distance dispersal biota [16, 25]. In respect to the problem of the initial arrival of D. antarctica and C. quitensis to Antarctica, the examples of oceanic island biotas and colonization of such islands from migration sources located thousands of kilometers away show us that the Drake Passage (ca. 900 km) cannot be regarded as an ultimate barrier for eventual long�distance migrations from southern South America to Maritime Antarctica, even considering such addi� tional obstacles as the Antarctic Polar Front and the Antarctic Circumpolar Current. Of course, the physical obstacles to migration of plants across the Drake Passage probably limited considerably the number of species that actually performed such random migrations. Another problem for migrants is to take a foothold in the new area and success� fully colonize the inhospitable Antarctic shores, which was possible only to hardy plants preadapt� ed to harsh environmental conditions. Thus, prob� ably much more species in fact migrated from time to time across the Drake Passage, but only two species happened to be preadapted to the new con� ditions, which partly explains the scarcity of the Antarctic angiosperm flora. Nothal floristic links: a legacy of Gondwana, migrations from the north, or long�distant dispersal phenomena? Recent studies convincingly demon� strated that many presumably Gondwanan groups in fact attained their present distribution through Laurasian migrations or/and long�distance disper� sal [11–14]. Even Nothofagus, the long�cherished icon of adepts of nothal vicariance models [26, 27], probably experienced long�distance dispersal events in the course of its evolution [28]. However, it does not mean that all Gondwana�based histor� ical�biogeographical explanations should be rejec� ted. In fact, the Nothal biota of the Southern Hemisphere shows a complicated mix of different biogeographical models and patterns: there we can find examples of the real legacy of Gondwana (but only in some ancient enough groups!), southward migrations from the north (so�called boreotropical migrations), and numerous cases of transoceanic long�distant dispersal phenomena [11, 14, 15, 29]. For further discussion see also reviews by Eskov in Russian [12, 13] and Mosyakin in Ukrainian [5] and references therein. The changing views on the history (or histories) of the Nothal floras were reflected also in the recent proposal to modify the system of Takhtajan’s floral kingdoms [30]. In particular, Cox [31] proposed to abandon the Antarctic Floral Kingdom and allo� cate its constituent parts to the neighboring king� doms. Cox justly indicated that «The Antarctic flo� ral Kingdom contains some (but not all) of the remains of a once�continuous southern Gondwa� na cool�temperate flora, now scattered into a relict distribution by the processes of plate tectonics, and present only where the persistence of cool, moist climates has allowed it to survive» [31]. We can add to that that some phytogeographic similarities of various parts of the Antarctic Kingdom are caused not only by plate tectonics events, but also, consid� erably, by long�distance dispersal events. Conse� quently, the Chile�Patagonian Region should be allocated to the Neotropical Kingdom. The Fernan� dezian Region and the Region of the South Sub� antarctic islands should be also placed there because of their long�distance dispersal links to the Neotropis. New Zealand and its surrounding is� lands should be transferred to the Australian Kingdom. Thus, appeals of Parnikoza et al. [1] to publica� tions emphasizing the high past biodiversity of Gondwana and its fragments in the Cretaceous and the Cenozoic has little or no implication to the question when and how modern Deschampsia and Colobanthus migrated to Antarctica. We know that Antarctica in the distant past was a much hospitable place than it is now; probably the great southern continent was the scene of evolution and migration of many important taxa now constituting the Not� hal biota [29, 32–34], but interpretations of these data from the viewpoint of the modern biotic situa� tion should be done carefully. It should be also evi� dent that a find of a presumably caryophyllaceous flower Caryophylloflora paleogenica G. J. Jord. & Macphail in the Middle to Upper Eocene of Tas� mania [35], mentioned by Parnikoza et al. [1], has nothing to do with the present�day distribution of Colobanthus (Caryophyllaceae) in Antarctica. Mo� reover, the taxonomic placement of that Eocene fossil in the family Caryophyllaceae sensu stricto is S.L. Mosyakin, L.G. Bezusko, A.S. Mosyakin 56 ISSN 0564–3783. Цитология и генетика. 2007. № 5 tentative; probably this ancient plant belonged to some group of caryophyllids in a wider sense. Late Pleistocene and Holocene history of the Arc� tic and boreal biota. Arctic, subarctic and Alpine plants are favorite models used in numerous mole� cular phylogeography studies, especially in Euro� pe. Reviews of these studies from the viewpoint of the problem of Pleistocene relicts and refugia and Holocene floral migration routes have been recen� tly published in Ukrainian [6, 7]; many other recent review publications are useful for a better vi� sion of the Late Pleistocene and Holocene history of the Arctic and boreal biota [4, 8, 9, 36–40]. Moreover, it has been shown that, contrary to simplistic views of exclusively gradual «step�by� step» dispersal of most plants [41], the real Late Pleistocene – Early Holocene recolonization of glacial and periglacial areas of Eurasia and North America by plants developed mostly according to the long�distance dispersal scenario [15]. At the same time, hundreds of molecular phylogeography studies and state�of�the�art paleobotanical investi� gations showed that the concept of the «glacial steamroller» (or, better to say in this case, an «ice� roller»?) that exterminated nearly all life in glaciated and adjacent areas is also probably an exaggera� tion. Thus, «tabula rasa» and «survival in situ» mo� dels are not mutually excluding; both these expla� nations work for some taxa and specific areas, de� pending on many factors [4, 6, 7, 9, 37, 38, 40, 42]. No surprise–life is much more diverse than our mental models of it. By analogy, comparing the available biogeogra� phic cases of Arctic and Antarctic vascular plants, we can assume that (1) survival of Deschampsia and Colobanthus in Antarctica in situ since pre�Pleis� tocene times has a very low probability, (2) their Holocene migration (or probably even several migration events) to Antarctica is the most feasible explanation, and (3) their survival in situ during the Last Glacial Maximum (LGM) since one of interglacials is less probable, but not excluded. Origins of native vascular plants of Antarctica: still in the mist. Here we will try to show some faults in discussion of Parnikoza et al. [1] when they attempt to prove the relict status of D. antarctica and C. quitensis in Antarctica and to pinpoint the age of their migrations. According to Parnikoza et al. [1], D. antarctica and C. quitensis migrated to Antarctica «during the Oligocene�Pliocene», when the southern continent was less isolated and its cli� mate was more favorable for naturalization of these taxa. Probably it was indeed less isolated and more favorable, but how is it related to the actual time of immigration of the two species in question? In the Cretaceous Antarctica was even less isolated from other Gondwana fragments and climatically more favorable than it was in the Oligocene�Pliocene and, judging from molecular�clock�based and fos� sil�calibrated age estimates, orders Caryophyllales and Poales, and probably even phylogenetically basal representatives of the families Caryophy� llaceae and Poaceae, have differentiated already in the Late Cretaceous [10, 43, 44]. Should we becau� se of that assume that Colobanthus and Descha� mpsia migrated to Antarctica already in the Late Cretaceous? Or probably in the Paleocene? Is there anybody voting for the Eocene? In the Oligocene palms were growing in Ukraine, but it does not necessarily mean that palms currently grown in Crimea are relicts and direct descendants of those Ukrainian Oligocene palms. Sometimes researchers (usually except geolo� gists and paleontologists) have problems with feel� ing the vastness of the geological timescale. It is simply nonsensical, from geological and paleobio� logical viewpoints, to guess as a migration age the age limit covering about 32 million years, or roughly a half of the whole Cenozoic [1]. The Early Oligocene started ca. 34 million years ago (Mya), while the Pliocene�Pleistocene boundary is cur� rently placed at ca. 1.8 Mya [45]. Moreover, it is hard (in fact, impossible) to believe that Antarctic populations of both species, D. antarctica and C. quitensis, remained unchanged since the Oligocene, Miocene, or even Pliocene, and developed no visible morphological or considera� ble genetic distinctions from their relatives on the South American continent. That notion simply denies evolution, adaptive or neutral. Parnikoza et al. [1] properly cited the results of Holderegger et al. [46], who found that Antarctic populations of D. antarctica show low genetic diversity. That fact may have several explanations, including the following most obvious ones: (1) Antarctic D. antarctica and C. quitensis are recent migrants that originated from limited founder stocks; the descendants of the founding popula� tions simply had no time for genetic differentiation in Antarctica (the most parsimonious explanation, Origins of native vascular plants of Antarctica: comments from a historical ... 57ISSN 0564–3783. Цитология и генетика. 2007. № 5 by the way); and (2) there is a constant gene flow and/or migrations between the isolated Antarctic island populations of D. antarctica and C. quitensis, mixing their gene pools to the level of «low genet� ic diversity» [46]. If these species are capable of self�pollination and/or asexual reproduction (and they are!), then the assumption of their long�term isolation in combination with a genetic «melting pot» simply loses ground. Let us consider just two examples: Chenopodium tomentosum Thouars (Chenopodiaceae) [47] and Rumex frutescens Thouars (Polygonaceae) [48, 49], both endemic to remote South Atlantic islands of the Tristan da Cunha (Tristan d’Acugna) group (ca. 2800 km from Africa and ca. 3200 km from the South American mainland), both evidently resulted from recent and single long�distance dis� persal events, both having very close relatives in South America (taxa of the Chenopodium ambro� sioides L. aggregate in the first case, and Rumex cuneifolius Campderá and other representatives of the predominantly South American subsection Cuneifolii Rechinger f. in the second), but still dif� fering from them to the species or at least sub� species level. Why Antarctic plants do not differ, specifically or at least subspecifically or varietally, from their South American conspecific relatives, if they parted several million years ago? Chwedorzewska [50], baseding on AFLP ana� lysis, reported that genetic diversity within the Antarctic populations of D. antarctica was greater than respective genetic diversity values within the analyzed Arctic populations of D. brevifolia R. Br. and D. alpina (L.) Roem. & Schult. sampled in Svalbard (Spitsbergen). Moreover, southern popu� lations of D. antarctica revealed less diversity than northern populations living in less harsh condi� tions. These preliminary data can be interpreted in several ways. As we have seen from other examples (see above), low genetic diversity values within a species in a particular area may indicate a recent arrival of the species to that area and the founder effect. On the other hand, in the case of D. antarc� tica even a high genetic diversity can be interpret� ed as either (1) in situ genetic differentiation in iso� lated fragmented habitats and gradual but limited dispersal «by step�stones» or (2) a result of several independent long�distance migration events. There is also another option involving a bottleneck effect, when large portions of a population were eliminat� ed due to high selective pressure of environmental conditions and/or random non�selective cata� strophic events, such as advance of glaciers, sea level oscillations, and extreme climatic episodes. A reliable answer favoring any of these options can be obtained only through phylogenetic and phylogeo� graphic studies involving wide�scale sampling of the species from all (or most of) known range frag� ments in Antarctica and many representative sam� ples from South America. Recent ITS analysis of C. quitensis demonstrated a relatively high genetic similarity among the studied Andean and Antarctic populations (sequence diver� gence = 1.17 %) despite the considerable geogra� phical distance (> 3300 km) [51]. It should be also noted that C. quitensis is a selfing species, which is also capable of asexual reproduction. Of course, there was considerable ecotypic differentiation re� vealed, which should be expected in plants inhabit� ing so extreme and geographically isolated habitats differing considerably in local ecological conditions. Gianoli et al. [51] quite logically assume Andean origin of Antarctic populations of Colobanthus and its recent dispersal by migratory birds. Of course, we agree with Parnikoza et al. [1] that the ITS region sequences are probably not good markers of recent microevolutionary changes in Colobanthus, and other methods should be applied instead of or in addition to ITS phylogeny. However, before obtaining new data it would be safer to stick to the most parsimonious explanations. If, as we have seen above, less than 5 million years was enough time in the Hawaii for evolution of endemic genera and spectacular evolutionary radiation of diverse species groups having in many cases just one recent ancestor per group that ar� rived by long�distance dispersal, if very recent migrants from South America to the islands of Tristan da Cunha were able to form distinct species differing from their South American relatives, then why, if we assume the «Oligocene�Pliocene» mig� ration of Deschampsia and Colobanthus to Antarc� tica, these Antarctic plants developed no distinc� tions from their South American ancestors? Of course, one may say that tropical islands are prob� ably more stimulating for evolutionary changes than the harsh Antarctic environment is. However, numerous examples demonstrate that in fact an escape from biotic competition enables dramatic island radiations, while abiotic environmental S.L. Mosyakin, L.G. Bezusko, A.S. Mosyakin 58 ISSN 0564–3783. Цитология и генетика. 2007. № 5 stress is the leading factor promoting such radia� tions. Consequently, the amazing evolutionary conservatism indirectly implied by the assumption of the «Oligocene�Pliocene» origin of the two Antarctic species is simply inexplicable. A recent molecular study of phylogenetic rela� tionships of Deschampsia antarctica using ITS shed little light to the problem. In the trees obtained by Fernández Souto et al. [52] D. antarctica was grouped with five other Deschampsia species, but showed out in three different positions: (1) as a clade sister to all other Deschampsia species (ex� cluding D. flexuosa (L.) Trin.), (2) as a sister group of D. cespitosa (L.) P. Beauv. – D. alpina (L.) Roem. & Schult. – D. sukatschewii (Popl.) Roshev., and (3) as sister to D. mejlandii C.E. Hubb. – D. christo� phersenii C.E. Hubb. In plain English it just means that the authors failed to pinpoint the phylogenet� ic position of D. antarctica, probably because of limitations of the methods or sequences used, or be� cause of inadequate taxon sampling. Fernández Souto et al. [52] seem to be surprised by the revealed fact that «Deschampsia does not appear monophyletic as D. flexuosa is not included in this clade». It probably escaped their attention that D. flexuosa has been since long ago considered genef� ically distinct from Deschampsia s. str. and trans� ferred into segregate genera, either Lerchenfeldia Schur (as L. flexuosa (L.) Schur) or Avenella Parl. (as A. flexuosa (L.) Drejer), and is currently treat� ed taxonomically as a member of Avenella. This little example shows that some experience in tradi� tional taxonomy is not unneeded even for molecu� lar taxonomic studies. Fascinating finds of a rich (of course, rich by Antarctic standards) fossil Neogene flora of the Meyer Desert Formation (the biostratigraphic age less than 3.8 Ma, which means an Early Pliocene or more recent age) in the Transantarctic Moun� tains prove that continental Antarctic in the Late Neogene was more suitable for terrestrial life than most people expected [32]. However, this fact has no bearing to proving the relict status of Colobanthus and Deschampsia in their present Antarctic ranges, as well as presence of broadleaf trees in boreal areas of Europe during interglacials does not nec� essarily mean that these trees survived in situ dur� ing glacial phases until the present day. Glacial and climatic history of Antarctica should be also considered in detail before making any sug� gestions on refugia of Antarctic vascular plants. Large�scale deglaciation in Antarctic coastal areas started with a general warming trend after 8.4 ka (thousand years ago). Before that many now ice� free Antarctic areas were glaciated [53–57]. It is evident from a multitude of sources that the global climatic history of the Late Pleistocene and Early Holocene was rather complex. The warming trend of the last deglaciation was interrupted by the Younger Dryas event that lasted for almost 1200 years (12.7–11.5 ka) and resulted in a reverse to almost glacial conditions. A short cold event occurred also around 8 ka. For example, the deglaciation on South Georgia commenced prior to 18.6 ka; colder conditions returned after 14 ka and lasted during the Younger Dryas (12.7–11.5 ka) without significant changes, while the transition to postglacial conditions occurred between 8.4 and 6.5 ka and was inter� rupted by a cold event that began ca. 7.8 ka and lasted for ca. 400 years [58]. It is highly improba� ble that Deschampsia and Colobanthus survived such dramatic events in situ without any response, e.g. reductions of their ranges. We should also not forget about eustatic sea level changes that reshaped and remodelled the coastal Antarctic areas and their biotas [59] and, conse� quently, influenced the potential and actual habi� tats of Deschampsia and Colobanthus; that makes their preservation in situ during diverse and dra� matic Pleistocene events even more improbable. However, a suggestion of survival of D. antarc� tica and/or C. quitensis during the LGM is no heresy. It might well be the case that these two angiosperms indeed survived the LGM in Antarctic islands, but it should be proved. In fact, the present climatic situation in Antarctica is rather unusual as compared to most periods of the Pleistocene, cor� responding roughly to interglacial conditions [53, 57], especially if compared to the Last Glacial Maximum, both in the Northern [60] and Southern [55, 59] hemispheres. Some range expansions of several Antarctic species southward were observed recently; it is usually viewed as a response to glob� al warming, although other causes, like normal cli� matic oscillations, should be also considered. Anyway, the present�day climatic conditions in Antarctica are much more favorable to plants than conditions there during the Last Glacial Maximum or other glacial maxima of the Pleistocene. Please Origins of native vascular plants of Antarctica: comments from a historical ... 59ISSN 0564–3783. Цитология и генетика. 2007. № 5 note that D. antarctica and C. quitensis (the first species being more widespread than the second) occupy now only a comparatively narrow northern strip of Maritime Antarctica, and these plants are already living virtually on the brink of survival. These considerations make the ideas of in situ sur� vival of D. antarctica and C. quitensis in Antarctica since the Paleogene or even since the Early Pleis� tocene rather problematic. Endemism and distribution patterns of various groups of plants and fungi (including lichens) occur� ring in Antarctica indicate that the Pleistocene survival was possible for some lichens, less proba� ble for mosses [61], and rather improbable for the two considered species of vascular plants [see dis� cussion in 62]. That was already evident to biogeo� graphers of the first half of the 20th century. For example, consider discussions in the classical books by Wulff [63, 64], who often favored vicari� ance�based mobilistic historical explanations of plant distribution patterns, but did not extend such explanations to the two extant native species of Antarctic angiosperms. The reasons of the scarcity of the vascular flora of Antarctica should be also reconsidered from the viewpoint of both ecological and migrational factors. As Aleksandrova [65] cor� rectly noted, the presence of just two species of flowering plants in Antarctica cannot be explained by extreme ecological factors only, because the ecological conditions for plants there are not worse than, for example, on the Franz Josef Land in the Russian Arctic. Consequently, we should consider such additional explanations as migrational obsta� cles and isolation of the region. Indeed, solid evidence would be definitely need� ed for supporting the idea of the pre�Pleistocene age of Colobanthus and Deschampsia in Antarctica, but, as we see, no such evidence is available yet, while the bulk of both direct and indirect available data discussed in the present article speak in favor of the Holocene or, at best, Late Pleistocene age of the present�day Antarctic flowering plants. Conclusions The suggestion of Parnikoza et al. [1] of an Oligocene�Pliocene origin of Deschampsia antarc� tica Desv. and Colobanthus quitensis (Kunth) Bartl. is most probably a gross overestimation. No data in their article, either literature or original, give con� vincing evidence supporting their concept of the Tertiary age and relict status of these plants in Antarctica. It might have been better to critically assess an ample set of available evidence from clas� sical historical biogeography, phylogeography, paleobotany, paleoclimatology and some other fields, not necessarily data regarding the two taxa and the territory considered, but also (and mainly) comparative data regarding other taxa and other areas with similar phytogeographical traits. Judging from both direct and indirect evidence and comparison with biogeographical analogues, the most reasonable and consistent with facts sug� gestion would be that of the Holocene or Late Pleistocene (last interglacial?) age of migration of the ancestral stock of D. antarctica and C. quitensis to Antarctica through bird�aided long�distance dis� persal events. Thus, these species (or one of them) are either Holocene migrants or, at best, relicts of a recent intergalical. In the last case they probably survived the Last Glacial Maximum (LGM) in coastal refugia in Maritime Antarctica or adjacent islands. However, these age estimations should be critically tested by (1) appropriate methods of molecular phylogeography involving extensively sampled plants from Antarctica and South Ame� rica, (2) molecular clock methods, if feasible, (3) direct paleobotanical (including paleopalynologi� cal) studies, (4) paleoclimatic reconstructions, and (5) comparison with similar cases documented for taxa with similar distribution/dispersal patterns. On� ly in these directions lies the positive and reliable answer to the long�intriguing question of the ori� gins of D. antarctica and C. quitensis in Antarctica. The problem of the origin of Antarctic vascular plants is a perfect model for integration of modern methods of molecular phylogeography and phylo� genetics, population biology, paleobiology and paleogeography for solving the long�standing enig� ma of historical phytogeography. РЕЗЮМЕ. Дан обзор проблемы происхождения аборигенных сосудистых растений Антарктики Des� champsia antartica (Poaceae) и Colobanthus quitensis (Caryophyllaceae) с точки зрения исторической фито� географии и родственных направлений науки. Неко� торые авторы считают, что эти растения в Антарктике являются реликтами третичных времен, а другие ис� следователи склоняются к концепции их недавней го� лоценовой миграции. Прямых данных (как ископае� мых, так и молекулярно�генетических) для решения этой проблемы пока что не хватает. Тем не менее, нет S.L. Mosyakin, L.G. Bezusko, A.S. Mosyakin 60 ISSN 0564–3783. Цитология и генетика. 2007. № 5 убедительного подтверждения третичного реликтового статуса этих растений в Антарктике. Вероятнее всего, D. antarctica и C. quitensis мигрировали в Антарктику в голоцене или позднем плейстоцене путем расселения на дальние расстояния с помощью птиц. Эта концеп� ция должна быть критически проверена с помощью соответствующих методов молекулярной филогеогра� фии, методов молекулярных часов (при возможнос� ти), непосредственных палеоботанических исследова� ний, палеоклиматических реконструкций и сравнения с таксонами, имеющими аналогичные особенности распространения или расселения. Проблема происхож� дения антарктических сосудистых растений является замечательной моделью для интеграции современных методов молекулярной филогеографии и филогенети� ки, популяционной биологии, палеобиологии и палео� географии для решения давней загадки исторической географии и эволюции растений. РЕЗЮМЕ. Подається огляд проблеми походження єдиних аборигенних судинних рослин Антарктики Deschampsia antarctica (Poaceae) та Colobanthus quitensis (Caryophyllaceae) з точки зору історичної фітогеогра� фії та споріднених напрямків науки. Деякі автори вва� жають, що ці рослини в Антарктиці є реліктами тре� тинних часів, а інші дослідники схиляються до кон� цепції їх недавньої голоценової міграції. Прямих да� них (як викопних, так і молекулярно�генетичних) для вирішення цієї проблеми поки що не вистачає. Проте, третинний реліктовий статус цих рослин в Антарктиці не має переконливого підтвердження. Цілком імовір� но, що D. antarctica та C. quitensis мігрували до Антарк� тики у голоцені або пізньому плейстоцені завдяки розселенню на далекі відстані за допомогою птахів. Ця концепція має бути критично перевірена за допо� могою відповідних методів молекулярної філогеогра� фії, методів молекулярного годинника (при можли� вості), безпосередніх палеоботанічних досліджень, па� леокліматичних реконструкцій та порівняння з таксо� нами, які мають аналогічні особливості поширення або розселення. 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Origins of native vascular plants of Antarctica: comments from a historical phytogeography viewpoint

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