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Eliömaantiede: kasvimaantiedeIII Jääkausi ja Suomen kasvisto
Jari Oksanen
Oulun yliopisto
SL 2015
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 1 / 76
Neutraali malli Jääkausi
Jääkausi
Pohjoisen Euroopan kasvillisuus tuhoutui kokonaan jään alle jajäätiköitymättömilläkin alueilla tapahtui suuria mullistuksia. Kaikki Suomenkasvit ovat saapuneet Suomeen jääkauden jälkeen, esimerkiksi kuusi jopapaljon ihmisen jälkeen. Jäätiköityminen on lisäksi vaikuttanut maaperänrakenteeseen ja muihin ympäristöoloihin pitkään jääkauden jälkeenkin.
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 2 / 76
Neutraali malli Jääkausi
Jääkausien aika
Pleistoseeni eli viimeiset 2 milj. vuotta: jatkuvaa jääkausien ja niidenvälikausien vuorotteluaInterglasiaali: jääkauden lämpimämpiä välikausiaInterstadiaalit: lyhyt tai viileä välijakso
Jäätikön ’vetäytyminen’ näkyy interstadiaalina lauhkeilla vyöhykkeillä
Viimeisin jääkausi 115 000 . . . 10 000 v sitten
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 3 / 76
Neutraali malli Jääkausi
Milankovićin syklitМилутин Миланковић, 1879 – 1958
Maan radan soikeus vaihtelee 100 000 v sykleissäMaan kiertoakselin pystyys vaihtelee 41 000 v sykleissäAika jolloin maa on lähinnä aurinkoa (perihelion) heilahtelee 21 000 vsykleissäNäiden syklien yhteisvaikutus aiheuttaa nykyiset jääkaudet ja niidenvälikaudetNyt maan akseli on 23.5◦ kulmassa, perihelion on tammikuussa9 000 v sitten akseli oli asteen vinompi ja perihelion oli heinäkuussa:auringon säteily oli heinäkuussa 8% voimakkaampi ja tammikuun 8%heikompi 45 leveysasteella kuin nyt
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 4 / 76
Neutraali malli Jääkausi
Lähimenneisyys
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 5 / 76
Neutraali malli Jääkausi
Viimeisin jääkausi
et al., 2000, 2002; Gataullin et al., 2001; Stein et al.,2002). It is assumed that the southern flank of the icesheet coalesced with the Scandianvian Ice Sheet near thenorthern tip of the Kanin Peninsula, but the exactboundary remains to be defined in the confluence zone(Fig. 16). In the Pechora Sea, further to the east, webelieve the ice front reached the Kolguev Line, whichdepicts the southern extent of the Late Weichselian tillon the Barents Sea shelf (Gataullin et al., 2001). In thesouthern Kara Sea the ice front was probably locatedalong the southern and eastern margin of the NovayaZemlya Through (Svendsen et al., 1999, 2004; Polyaket al., 2000). Further to the north the LGM limit hasbeen mapped across the shallow shelf near the southernslopes of the St. Anna Through (Polyak et al., 2002;Stein et al., 2002). In general, the area on the proximalside of the inferred ice sheet limit is characterized by arough morainic relief, whereas on the distal side theseafloor is graded and in places underlain by thicksequences of marine sediments not covered by till.Furthermore, incised river channels and the widespreadoccurrence of permafrost indicate that this area has been
subaerially exposed at a time when the sea level wasmuch lower than today.
Based on the record from Severnaya Zemlya it seemsclear that the Barents-Kara Ice Sheet did not reach thisarchipelago and that local glaciers were probably evensmaller than today (Makeyev et al., 1979; Bolshiyanovand Makeyev, 1995; Raab et al., 2003). Our investiga-tions on the mainland suggest, however, that the NWcoast of Taimyr was affected by a Late Weichselian iceadvance directed from the continental shelf (Alexander-son et al., 2002). One possible interpretation is that theadvancing ice formed part of the Barents-Kara IceSheet. In that case the ice sheet must have blocked thenorthbound drainage in West Siberia, as well as in theEuropean part of the Russian mainland (Fig. 16). Asdiscussed previously, however, aeolian sedimentationand growth of ice wedges during this time span excludethe possibiliy that a pro-glacial lake flooded the lowlandareas along the Arctic coastline. Possibly the erodedchannels on the Kara Sea shelf (Stein et al., 2002)contained rivers that flowed towards the Arctic Oceanduring most of the LGM period. The ice advance that
ARTICLE IN PRESS
Fig. 16. A reconstruction of the Eurasian ice sheets at the Late Weichselian glacial maximum (LGM). The eastern limit of the Scandinavian Ice Sheetand the southern and eastern limit of the Barents-Kara Ice Sheet are to a large extent based on our own investigations. The northern and western icesheet limit on the Barents Sea shelf is drawn according to the reconstruction by Landvik et al. (1998). The ice limits for the rest of Europe are takenfrom various sources (e.g. Ehlers and Gibbard, 2004). The presence of restricted valley glaciers on the Putorana Mountains is indicated by hatchedlines. Notice that the glacier distribution over Iceland, Greenland, Alps and other mountain areas are not shown on this reconstruction.
J.I. Svendsen et al. / Quaternary Science Reviews 23 (2004) 1229–1271 1257
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 6 / 76
Neutraali malli Jääkausi
Jääkausien aikaan
Kasvillisuustyypit ”siirtyivät etelään” jäätikön edestäIlmasto aridimpi: aroja ja aavikoita ennemmän, sademetsätpirstoutuivat
Jäätikön edustalla ’tundra-aro’Suurten herbivorien aiheuttama? (Sergei Zimov, Зимов СергейАфанасьевич)http://www.pleistocenepark.ru/en/
Yhtenäiset kasvillisuusvyöhykkeet pirstoutuivatRefugiot: Skandinavia? Amazon?
Havaittu myös kosteampia jaksoja (’pluviaali’)Osa lajeista hävisi sukupuuttoon, ainakin mantereilta
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 7 / 76
Neutraali malli Jääkausi
Vyöhykkeet siirtyvät etelään?
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 8 / 76
Neutraali malli Jääkausi
Vyöhykkeet siirtyvät . . . etelään?
Fig. 2. Simulated aNPP d 0 ka BP. Simulated aNPP values for eleven aggregate PFTs. Shading indicates the aNPP level, with white areas having zero aNPP for the PFT and shadedareas having non-zero aNPP; highest aNPP values are indicated by yelloweorange colours as indicated in the legend to the figure.
J.R.M. Allen et al. / Quaternary Science Reviews 29 (2010) 2604e2618 2609
Fig. 5. Simulated annual net primary productivity d 21 ka BP. For explanation see Fig. 2.
J.R.M. Allen et al. / Quaternary Science Reviews 29 (2010) 2604e26182612
JRM Allen et al. Quaternary Science Reviews 29 (2010) 2604–2618.Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 9 / 76
Neutraali malli Jääkausi
Ephedra distachya: tundra-aron relikti?
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 10 / 76
Neutraali malli Jääkausi
Mammuttiaro?Zimov SA et al. American Naturalist 146, 765–794; 1995
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 12 / 76
Neutraali malli Jääkausi
Jääkautiset kasvillisuusvyöhykkeet
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 13 / 76
Neutraali malli Jääkausi
Jääkaudesta toipuminen alkaa
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 14 / 76
Neutraali malli Jääkausi
Melkein nykyiset vyöhykkeet
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 15 / 76
Neutraali malli Jääkausi
Jääkaudesta toipuminen Suomessa
Varhaiset lajit arktisiaPreboreaalikausi 8 000–6 800 eaa:Koivuvaltaisia metsiä, jalojalehtipuitaBoreaalikausi 6 800–5 500 eaa:kuivia, mäntyvaltaisia metsiäAtlanttinen kausi 5 500–2 500 eaa:lämmin ja kostea, kuusi saapuuSubboreaalinen (2 500–500 eaa) jasubatlanttinen (500 eaa —): ilmastoviilenee ja mantereistuu, kuusi leviää
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 16 / 76
Neutraali malli Jääkausi
Kuusen invaasio
Rise of the boreal ecosystem in Fennoscandia
631
© 2009 The Authors. Journal compilation © 2009 British Ecological Society,
Journal of Ecology
,
97
, 629–640
each pollen type by the ratio of the added and counted marker grainsand the PAR values were calculated from the pollen concentrationvalues by multiplying the concentration of each subsample by thesedimentation rate (Davis & Deevey 1964).
The modern PAR values were defined for eight sites in Fennoscandiato explore the correlation of the PAR values and the three population
sizes (Table 1). The value for Lake Orijärvi was defined by cal-culating the average PAR of the 14 pollen samples dating from the20th century. The rest of the sites have a lower number of pollen samplesanalysed from the top sediment section and their modern PARvalues were defined as the average of the last 150 years. This timeperiod was chosen due to the fact that the age–depth model used for
Fig. 1. The study area and the modernabundance pattern (volume m3 ha–1) of Piceaabies in Fennoscandia. The isolines indicatethe approximate Holocene spread patternof P. abies from eastern Finland to westernSweden with 1000 year. intervals (adoptedfrom Giesecke & Bennett 2004). The study sitesare marked with dots. A = Lake Kirkkolampi,B = Lake Orijärvi, C = Lake Laihalampi,D = Lake Nautajärvi, E = Lake Klotjärnen.The P. abies volume map is reproduced andpublished with the permission of copyrightholders, the Finnish National Forest Inventory,the Norwegian National Forest Inventoryand the Swedish National Forest Inventory.
Table 1. The modern PAR (grains cm–2 year–1) and biomass values of Picea abies in Finland and Sweden, at sites ranging from P. abies-dominated southern boreal forest to beyond the northern distribution limit of P. abies in northern Fennoscandia. The modern P. abies above-ground biomass values (t ha–1) were obtained from the National Forestry Inventory Statistics and are shown for zones ranging from 500 to4500 m measured from the shores of the lakes (Seppä et al. 2009). The modern PAR value is defined as the average of the last 150 years
SiteLatitude (North)
Longitude (East)
Size (ha)
Modern PAR
Modern biomass 500 m range
Modern biomass 1000 m range
Modern biomass 4500 m range
Laihalampi 61°29′18″ 26°05′ 25.5 732 30,02 30,10 43,50Nautajärvi 61°48′ 24°41′ 18.9 791 29,54 33,36 32,02Kirkkolampi 61°47′ 30°00′ 72 675 No data No data No dataOrijärvi 61°40′ 27°14′ 25.9 690 10,89 21,25 21,75Klotjärnen 61°49′ 16°32′ 2 612 No data No data No dataAkuvaara 67°07′ 27°41′ 4 29 0 0 0Tsuolbmajavri 68°41′ 22°05′ 14 8 0 0 0Toskaljavri 69°12′ 21°28′ 100 5 0 0 0
Seppä
etal.,
JEco
l97,62
9–640(2009)
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 17 / 76
Neutraali malli Jääkausi
Synkät mustat metsät
Rise of the boreal ecosystem
in Fennoscandia633
© 2009 T
he Authors. Journal com
pilation © 2009 B
ritish Ecological Society, Journal of E
cology, 97, 629–640 Fig. 2. The general PAR values of Picea abies at the five study sites, indicated on the right-hand side of the y-axis. The sites are arranged along the direction of the spread from the right (east) to the left (west).The PAR values of Tilia cordata are shown on the left-hand side of the y-axis to reflect its population dynamics during and after the colonization process of P. abies in Fennoscandia. The arrows indicatewhen the P. abies PAR reached the value comparable with the modern at each site and the light and dark vertical lines point to the periods of maximum P. abies and T. cordata populations. Two curvesindicating the Holocene climate trends are shown on the left. The NGRIP δ18O record is derived from an ice core in central Greenland and indicates a general temperature pattern in the North Atlantic region(Johnsen et al. 2001). Other temperature records confirm that this curve in general is valid for northern Europe as well. The southern Swedish δ18O record is obtained from isotopic composition of acalcareous lake sediment core and reflects main trends both in temperature and hydrological conditions (Hammarlund et al. 2003; Seppä et al. 2005). HTM denotes the approximate end of the warm, dryand relatively stable mid-Holocene period in northern Europe.
Seppä et al., J Ecol 97, 629–640 (2009)
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 18 / 76
Neutraali malli Jääkausi
Jääkauden geologiset jäljet
Jää kuluttaa kalliota ja kuljettaa eteenpäin irtainta maata jasiirtolohkareita
Moreeni ja moreenimuodostumat: drumliinit, reunamoreenitLajittunut aines jäätikköjokiin: harjut, deltat, sandurit
Jäätikön ’vetäytymisen’ jälkeen veden pinta korkeallaRavinteet eivät huuhtoutuneet supra-akvaattisilta alueilta: vaaranlaetRavinteikkaita mereisiä savisedimenttejä rannikolle
Jäänjakajan alueella maaperä vanhaa: rapautunut, jopa lateritisoitunut
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 19 / 76
Neutraali malli Suomen kasvisto
Suomen kasviston elementit
Kasvit ovat saapuneet Suomeen jääkauden jälkeen. Ne ovat kehittyneetmuualla ja talvehtineet jäätiköitymättömien alueiden ulkopuolella. Kasvitovat tulleet Suomeen kaikista ilmansuunnista – jopa pohjoisesta.Leviämistien ja nykyisen esiintymisen mukaan voidaan muodostaakasvistollisia ryhmiä.
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 20 / 76
Neutraali malli Suomen kasvisto
Lajistolliset alueet
Panbiogeografia: lajistonryhmittäminen nykyisenlevikin perusteellaIlmasto ja/tai historia:skaalasta riippuenFennoskandian floora ∼kasvillisuusvyöhykkeetEtelä jakaantuu syväänetelään ja eteläsuomalaiseenvyöhykkeeseen, pohjoinenSkandinavianvuoristoalueeseen(Kölivuoristo) jaalavampaan Lappiin
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 21 / 76
Neutraali malli Suomen kasvisto
Kasvien levikkityypit: panbiogeografia
Tavallisin tyyppi: Koko maaIlmastolliset vyöhykkeet:
Lounaiset (eteläiset) lehto- ja ketolajitPohjoiset tunturikasvitItäiset taigalajit: ”vain Suomessa”
Erikoiset habitaatit: disjunktioVaateliaat lehto-, serpentiini- ja kalkkilajitKäsivarsi näkyvin
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 22 / 76
Neutraali malli Suomen kasvisto
Elementae florae fennicae I
Ottavat huomioon kasvien koko levikin, myös Suomen ulkopuolella
Kasvien leviämistie Suomeen: sama levikki, mutta laji voi olla itäinen(Kannaksen kautta) tai läntinen (Ahvenanmaan kautta)
Usein hyvin spekulatiivinen: alkuperää ja reittiä ei tiedetä
Spekulaation perustana myös autekologiset vaatimukset
Aarno Kalela (1961) ja Olov Gjærevoll (1973)
Seuraavian karttojen data http: // www. gbif. org/ , ohjelmallinen yhteys R-paketti spocckarttaprojektio Lambert Azimuthal Equal Area
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 23 / 76
Neutraali malli Suomen kasvisto
Elementae florae fennicae II
Läntiset lajit: suosivat mereistä ilmastoaÄärimmäisen oseaaniset vainLänsi-NorjassaSuboseaanisia myös Suomessa (10):Blechnum spicant, Myrica gale
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Hymenophyllum wilsonii
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 24 / 76
Neutraali malli Suomen kasvisto
Elementae florae fennicae III
Eteläiset, suboseaaniset lajithemiboreaalisessa Suomessa (200)
Lathraea squamaria, Mercurialis perennis,Primula veris, Ranunculus bulbosusLounaisia lehtolajeja
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Ranunculus bulbosus
Mantereis-eteläiset lajit: pääalue Venäjälläja Länsi-Siperiassa, meillä eteläboreaalisia
Spergula morisonii, Filipendula vulgaris,Poa compressa, Hepatica nobilis,Calamagrostis arundinacea
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Calamagrostis arundinacea
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 25 / 76
Neutraali malli Suomen kasvisto
Elementae florae fennicae IV
Itäiset lajit (100): Venäjäntaigavyöhykkeeltä
Picea abies,Chamaedaphnecalyculata, Carexglobularis
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Carex globularis
Boreaalis-hemiarktinen ryhmä: myösKeski-Euroopan vuoristoissa ja arktisillaalueilla
Andromeda polifolia, Betula nana,Empetrum nigrum ssp. hermaphroditum
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Petasites frigidus
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 26 / 76
Neutraali malli Suomen kasvisto
Elementae florae fennicae V
Tunturikasvit: puurajan yläpuolellaHeterogeeninen habitaattiryhmäKalkkialustan lajit Suomessa harvinaisia jadisjunktiivisia, Kölivuoristossa yleisiä
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Cryptogramma crispa
Merenrantakasvit: habitaattityyppiElementtinä heterogeeninen: eteläisiä (Crambe maritima), arktisia(Puccinellia phryganodes), endeemisiä (Deschampsia bottnica)
’Indifferentit’: ihmisen seuralaisia
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 27 / 76
Neutraali malli Suomen kasvisto
Euroopan kasvistoelementitFinnie TJR et al., J. Biogeogr. 34, 1848–1872; 2007
Numeerinen analyysi Atlas Florae Europaeae, 4420 50× 50 km ruutua,2762 lajia (20% Euroopan lajistosta)20 floristista elementtiä, nimetty tyypillisen lajin mukaanJäätiköityneiden alueiden metsät: boreaaliset (Equisetum sylvaticum),keskieurooppalaiset (Lychnis flos-cuculi, nyk. Silene) jalänsieurooppalaiset (Ranunculus bulbosus)Arktis(-alpiiniset) lajit: Oxyria digynaAtlanttiset merenrannat: Cochlearia danicaVuoristomassiivit: Pyreneet (Erysimum duriaei), Alpit (Salixserpillifolia), Karpaatit ja Tatra (Rumex alpinus)Välimeren alue ei floristisesti yhtenäinen vaan erittäin monta erillistäelementtiäKaakkoiseurooppalaiset ja itäiset: Unkari – Bulgaria (Alyssumalyssoides), Bulgaria (Dianthus moesiacus), Krim (Dianthus capitatus)
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 29 / 76
Neutraali malli Suomen kasvisto
Euroopan kasviston elementit
Equisetum sylvaticum
Ranunculus bulbosus
Lychnis flos−cuculi
Oxyria digyna
Erysimum duriaei
Salix serpillifolia Rumex alpinus
Cochlearia danica
Alyssum alyssoides
Dianthus moesiacus
Dianthus capitatus
Osyris alba
Silene scabrifolia
Sarcocapnos enneaphyllaLobularia maritima
Ostrya carpinifolia
Ranunculus psilostachys
Brassica cretica
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 30 / 76
Neutraali malli Suomen kasvisto
Kasvistoelementin levikkiOxyria digyna -ryhmä
extending into adjacent regions (e.g. Dianthus monspessul-
anus, Erucastrum nasturtiifolium, Saponaria ocymoides) or
have distributions in this area with outliers elsewhere. In
extreme cases there are outlying populations in the British
Isles (Arabis scabra, Meconopsis cambrica, Saxifraga hirsuta),
Greece (Bufonia paniculata, Paronychia polygonifolia) or even
the Crimea (Iberis saxatilis). However, many species have
restricted ranges within the core area of the group, and
some are endemic to single massifs such as the Pyrenees
(e.g. Saxifraga aquatica), the Massif Central and Cevennes
Figure 6 The distribution of species in theOxyria digyna element.
Figure 7 The distribution of species in theErysimum duriaei element.
Floristic elements in European plants
Journal of Biogeography 34, 1848–1872 1857ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 31 / 76
Neutraali malli Suomen kasvisto
Euroopan kasvisto: puut
Betula pubescensLarix sibiricaPicea abies
Pinus sylvestrisP. sibirica
Carpinus betulaFagus sylvatica
Quercus petraea
Betula pendulaQuercus roburUlmus glabra
Quercus mas
Larix deciduaPinus cembraP. uncinata
Abies albaAlnus viridisPinus mugo
Taxus baccataPopulus albaP. canescens
P. nigraQuercus pubescensCarpinus orientalis
Celtis caucasicaFagus orientalisQuercus spp x 4
Laurus nobilisPinus halepensis
P. pineaQuercus coccifera
Q. ilexPinus pinaster
Quercus suber + 5Abies nebrodensis
Alnus cordataQuercus congesta
Q. sicula
Castanea sativaCeltis australis
Ostrya carpinifoliaQuercus cerris
Abies cephalonicaPinus heldreichii
Platanus orientalis
Pinus brutiaQuercus macrolepis
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 32 / 76
Neutraali malli Suomen kasvisto
Eurooppalaiset elementit Suomessa
Equisetum sylvaticum
Ranunculus bulbosus
Lychnis flos−cuculi
Oxyria digyna
Erysimum duriaei
Salix serpillifolia Rumex alpinus
Cochlearia danica
Alyssum alyssoides
Dianthus moesiacus
Dianthus capitatus
Osyris alba
Silene scabrifolia
Sarcocapnos enneaphyllaLobularia maritima
Ostrya carpinifolia
Ranunculus psilostachys
Brassica cretica
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 33 / 76
Neutraali malli Suomen kasvisto
Eurooppalaiset elementit Suomessa I
Valtaosa Suomen lajeista boreaalisia taigalajeja (Equisetumsylvaticum), keskieurooppalaisia (Silene flos-cuculi) ja Skandinaviantunturikasveja ja muita pohjoisia lajeja (Oxyria digyna)
Eurooppalaiset lajiryhmät vastaavat osittain perinteisiä suomalaisiaelementtejä, mutta vain likipitäenEquisetum sylvaticum: Suuri osa yleisistä, koko maassa tavattavistalajeista
Actaea rubra (erythrocarpa), Alnus incana, Trollius europaeusSilene (Lychnis) flos-cuculi: Suurin elementti, lajit eteläisiä
Actaea spicata, Alnus glutinosa, Anemone nemorosa, Caltha palustrisRanunculus bulbosus: Lounais-Suomen suboseaaninen elementti
Dryopteris dilatata, Saxifraga granulataOxyria digyna: Tunturikasvit ja muut pohjoiset lajit
Salix lanata, Saxifraga oppositifolia
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 34 / 76
Neutraali malli Suomen kasvisto
Eurooppalaiset elementit Suomessa II
Cochlearia danica: merenrantakasvit ja oseaaniset lajitCakile maritima, Honckenya peploides, Myrica gale
Salix serpillifolia: muutama Alppien laji ulottuu Suomeen (Pulsatillavernalis), mutta myös monet Oxyria digyna -lajit AlpeillaRumex alpinus: Joitain Karpaattien lajeja ulottuu Suomeen
Aconitum napellus, Asplenium viride, Clematis alpina
Alyssum alyssoides: muutamia kulttuurin seuralaisia, enimmäkseenharvinaisia osin vakiintuneita tulokkaita
Chenopodium, Isatis tinctoria
Dianthus capitatus: kulttuurin seuralaisia, usein kuivilla paikoillaBunias orientalis, Silene viscosa
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 35 / 76
Neutraali malli Suomen kasvisto
Jääkausi ja levikkiryhmät
Spekulatiivinen ryhmittely nykylevikin mukaanEtenkin tunturikasvien disjunktioiden selittäminen KölivuorillaSuomessa reliktilevikkejä jääkauden lämpö- ja kylmäjaksoiltaKylmärelikteiksi väitettyjä tunturikasveja: Dryas octopetala, Arnicastricta (alpina)Ruijanesikkoryhmä: Oulun seudun ’arktisia lajeja’’Kylmänkestävät pioneerit’ itse asiassa paahderinteiden harjukasveja:Anthyllis vulneraria ssp. fennica, Oxytropis campestris ssp. sordida’Lämpökauden reliktejä’ kuten Cladium mariscus: levikkien reunatyleensä repaleisia
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 36 / 76
Neutraali malli Suomen kasvisto
Lapinvuokko — reliktilevikki?
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 39 / 76
Neutraali malli Suomen kasvisto
Refugiot
Alpiinisten kasviendisjunktio:glasiaalirefugiotToisaalta evidenssiolemassa olostaToisaalta eivät tarpeennykylevikin selittämiseksi
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 40 / 76
Neutraali malli Suomen kasvisto
Kuusi idästä, mutta Norjaan myös Lofoottien refugiosta
but all populations from the northern Europeanrange share a similar old and ancestral nad1mtDNA variant (20).
A likely explanation for the distribution pat-terns of haplotypes A and B is that scatteredpopulations carrying A survived the LGM in mi-croenvironmentally favorable pockets in westernNorway, where proximity to the relatively tem-perate conditions of the Atlantic Ocean may havefavored survival during glacial times. After cli-mate warming, colonization of Scandinaviastarted first from local western stands that initiallyexpanded slowly and eventually mixed with pop-ulations arriving from the east. Rapidly risingsea levels and complex glaciated terrain probablycontributed to the scattered populations on theAtlantic coast remaining isolated and becomingfixed for haplotype A. As spruce can persist forhundreds of years by vegetative propagationunder unfavorable conditions (22), it is possiblethat these trees produced no pollen, leaving notrace in the palynological record.
To test whether spruce trees carrying hap-lotype A are indeed early Scandinavian survi-vors, we used aDNAextracted from lake sediments(23, 24) from the Trøndelag region (63°N; centralNorway) and on the island of Andøya (69°N;northernNorway). Such environmental aDNA (25)is shown to be local in origin and provides a proxyfor plant paleo-community reconstruction thatexhibits more similarity to macrofossils than topollen records (26, 27). We also extracted and
analyzed aDNA from the oldest spruce pollenfound in the Trøndelag core [6300 cal. yr B.P.;(16)]. We could not recover mtDNA from thedeepest Andøya samples, likely due to the rel-atively low–copy number of mtDNA comparedto chloroplast DNA (cpDNA) in plant tissues(see below). In contrast, we detected the sprucemtDNA haplotype A from the deepest samples atTrøndelag dating to 10,300 and 6500 cal. yr B.P.and from spruce pollen 6300 years old (Fig. 2and table S5A).
These early occurrences of haplotype A inTrøndelag indicate that spruce was alreadypresent in the region during the early Holocene,much earlier than the first wave of colonizationinferred from pollen analyses (~3000 cal. yr B.P.)(19). Our data are reinforced by a recent report oflate-glacial and early-Holocene pollen andstomata of Pinus and Picea in the Dovre moun-tains in central Norway, which also indicate localpresence of the two taxa (28).
To further test where, and for how long,spruce trees might have survived in Scandinavia,we focused our studies at Andøya (Fig. 1A). This is-land is particular with regards to LateWeichselianand Holocene paleoenvironments, due to its long,continuous lacustrine sedimentary records, itsearly deglaciation at ~26,000 cal. yr B.P. (29),and its location only a few hundred kilometersnorth of the modern northern limit of spruce hap-lotype A. Thus, at a time when almost the wholeof Scandinavia was covered by ice, Andøya’s
northern tip harbored a nunatak ecosystem [see(29–31)]. Previous work on the early vegetationhistory of Andøya is based on pollen records, andto a lesser extent on macrofossils (32). Here, wecombinedmacrofossil and aDNAanalyses of lakesediments to assess past local vegetation (16, 33)to amplify short fragments of cpDNA [generally<100 bp; thus, shorter than the mtDNA mh05fragment (16)]. Chloroplast DNA is alsomore com-mon than mtDNA in many plant tissues (34, 35),increasing the chances of DNA survival and de-tection in older sediments.
Results from both macrofossil and cpDNAanalyses indicate the presence of a polar desertor open pioneer vegetation community from~ 22,000 cal. yr B.P. (fig. S3). Tundra herb di-versity increased with a climatic warming around15,000 cal. yr B.P., and there were further in-creases and changes in diversity in the Holocene,including the establishment of boreal speciessuch as rowan (Sorbus aucuparia), generally re-flecting previous vegetation reconstructions [see(29–31)]. The most notable result was the findingof DNA of pine (Pinus sp.) in sediments dating to~22,000 and ~19,200 cal. yr B.P., and spruce(Picea sp.) in sediments dating to ~17,700 cal. yrB.P. (Fig. 2 and fig. S3).
There is little knowledge about aDNA ta-phonomy and the processes that may deliver plantDNA to lake sediments. However, it is likelythat tree aDNA at Andøya and Trøndelag re-flects contemporaneous plant biomass derived
Fig. 1. (A) Reconstruc-tions of stages in thedevelopment of theScandinavian Ice Sheet45,000 to 9500 cal. yrB.P. compiled from arange of sources (16)A, Andøya; T, Trøndelag.(B) Geographical dis-tribution of mitochondrialmh05 haplotypes A (darkblue circles) and B (redcircles) in Norway sprucepopulations. Size of thecircles is proportional topopulation size (centeredwhite dots indicate pop-ulations with N < 10; ta-ble S1). Arrows suggestpostglacial movementsof the two haplotypesafter the LGM. The oliveshading shows the naturalrange of Norway spruce.
2 MARCH 2012 VOL 335 SCIENCE www.sciencemag.org1084
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Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 41 / 76
Neutraali malli Suomen kasvisto
Primula sibirica -ryhmä
Primula nutans var. jokelae -ryhmä pedanttisille
Perämeri ja VienanmeriMerenrantalajeja: Carexhalophila, C. paleacea, Hippuristetraphylla, Potentilla anserinassp. egedii, PuccinelliaphryganodesLöyhemmin Carex glareosa, C.mackenzieiTulvamailla Alopecurusarundinaceus, Arctophila fulva,Catabrosa aquatica, Salixtriandra
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 42 / 76
Neutraali malli Suomen kasvisto
Primula nutans: kliini Siperiasta?
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 44 / 76
Neutraali malli Tulokaslajit
Suomen tulokaslajit
Kaikki kasvit ovat saapuneet maahan vasta jääkauden jälkeen, muttaihminen on kiihdyttänyt kasvilajien invaasiota. Osa alkuperäisen näköisistäluonnonkasveista saattaa olla vanhoja kulttuuritulokkaita, ja joidenkin lajientiedetään varmasti saapuneen vasta hiljan. Joitain kasvilajeja pidetään jopahaitallisina invaasiolajeina. Kasvilajit saavunta heijastaa kulttuurin jakaupan muutoksia ja kasveissakin voidaan erottaa kulttuurikerrostumia.
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 45 / 76
Neutraali malli Tulokaslajit
Alkuperäiset lajit ja tulokkaat
Kaikki putkilokasvit Ilman uustulokkaita Ilman muinaistulokkaita
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 46 / 76
Neutraali malli Tulokaslajit
Eri-ikäisiä tulokkaita
Muinaistulokkaat eli arkeofyytit saapuneet ennen 1600-lukuaVanhan asutuksen seuralaisia: Aethusa cynapium, Sisymbriumofficinale, Chelidonium majusLuonnonvaraiselta vaikuttavia niitty- ja lehtokasveja: Achilleamillefolium, Carum carvi, Anthriscus sylvestris (mutta voivat ollaalkuperäisiäkin. . . )
Uustulokkaat eli neofyytitJoukossa hyvinkin vakiintuneen oloisia kasveja: Galium album,Matricaria matricarioides
Satunnaiset kasvit: eivät pysty muodostamaan pysyvää, lisääntyvääkantaa
Voivat olla pysyviä, jos immigraatio vakaa: esim. pellavan rikkaruohotitse asiassa satunnaisia
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 47 / 76
Neutraali malli Tulokaslajit
Vanha kulttuuri
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 49 / 76
Neutraali malli Tulokaslajit
Nuori tulokas: Kaisaniemi 1849
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 50 / 76
Neutraali malli Tulokaslajit
Kanadanvesirutto (Elodea canadensis)
R. K. Heikkinen et al. (2009) BioRisk2, 1–32
vaalea: havaittu ennen v 1985
tumma: havaittu v 1985
jälkeen
Levisi Helsinginkasvitieteellisestäpuutarhasta 1800-luvunlopussa1900-luvulla reheviinEtelä-Suomen järviin, nytjo KuusamossaLeviämisen mekanismitja järvien valintatuntematon
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 51 / 76
Neutraali malli Tulokaslajit
Vieraslajit
Lähde: Sami Aikio /Kastikka-tietokanta
Vieraslajit (invaasiolajit) vakava uhkamonella alueella: Esim. Uuden-Seelanninputkilokasveista puolet tulokkaita ja 400ongelmallisiaVieraslajit haittana eurooppalaistentuomalla talousmuodolle (laidunnus, viljely),mutta leviävät myös luontaisiinhabitaatteihinVakavin uhka temperaattisessa jamediterraanisessa ilmastossaSuomessa kiinnitetty huomiota viimevuosina: Vieraslajistrategia 30.3.2011Havainnot keskittyvät suurtenasutuskeskusten ja teiden lähelle – muttaniin havaitsijatkin
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 52 / 76
Neutraali malli Tulokaslajit
Suomen vieraslajeja
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 53 / 76
Neutraali malli Tulokaslajit
Kansallinen vieraslajistrategia (2011)http://www.mmm.fi/fi/index/etusivu/ymparisto/luonnonmonimuotoisuus/vieraslajit.html
Kansallinen vieraslajistrategia pyrkii estämään vieraslajien leviämisenVieraita lajeja ei saa siirtää tai istuttaa luontoon ja esiintymät ontuhottavaMonen vieraslajin lähteenä kasviharrastajat (puutarhat, terraariot,akvaariot), mutta myös kauppa ja liikenne, riistaeläimiä ja vesieläimiäjopa istutettuTodelliset ongelmat eläimissä, joissa tahallisia siirtoistutuksia(aasialainen supikoira), tarhakarkulaisia (amerikkalaiset minkki,piisami, majava) että liikenteen mukana tulleita (Kaspianmerenpetovesikirppu, amerikkalainen kampamaneetti), muttakasvipuolellakin joitain mahdollisia pahiksiaVieraslajistrategiassa etenkin ihmiselle ja luonnolle selvästi haitallisialajeja, ml. tautejaVakavimmille ehdotetaan lakiin perustuvaa hävittämistäPutkilokasveja 24, joista erittäin haitallisia kurtturuusu ja jättiputket
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 54 / 76
Neutraali malli Tulokaslajit
Kasvien pahiksia: Vesirutto Elodea canadensis
E. canadensis laajalle levinnyt, myös E. nuttallii tuloillaan PerämerenympäriAmerikkalainen laji, joka karkasi Helsingin kasvitieteellisestäpuutarhasta, mutta nykyesiintymissä varmaan myös akvaarioistaluontoon laskettujaVoi muodostaa todellisia massaesiintymiä, jolloin varjostaa kesälläpohjan ja kuolleet kasvit kuluttuvat lahotessaan hapenLeviää varren kappaleista, joita siirtänevät vesilinnut, veneilijät jakalastajatEi suvullista lisääntymistä SuomessaYmpäristövaatimukset huonosti tunnettuja, mutta usein korkeahko pHja alkaliniteettiVoi esiintyä Lapin läänin eteläosiin astiKansallinen vieraslajistrategia: paikallisesti tarkkailtava (Itämeressä)
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 55 / 76
Neutraali malli Tulokaslajit
Kasvien pahiksia: kurtturuusu, Rosa rugosa
Itäaasialainen pohjoisen Tyynenmeren hiekkarantojen lajiMeillä koristekasvikarkulainen, jonka perusmuoto levinnyt Itämerenhiekkarannoille, missä voi kasvaa osin hiekan sisään hautautuneenalaajoina kasvustoinaSaattaa vaikuttaa muihin hiekkarantakasveihin ja hiekkarantojeneläimiin ja ihmisen mielestä kiusallinen: piikikäs pensas uimarannallaPidetään uhkana hiekkarantojen biodiversiteetille: vaikuttaa muihinlajeihinKauniit kukat, joten ei pidetä Suomen luontoon sopivanaLeviää siemenistä (kiulukat)Esiintyy karkulaisena myös ihmisen muuttamassa ympäristössä(kaupungit, tienvarret)Kansallinen vieraslajistrategia: erittäin haitallinenRunsaasti viljelylajikkeita, joista kaikki eivät ole leviäviä
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 57 / 76
Neutraali malli Tulokaslajit
Lupiini, Lupinus polyphyllus
Pohjoisamerikkalainen koristekasvi, levinnytSuomessa erittäin nopeasti etenkin teiden varsille,mutta myös niityilleEsiintyy vain ihmisen muokkaamissa ympäristöissä,joissa kuitenkin muita arvokkaina pidettyjä ihmisenseuralaislajeja ja joita halutaan suojellaperinnebiotooppeinaLeviää siemenistä, muodostaa laajoja kasvustoja,joissa muiden lajien lajimäärä alentuuHernekasvi: typen yhteyttäjäTielaitos suosi pitkään: tienvarret niitettiin vastalupiinin kukinnan jälkeen (nyt toisin)Komeakukkainen, joten pidetään Suomen luontoonsopimattomana (VLS: "maisemallisesti haitallinen")
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 59 / 76
Neutraali malli Tulokaslajit
Kasvien pahiksia: jättiputket, Heracleum mantegazzianumH. persicum, H. sosnowskyi
Kaukasialaisia koristekasvikarkulaisia, käytetty(kuulema) myös rehukasveinaMeillä tavallisin H. mantegazzianum, mutta myös”Stalinin kosto” (Месть Сталина, H. sosnowskyi)tavattu KarjalassaMeillä yleensä paikallisia eivätkä ole useinkaanlevinneet laajalti alkuperänsä ulkopuolellaKarjalan kannaksella niityt voivat olla näidenvallassa ja Pohjois-Norjassa runsas kulttuurinseuralainen ("Tromsø palm")Ihmiselle ja eläimille haitallinen: aiheuttaa UV-valonkanssa kivuliaita ihottumia ja pysyviä ihomuutoksiasekä palovamman kaltaisia oireitaKansalllinen vieraslajistrategia: erittäin haitallinen
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 61 / 76
Neutraali malli Tulokaslajit
Paimenmatara, Galium album
VLS pitää tulokkaana: ollut kuitenkin Suomessa ainakin 1800-luvultaAlkujaan itäinen (Kannas), mutta levinnyt nopeasti heinänviljelynmyötä 1900-luvulla (kuten ojakärsämö, Achillea ptarmica)Pääasiallinen haitta: risteytyy alkuperäisen keltamataran (Galiumverum) kanssa ja muodostaa hybridinotomorfin G.× pomeranicum,joka risteytyy paimenmataran kanssaAlkuperäinen G. verum hävinnyt laajoilta alueilta, jäljellä vainvaaleankeltaisia, särmävartisia risteymiä
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 62 / 76
Neutraali malli Tulokaslajit
Vieraslajeille alttiit habitaatit
Ihmisen muuttamat habitaatit ("kulttuurimaisemat")Monet lajit alkuperäisellä paikallaanVoivat syrjäyttää muita ihmisen tuomia lajeja, joista joitain pidetäänarvokkaina ja suojeltavinaJopa tienvarsia pidetään nykyään arvokkaina niitty- ja ketolajienhabitaattinaPaikallisesti esim. taimistojen rikkaruohoja: rikkanenätti (Rorippasylvestris), amerikanhorsmat (Epilobium adenocaulon, E. ciliatum)
Vesistöt: esim Elodea, Ceratophyllum demersumPuronvarret, myös lehdoissa
Jättipalsami (Impatiens glandulifera), ruttojuuri (Petasites hybridus),isosorsimo (Glyceria maxima)
Merenrannat, etenkin hietikot
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 64 / 76
Neutraali malli Tulokaslajit
Invasiivisuuden arviointiT Tyler et al, Nordic J Bot 33, 300–317 (2015)
Kilpailukyky luonnossa: heikoimmat häiriöalueilla ja ihmisenmuuttamassa kasvillisuudessa, vahvemmat metsissä ja soillaPopulaatiotiheys: vahvimmat muodostavat laajoja yhtenäisiäkasvustojaLeviämiskyky: monet tulokkaat tuskin tuottavat leviäimiä ja jäävätpaikallisiksiRisteytyminen alkuperäisten lajien kanssaMaahanmuuton ikä: tuoreimmat tulokkaat pahimpia – vanhat josaavuttaneet tasapainonMaantieteellinen etäisyys: mitä kauempaa, sitä epäilyttävämpääYleisyys nyt
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 68 / 76
Neutraali malli Tulokaslajit
Liikenne ja kasvien leviäminen
Vanhoja kauppateitä pitkin: VienastaChaerophyllum prescottii, viikinkien mukanaCardamine parviflora, Allium ursinumVenäläiset varuskunnat: Bunias orientalis,Berteroa incanaSaksalaiset sotajoukot: Arabidopsis arenosaTielaitos: Festuca duriuscula, Lupinuspolyphyllus, Silene tatarica, LotuscorniculatusVR: Senecio viscosus, Lepidium densiflorum,Erigeron canadensis
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 69 / 76
Neutraali malli Tulokaslajit
Kaupunkien kasvit
Kaupungit habitaattien mosaiikkiMyös metsät kulttuurivaikutteisia: heiniä, koiran- ja vuohenputkeaVanhoissa kaupungeissa vanhan kulttuurin seuralaisiaMerkillisiä maannoksia: rakennusjätteitä, saasteita, pohjamaataErittäin suuri putkilokasvidiversiteetti — kaikki eli γ = β + α korkeat
Helsingin Suurkirkon portailla 74 putkilokasvilajia (1996)Oulussa 1017 lajia (2001), joista 402 satunnaisia ja 106 karkulaisia
Useita historiallisia kerrostumia, joista kiintoisin lähes sukupuuttoonajautunut Toppilan painolastikasvistoOulun kaupungin nimikkokasvi hietikkopitkäpalko (Arabidopsis arenosa)on radanvarsia pitkin leviävä saksalaistulokas
Jari Oksanen (Oulun yliopisto) Biogeo: KAMA 2015 75 / 76
