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Ancient subalpine clonal spruces (Picea abies) – sources of postglacial vegetation history in the Swedish Scandes
Mid Sweden University, Faculty of Science, Technology and Media, Department of Natural Sciences, Engineering and Mathematics. (Växtekologi)
Institutionen för ekologi, miljö och geovetenskap, Umeå universitet.
2011 (English)In: Arctic, ISSN 0004-0843, E-ISSN 1923-1245, Vol. 64, no 2, p. 183-196Article in journal (Refereed) Published
Abstract [en]

This study addresses the long-standing issue of postglacial immigration of Picea abies (Norway spruce) into Scandinavia. Methodologically, the main focus is on megafossil tree remains (wood and cones) of spruce and other species, retrieved from the treeline ecotone (Swedish Scandes), as a tool for vegetation reconstruction. Radiocarbon dating of megafossils, preserved in the soil underneath layering clonal groups of Picea abies, provide the core data. Living spruce clones, with in some cases likely continuity back to the early Holocene (9500 cal. yr BP onwards), were found at high-elevations. First postglacial arrival to the Swedish Scandes at this stage concurs with previous megafossil inferences. This is several millennia earlier than inferred from pollen data and very soon after regional deglaciation. Persistence of some individual Picea clones since the early Holocene thermal optimum and up to the present is indicative of permanently open or semi-open spots in the high-mountain landscape, also at times when treelines in general were much higher than present. Initially, Picea clones appear to have existed in a regional no-analogue vegetation matrix of widely scattered pine (Pinus sylvestris), mountain birch (Betula pubescens ssp. czerepanovii), Siberian larch (Larix sibirica) and thermophilic broadleaved deciduous species. In response to subsequent neoglacial cooling, the alpine character of the landscape has been enhanced through a lowered pine treeline and disappearance of larch and thermophiles. Spruces, which escaped fire and other calamities, endured due to their inherent phenotypic plasticity. Increasing climatic harshness throughout the Holocene conserved them as crippled krummholz, protected from winter stress by almost complete snow coverage. Appearance of Picea abies exclusively in the west, shortly after the deglaciation, could suggest that it has immigrated from “cryptic” ice age refugia much closer to Scandinavia than conventionally thought.

Place, publisher, year, edition, pages
2011. Vol. 64, no 2, p. 183-196
Keywords [en]
Picea abies, clones, megafossils, immigration, Holocene, cryptic refugia, Swedish Scandes.
National Category
Ecology
Identifiers
URN: urn:nbn:se:miun:diva-12666ISI: 000292226100004Scopus ID: 2-s2.0-79958114279OAI: oai:DiVA.org:miun-12666DiVA, id: diva2:377346
Available from: 2010-12-14 Created: 2010-12-14 Last updated: 2017-12-11Bibliographically approved
In thesis
1. Treeline dynamics in short and long term perspectives: observational and historical evidence from the southern Swedish Scandes
Open this publication in new window or tab >>Treeline dynamics in short and long term perspectives: observational and historical evidence from the southern Swedish Scandes
2010 (English)Licentiate thesis, comprehensive summary (Other academic)
Abstract [en]

Against the background of past, recent and future climate change, the present thesis addresses elevational shifts of alpine treelines in the Swedish Scandes. By definition, treeline refers to the elevation (m a.s.l.) at a specific site of the upper trees of a specific tree species, at least 2 m tall.

Based on historical records, the first part of the thesis reports and analyzes the magnitude of treeline displacements for the main trees species (Betula pubescens ssp. czerepanovii, Picea abies and Pinus sylvestris) since the early 20th century. The study covered a large and heterogeneous region and more than 100 sites. Concurrent with temperature rise by c. 1.4 °C over the past century, maximum treeline advances of all species amount to about 200 m. That is virtually what should be predicted from the recorded temperature change over the same period of time. Thus, it appears that under ideal conditions, treelines respond in close equilibrium with air temperature evolution. However, over most parts of the landscape, conditions are not that ideal and treeline upshifts have therefore been much smaller. The main reason for that discrepancy was found to be topoclimatic constraints, i.e. the combined action of geomorphology, wind, snow distribution, soil depth, etc., which over large parts of the alpine landscape preclude treelines to reach their potential thermal limit.

Recorded treeline advance by maximum 200 m or so over the past century emerges as a truly anomalous event in late Holocene vegetation history.

The second part of the thesis is focused more on long-term changes of treelines and one specific and prevalent mechanism of treeline change. The first part of the thesis revealed that for Picea and Betula, treeline shift was accomplished largely by phenotypic transformation of old-established stunted and prostrate individuals (krummholz) growing high above the treeline. In obvious response to climate warming over the past century, such individuals have transformed into erect tree form, whereby the treeline (as defined here) has risen. As a means for deeper understanding of this mode of positional treeline change, extant clonal spruces, growing around the treeline, were radiocarbon dated from megafossil remains preserved in the soil underneath their canopies. It turned out that Picea abies in particular may attain almost eternal life due to its capability for vegetative reproduction and phenotypic plasticity. Some living clones were in fact inferred to have existed already 9500 years ago, and have thus persisted at the same spot throughout almost the entire Holocene. This contrasts with other tree species, which have left no living relicts from the early Holocene, when they actually grew equally high as the spruce. Thereafter they retracted by more than 300 m in elevation supporting that also on that temporal scale, treelines are highly responsive to climate change.

The early appearance of Picea in the Scandes, suggests that Picea “hibernated” the last glacial phase much closer to Scandinavia than earlier thought. It has also immigrated to northern Sweden much earlier than the old-established wisdom.

The experiences gained in this thesis should constitute essential components of any model striving to the project landscape ecological consequences of possible future climate shifts.

Place, publisher, year, edition, pages
Sundsvall: Department of Natural Sciences, 2010
Series
Mid Sweden University licentiate thesis, ISSN 1652-8948 ; 51
Keywords
Betula pubescens ssp. czerepanovii, Picea abies, Pinus sylvestris, climate change, monitoring, treeline advance, clones, megafossils, immigration, Holocene, cryptic refugia, Swedish Scandes
National Category
Ecology
Identifiers
urn:nbn:se:miun:diva-12670 (URN)978-91-86694-09-8 (ISBN)
Presentation
2010-12-10, Q221, Campus, Östersund, 13:00 (Swedish)
Opponent
Supervisors
Available from: 2010-12-15 Created: 2010-12-14 Last updated: 2013-02-18Bibliographically approved
2. Treeline dynamics in short and long term perspectives: observational and historical evidence from the southern Swedish Scandes
Open this publication in new window or tab >>Treeline dynamics in short and long term perspectives: observational and historical evidence from the southern Swedish Scandes
2013 (English)Doctoral thesis, comprehensive summary (Other academic)
Abstract [en]

Treelines in high-mountain regions are constrained by heat deficiency, although the working mechanisms are still not entirely understood. Observational and paleoecological studies on treeline performance may contribute to increased understanding of the treeline phenomenon in general. The present thesis addresses elevational shifts of alpine treelines in the Swedish Scandes. By various analytical tools, the studies embrace widely different temporal scales.

The concept treeline refers to the elevation (m a.s.l.) at a specific site of the upper individual tree of a certain tree species, at least 2 m tall. All the principal tree species in the Scandes are concerned, i.e. mountain birch (Betula pubescens ssp. czerepanovii), Norway spruce (Picea abies) and Scots pine (Pinus sylvestris).

Paper I deals with regional treeline dynamics at more than 100 sites over the past 100 years. Concurrent with temperature rise by c. 1.4 °C over the same period, maximum treeline advances of all species amount to about 200 m. Thus, under ideal conditions, treelines respond in close equilibrium with air temperature evolution. However, over most parts of the landscape, treeline upshifts have been much smaller than 200 m, which relates to the combined action of geomorphology, wind, snow distribution and soil depth. After 1975, the birch has lost its role as the most rapidly advancing tree species, being superseded by pine and spruce.

Paper II is a short-term (2005/2007-2010/2011) study of mountain birch treeline performance along a regional maritimity-continentality gradient. Upshift by 3.0 yr-1 in the maritime part of the gradient contrasts to retreat by 0.4 m yr-1 in the continental part. In the latter area, earlier and more complete melting of late-lying snow patches has seemingly progressed to a state when soil drought sets back the vigour of existing birches and precludes sexual regeneration and upslope advance of the treeline. In the maritime area, extensive and deep snow packs still exist above the treeline and constrain its position, although some release is taking place in the current warm climate.

Paper III explores treeline change by phenotypic transformation of old-established stunted and prostrate spruce individuals (krummholz) growing high above the treeline and is based on analyses of radiocarbon-dated megafossils, preserved in the soil underneath clonal groups of spruce. Living spruce clones, which in some cases may date back to the early Holocene (9500 cal. yr BP), suggests that spruce immigrated from “cryptic” ice age refugia much closer to Scandinavia than conventionally thought. As the krummholz form presupposes open and windy habitats, it is inferred that permanently open spots prevailed in the high-mountain landscape even during periods when treelines in general were much higher than today.

Paper IV reports radiocarbon dates of wood samples, retrieved from newly exposed glacier forefields at three main sites, located high above the modern treelines and embracing the entire Swedish Scandes. It appears that pine colonized early emerging nunataks already during the Late Glacial. Around 9600-9500 cal. yr BP a first massive wave of tree establishment, birch and pine, took place in “empty” glacier cirques. Both species grew 400-600 m above their present day treeline position and accordingly, the summer temperatures may have been 3.5 °C warmer than present (uncorrected for land uplift). During the entire interval 9600 to 4400 cal. yr BP, birch prospered 100-150 m above the uppermost pines. In response to Neoglacial cooling, treelines of both birch and pine descended until their final disappearance from the record 4400 and 5900 cal. yr BP, respectively. Thereafter, these habitats experienced increased snow accumulation and glacier inception.

Abstract [sv]

Avhandlingen belyser förskjutningar i olika tidsskalor av den alpina trädgränsens läge i de svenska Skanderna. Trädgränsen definieras som den högsta nivån (m ö.h.) för minst 2 m höga individer av en viss art i en definierad del av en fjällsluttning. Avhandlingen består av fyra separata uppsatser, publicerade i olika välrenommerade vetenskapliga tidskrifter. 

I Paper I analyseras förändringar av trädgränserna för fjällens vanligaste trädarter, fjällbjörk, gran och tall (Betula pubescens ssp. czerepanovii, Picea abies och Pinus sylvestris) mellan 1915 och 2007. Undersökningen omfattar ett 8000 km2 stort, naturgeografiskt heterogent område, med mer än 100 lokaler. De maximala trädgränsförskjutningarna för samtliga trädarter uppgår till omkring 200 m. Resultaten motsvarar de trädgränsförskjutningar som teoretiskt kan förutsägas utifrån den temperaturhöjning med cirka 1,4 °C som skett under samma tidsperiod, förutsatt ideala förhållanden. För större delen av området råder emellertid andra, icke-ideala förhållanden, varför trädgränsernas uppflyttning i allmänhet blivit avsevärt mindre än 200 m. Den främsta anledningen till detta är lokala topoklimatiska begränsningar, d.v.s. kombinerade effekter av geomorfologi, vind, snöfördelning, jorddjup, etc., som i stora delar av det starkt brutna fjällandskapet mer eller mindre effektivt hindrar träden från att nå sina potentiellt högst belägna växtplatser betingade av temperaturen. Efter 1975 har björken förlorat sin roll som arten med den snabbast expanderande trädgränsen. I stället har tall och gran avancerat med större hastighet. Det innebär att även om klimatets uppvärmning fortsätter, så kommer det subalpina björkbältet att expandera i avsevärt mindre omfattning än vad som ofta förebådats. Möjligtvis kommer det att ersättas av tall.

Paper II behandlar björkens trädgränsdynamik under perioden 2005/2007-2010/2011 längs en regional klimatgradient med avseende på maritimitet/kontientalitet. Trädgränsen har under den aktuella perioden avancerat 3,0 m/år i den maritimt präglade delen av gradienten, vilket kontrasterar signifikant mot en sänkning med 0,4 m/år i området med mer kontinentalt klimat. Skillnaderna diskuteras i termer av klimatförändringens varierande effekter på snötäckets utbredning och varaktighet och dess inverkan på markfuktigheten. En allt tidigare total utsmältning av snölegorna i de kontinentala områdena har av allt att döma resulterat i vattenbrist under sommaren. Torka medför reducerad vitalitet för existerande björkar och förhindrar både sexuell förökning och uppflyttning av trädgränsen. I de maritima delarna kvarligger alltjämt mycket snö under en stor del av sommaren. Trädgränsens position har därför kunnat bibehållas eller flyttas upp. Vissa omständigheter tyder på att trädgränsens stigning i högre grad har varit baserad på fröföryngring efter 1975, jämfört med perioden 1915-1975.

Utgångspunkten för Paper III är erfarenheter från Paper I, som visar att trädgränsens uppflyttning för gran och björk huvudsakligen är resultatet av ökad höjdtillväxt av äldre, i vissa fall flertusenåriga, mer eller mindre buskformiga individer (krummholz), som vuxit på nivåer långt ovanför trädgränsen. Som ett svar på de senaste hundra årens varmare klimat har dessa antagit trädform, varigenom trädgränsen höjts. För en fördjupad förståelse av den här mekanismen har megafossil, d.v.s. grova vedrester bevarade i marken under gamla grankloner i trädgränsekotonen, 14C-daterats. Resultaten tyder på att granar i exponerad fjällmiljö kan uppnå i det närmaste ”evigt” liv genom sin förmåga till vegetativ förökning och möjligheten att växla mellan busk- och trädform i takt med klimatets växlingar. Vissa nu levande kloner existerade av allt att döma redan för 9500 år sedan. Den nu dokumenterat tidiga förekomsten av gran, bekräftar den på senare tid allt tydligare bilden av granen som en tidig invandrare till fjällkedjan. Möjligtvis har granen ”övervintrat” den senaste istiden närmare Skandinavien än vad som till helt nyligen varit den gängse uppfattningen.

Paper IV behandlar en för Skandinavien ny metod för historisk trädgränsrekonstruktion. I uppsatsen analyseras 14C-dateringar av totalt 78 större veddelar (megafossil) som nyligen exponerats i anslutning till smältande glaciäris och ”perenna” snölegor i tre huvudområden, Helags-Sylarna, Tärna och Abisko, högt ovanför dagens trädgräns. Det framkommer att tall (Pinus sylvestris) koloniserade tidigt framsmälta nunatakker redan under senglacial tid. För omkring 9600-9500 år sedan inträffade en första massiv våg av björk- och talletablering i isfria glaciärnischer. Båda arterna växte 400-600 m ovanför sina nuvarande trädgränspositioner, i ett klimat som kan ha varit 3,5 °C varmare än idag.

Under intervallet 9600 till 4400 BP uppträdde björken i ett 100-150 m brett bälte ovanför de översta tallarna. Som ett svar på klimatets successiva avkylning under senare delen av Holocen sänktes både björkens och tallens trädgränser i de aktuella miljöerna, till dess de för 4400 respektive 5900 år sedan helt försvann från lokaler där glaciärer och perenna snöfält började bildas. De analyserade trädresterna, som länge bevarats av glaciäris och perenn snö representerar en period med ett klimat långt varmare än under det senaste århundradet. Med denna analogi från det förflutna kan det därför antas att i en framtid där sommartemperaturerna rent hypotetiskt är 3,5 °C högre än i nutiden, skulle trädgränserna lokalt kunna flyttas upp med ungefär 600 m.

Place, publisher, year, edition, pages
Sundsvall: Kopieringen Mid Sweden University, 2013. p. 136
Series
Mid Sweden University doctoral thesis, ISSN 1652-893X ; 143
Keywords
Betula pubescens ssp. czerepanovii, Picea abies, Pinus sylvestris, Larix sibirica, climate, Betula pubescens ssp. czerepanovii, Picea abies, Pinus sylvestris, Larix sibirica
National Category
Natural Sciences
Identifiers
urn:nbn:se:miun:diva-18458 (URN)978-91-87103-63-6 (ISBN)
Public defence
2013-03-15, Sal G1352,, Mittuniversitetet, Östersund, 10:15 (Swedish)
Opponent
Supervisors
Available from: 2013-02-18 Created: 2013-02-12 Last updated: 2013-04-22Bibliographically approved

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