Between rear and leading edge [Elektronische Ressource] : juniper woodlands on the southern Tibetan Plateau ; a high mountain forest-line-ecosystem under environmental change / vorgelegt von Lars Opgenoorth

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Between Rear and Leading EdgeJuniper woodlands on the southern Tibetan PlateauA high mountain forest-line-ecosystem under environmental changeLars OpgenoorthBetween Rear and Leading EdgeJuniper woodlands on the southern Tibetan PlateauA high mountain forest-line-ecosystem under environmental changeDissertationzurErlangung des Doktorgradesder Naturwissenschaften(Dr. rer. nat.)demFachbereich Biologieder Philipps-Universität Marburgvorgelegt vonLars Opgenoorthaus DüsseldorfMarburg/Lahn 2009Vom Fachbereich Biologie der Philipps-Universität Marburgals Dissertation am 11.09.2009 angenommen.Erstgutachterin: Prof. Dr. Birgit ZiegenhagenZweitgutachter: PD Dr. Karsten WescheTag der mündlichen Prüfung am 11.09.2009List of PapersThis thesis is based on the following papers, which are referred to in the text by the Roman numerals I-IV. In addition a research proposal to the Volkswagen foundation is included as part of the outlook. It will be referred to with the roman numeral V.I Miehe, G., Miehe, S., Will, M., Opgenoorth, L., La Duo, Tsering Dorgeh and Liu, J.(2008): An inventory of forest relicts in the pastures of Southern Tibet (Xizang A.R.,China). Journal of Plant Ecology 194: 157–177.II Kaiser, K., Opgenoorth, L., Schoch, W. and Miehe, G. (2009): Charcoal and fossil woodfrompalaeosols,sedimentsandartificialstructuresindicatingLateHolocene woodland decline in southern Tibet (China).

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Between Rear and Leading Edge
Juniper woodlands on the
southern Tibetan Plateau
A high mountain forest-line-ecosystem
under environmental change
Lars OpgenoorthBetween Rear and Leading Edge
Juniper woodlands on the
southern Tibetan Plateau
A high mountain forest-line-ecosystem
under environmental change
Dissertation
zur
Erlangung des Doktorgrades
der Naturwissenschaften
(Dr. rer. nat.)
dem
Fachbereich Biologie
der Philipps-Universität Marburg
vorgelegt von
Lars Opgenoorth
aus Düsseldorf
Marburg/Lahn 2009Vom Fachbereich Biologie der Philipps-Universität Marburg
als Dissertation am 11.09.2009 angenommen.
Erstgutachterin: Prof. Dr. Birgit Ziegenhagen
Zweitgutachter: PD Dr. Karsten Wesche
Tag der mündlichen Prüfung am 11.09.2009List of Papers
This thesis is based on the following papers, which are referred to in the text by the Roman
numerals I-IV. In addition a research proposal to the Volkswagen foundation is included as
part of the outlook. It will be referred to with the roman numeral V.
I Miehe, G., Miehe, S., Will, M., Opgenoorth, L., La Duo, Tsering Dorgeh and Liu,
J.(2008): An inventory of forest relicts in the pastures of Southern Tibet (Xizang
A.R.,China). Journal of Plant Ecology 194: 157–177.
II Kaiser, K., Opgenoorth, L., Schoch, W. and Miehe, G. (2009): Charcoal and fossil
woodfrompalaeosols,sedimentsandartificialstructuresindicatingLateHolocene
woodland decline in southern Tibet (China). Quaternary Science Reviews 28: 1539–
1554.
III Opgenoorth, L., Vendramin, G., Mao, K., Miehe, G., Miehe, S., Liepelt., S. Liu, J.
and Ziegenhagen, B. (2009): Tree endurance on the Tibetan Plateau marks
the world’s highest known tree line of the Last Glacial Maximum. New Phytologist.
In press.
IV Opgenoorth,L.(2009):Identificationandcharacterizationofmicrosatellitemarkers
for Juniperus tibetica using next generation sequencing. Conservation Genetics
Resources. In press.
V Opgenoorth, L.: „Adaptation at the limits” or “the limits of Adaptation” - Tracing
abiotic stress-tolerance in Tibetan juniper trees along altitudinal and ecological
gradients. Research proposal to the Volkswagen Foundation.
Paper I through IV are reprints made available with permission from Springer (I, IV), Else-
vier (II), and Wiley-Blackwell (III).6Contents
Foreword ..............................................................................................................................9
1. Introduction .....................................................................................................................11
2. Results and Discussion ..................................................................................................14
2.1 The forest relics in the pastures of Southern Tibet (Xizang A.R.,China) – Paper I ...14
2.2 Late Holocene woodland decline in southern T ibet (China) – Paper II ........................15
2.3 Tree endurance on the Tibetan Plateau marks the world’s highest known tree
line of the Last Glacial Maximum – Paper III ............................................................16
2.4 Identificationandch aracterizationofmicrosa tellitemarkerusingnextgeneration
sequencing - Paper IV .............................................................................................17
3. Conclusions ....................................................................................................................19
4. Outlook ...........................................................................................................................20
4.1 „Adaptation at the limits” or “the limits of Adaptation” - Tracing abiotic stress
tolerance in Tibetan juniper trees along altitudinal and ecological gradients -
Research Proposal V ...............................................................................................20
5. Deutsche Zusammenfassung .........................................................................................22
6. Acknowledgements ........................................................................................................24
Bibliography .......................................................................................................................25
Paper I ................................................................................................................................31
Paper II ...............................................................................................................................55
Paper III ..............................................................................................................................73
Paper IV .............................................................................................................................87
Research Proposal V .........................................................................................................93
Erklärung ..........................................................................................................................107
Curriculum Vitae ...............................................................................................................109
78Foreword
The framing of this dissertation comes in a time of severe makeup and essentially amal-
gamation of the dif ferent fields of paleoecology , biogeography , conservation genetics and
eventually evolutionary biology mainly due to the recent revolutionary advances in sequenc-
ing technology but also due to important advances in our understanding of the natural pro-
cesses. It is almost symbolic that these seem to culminate in the year of Darwin
–withevolutionaryprocessesfinallyearningthefocustheyhavealwaysdeserved.
The makeup of the present thesis reflects this recent development only in parts: The char-
coal analytical study (Paper II) as well as the phylogeographic study (Paper III) of this work,
were designed in a “pre-next generation sequencing technology” context. Both studies
would today have been designed differently. The charcoal analytical approach would be
escorted or even replaced in large parts by shotgun sequencing approaches that utilize
ancient DNA. The advantages of this approach are the wider distribution of DNA in different
archives such as glaciers, permafrost and palaeosols, their superior power to determine
taxa on higher levels, and in case of good conservation, their potential use in evolutionary
contexts such as deep phylogeographies. Our first approaches using this technology are
currently underway: Pleistocene and Holocene soil samples from Paper II have been suc-
cessfully screened for aDNA recently in the lab of Eske Willerslev.
Phylogeographic studies on the other hand will change to the effect that the data sets will
increase drastically. While it seemed quite advanced in the beginning of my study to se-
quence the whole set of our samples instead of using RFLP’s (as was anticipated in the
study proposal) it already shows that such single locus data sets can hardly be published
anymore. One reason is that it is so much easier to produce larger data sets now. The es-
sential reason however is that multi-locus analysis from different genomes – or eventually
whole genomes - make the analysis much more reliable and enable us to implement more
complex analytical approaches such as coalescent simulation. These approaches bring us
as close as ever to being able to check a priori hypothesis in a historical context – much like
in the fields of experimental science. Neverthel ess, even (quasi-) experimental research
approaches need good background information for hypothesis building and testing.
The present study lies in the tradition of expedition science in the remoter places of this
planet and is intended to provide such background information on the forest history of the
Tibetan Plateau. It also intends to give an outlook how hypotheses based on this ground-
work can be tested in future experimental designs. In this respect, I take the freedom to
includearesearchproposaltotheV olkswagenFoundationasfinalpartofthisthesis.
On a more personal note, the makeup of this thesis also reflects my scientific “evolution”
with the two first papers reflecting my education in physical Geography and the last two pa-
persaswellastheresearchproposalreflectingmyswitchtomolecularecology .
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