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The Yunnan Biodiversity Hotspot Project __________________________________________________________________________________________________________________________________________________________________

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Impressum

Unpublished Symposium Volume

International Symposium

“The Yunnan Biodiversity Hotspot Project – Program, Objectives, and Application”

April 02 – 06 2016, Dresden, Germany

Copyright: Senckenberg Natural History Collections Dresden & Technical University Dresden, Germany

Edited by: Lutz Kunzmann (Senckenberg)

Cover and project logo: Markward Fischer (Senckenberg)

Maps: Susann Stiller (Senckenberg)

Layout: Felix P. Herrmann (TU Dresden)

International Symposium, Dresden, April 02–06 2016 __________________________________________________________________________________________________________________________________________________________________

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Content

Program 3

List of participants 7

Scientific committee / Organizing team 11

Preliminary Project Outline and Drafts of Group Proposals 12

A. Brief Project Outline 12

B. Drafts of Superordinated Project Proposals, Group Proposals and Abstracts of

Subprojects 13

Project summary

0. Synthesis and integration 15

0.1. Additional overview presentations 15

1. The physical basis 16

1.1. Current state of geo- and climate diversity 16

1.2. Uplift history and climate reconstruction 17

2. Evolution of Biodiversity – the fossil record 18

2.1. Standard section/drill core to study vegetation (pollen), climate (incl. glacial

history) using a multiproxy approach (incl. isotopes, biomarkers, etc.) 18

2.2. Neogene biota: paleo-biodiversity, paleoecology, biomarkers, paleoclimate,

stratigraphy, biogeography, altitudinal zonation 21

2.3. The hominoids 30

3. Evolution of Biodiversity – the genomic record 32

3.1. Plant groups 32

3.2. Invertebrate groups 42

3.3. Vertebrate groups 48

4. Present-day diversity distribution in Yunnan – a macroecology and

modelling analysis of present-day patterns 49

5. Future threats 53

Field trips 54

Map with field trip destinations 55

Practical paleontological field work part 1 56

Practical paleontological field work part 2 72

Useful information 81


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Program

Friday & Saturday – Arrival and Welcome

Date Time Program item (early arrivals) Program item (late

arrivals)

April 1

afternoon Arrival at airport Dresden,

transport to hotel, check-in

19:00 – 22:00 Dinner

April 2

8:30 – 13:00 Visit Senckenberg Natural History

Collections Dresden (collections,

labs)

Arrival at airport or main

station Dresden, hotel

check-in

13:00 – 14:00 Lunch

14:00 – 18:00 Transport to hotel, afternoon:

sightseeing and shopping

possibility

Afternoon: sightseeing and

shopping possibility

18:00 – 22:00 Welcome Party in the exhibition building Japanese Palace with

visit to the Senckenberg special exhibition 'Tibet'

Sunday - Symposium I

April 3 Workshop: oral presentations and discussion (day 1)

8:00 – 9:00 Registration, upload presentations

9:00 – 9:20 Welcome talks (Lutz Kunzmann, Volker Mosbrugger, Zhou Zhe-

Kun, Technical University Dresden representative)

9:20 – 9:40 Zhe-Kun Zhou: Evolution and extinction of some plants and

their response to Asia Monsoon climate.

9:40 – 10:00 Sun Hang: xxx

10:00 – 10:30 Volker Mosbrugger/Zhe-Kun Zhou/Lars Opgenoorth/Georg

Miehe: The Yunnan Biodiversity Hotspot Project: state-of-the-art

and application procedure

ongoing 1.2. Uplift history and climate reconstruction, geo- and climate

diversity

- Andreas Mulch


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- Bob and Teresa Spicer / Torsten Utescher

- Cheng-Long Deng

11:00 – 11:20 Tea and coffee break

ongoing 2.1. Standard section/drill core to study vegetation (pollen),

climate (incl. glacial history) using a multiproxy approach (incl.

isotopes, biomarkers, etc.)

- Torsten Utescher

- Wei-Ming Wang

- Martina Stebich

- Christian Rolf

ongoing 2.2. Neogene leaf floras: stratigraphy, taxonomy, biogeography,

climate and altitudinal zonation

- Lutz Kunzmann

- Tao Su

- Torsten Wappler

- Karolin Moraweck

- Ulf Linnemann

ongoing The hominoids

- Christine Hertler

- Tina Lüdecke

ongoing Present-day diversity distribution in Yunnan – a macroecology

and modelling analysis of present-day patterns

- Alice Hughes

- Matthew Forrest

13:00 – 14:00 Group photo and Lunch

14:00 – 15:30 Evolution of Biodiversity / the genomic record

3.1. Plant groups

- Georg Miehe

- Anita Roth-Nebelsick/Wilfried Konrad/Stefan Wanke

- Jian-Quan Liu/Lars Opgenoorth

3.2. Invertebrates

- Joachim Schmidt/Lars Opgenoorth

- Sonja Wedmann

- Martin Brändle

3.3 Vertebrates

- Martin Päckert


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16:00 – 18:00 "World Café" (round-table discussion):

- Discussion within the project groups (organized by PI)

- Networking between project groups

18:00 – 18:30 Summary first day: Volker Mosbrugger & Zhe-Kun Zhou

18:30 – 19:00 Break (hotel)

19:30 – 22:00 Dinner (Italian restaurant)

Monday - Symposium II

April 4 Workshop: discussion and synthesis

(day 2)

8:30 – 10:00 Presentation of results of “World Café” (PIs) and general

discussion


10:20 – 12:00 Synthesis and coordination of project groups (Volker

Mosbrugger, Zhe-Kun Zhou, Torsten Utescher, Georg

Miehe)

12:00 – 13:00 Lunch

13:00 – 14:30 Final discussion and outlook (Volker Mosbrugger, Zhe-Kun

Zhou)


15:00 – 18:00 Visit botanical garden of the Institute of Botany, Technical

University Dresden (guide: Barbara Ditsch, curator)

18:30 – 19:00 Break (hotel)

19:30 – 22:00 Symposiums dinner (Spanish restaurant) with Closing remarks

(Volker Mosbrugger)


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Tuesday - Field trip I

April 5 Practical paleontological field work and group discussions

(day 1)

8:00 – 10:00 Travel to MIBRAG opencast mine Vereinigtes Schleenhain

10:00 – 14:00 Field work in Eocene and Oligocene sections (with lunch break)


14:30 – 16:30 Return to Dresden (hotel)

16:30 – 19:00 Break (hotel)

19:30 – 22:00 Dinner (Saxonian restaurant)

Wednesday - Field trip II

April 6 Practical paleontological field work and group discussions

(day 2)

7:00 – 9:00 Travel to VATTENFALL opencast mine Welzow-Süd

9:00 – 13:00 Field work in Miocene sections

13:00 – 14:00 Lunch

14:30 – 16:30 Return to Dresden (hotel)

16:30 – 19:00 Break (hotel)

19:30 – 22:00 Dinner (historical Dresden restaurant) and Farewell

Thursday - Departure

April 7

morning Hotel check-out, transport to airport


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List of participants

Martin BRÄNDLE [email protected]

Philipp University of Marburg, Marburg, Germany

Cheng-Long DENG [email protected]

Institute of Geology and Geophysics, CAS, Beijing, PR China

Matthew FORREST [email protected]

Senckenberg, Frankfurt, Germany

Christine HERTLER [email protected]

Heidelberg Academy of Sciences and Humanities, Heidelberg, Germany

Sylvia HOFMANN [email protected]

Natural History Museum Erfurt, Erfurt, Germany

Yong-Jiang HUANG [email protected]

Kunming Institute of Botany, CAS, Kunming, PR China

Alice HUGHES [email protected]

Xishuangbanna Tropical Botanical Garden, Yunnan, PR China

Phyo Kay KHINE

Philipp University of Marburg, Germany

Annette KLUSSMANN-KOLB [email protected]


Wilfried KONRAD [email protected]

Technical University of Dresden

Lutz KUNZMANN [email protected]

Senckenberg, Dresden, Germany

Zhimin LI [email protected]



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Ulf LINNEMANN [email protected]


Jian-Quan LIU [email protected]

School of Life Science, Lanzhou University, Lanzhou, PR China

Tina LÜDECKE [email protected]


Georg MIEHE [email protected]


Karolin MORAWECK [email protected]


Volker MOSBRUGGER [email protected]


Andreas MULCH [email protected]


Christoph NEINHUIS [email protected]

Technical University of Dresden, Germany

Lars OPGENOORTH [email protected]


Martin PÄCKERT [email protected]


Christiane RITZ [email protected]

Senckenberg, Görlitz, Germany

Christian ROLF [email protected]

Leipniz Institute for Applied Geophysics, Hannover, Germany


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Anita ROTH-NEBELSICK [email protected]

State Museum of Natural History Stuttgart, Stuttgart, Germany

Joachim SCHMIDT [email protected]

University of Rostock, Rostock, Germany

Teresa SPICER [email protected]

Institute of Botany, CAS, Beijing, PR China

Martina STEBICH [email protected]

Senckenberg, Weimar, Germany

Tao SU [email protected]


Hang SUN [email protected]


Torsten UTESCHER [email protected]

Senckenberg, Frankfurt; University of Bonn, Germany

Truong Van DO [email protected]

Vietnam National Museum of Nature, Ha Noi, Vietnam

Wei-Ming WANG [email protected]

Institut of Geology and Palaeontology, Nanjing, PR China

Yun WANG [email protected]


Stefan WANKE [email protected]

Technical University of Dresden, Germany

Torsten WAPPLER [email protected]

University of Bonn, Bonn, Germany


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Sonja WEDMANN [email protected]


Karsten WESCHE [email protected]


Zhe-Kun ZHOU [email protected]



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Scientific committee

Volker Mosbrugger Senckenberg Society for Nature Research, Frankfurt/M., Germany

Georg Miehe Philipp University of Marburg, Marburg, Germany

Torsten Utescher Senckenberg Society for Nature Research Frankfurt/M., / Universität

Bonn, Germany

Lutz Kunzmann Senckenberg Natural History Collections Dresden, Germany

Christoph Neinhuis Technical University Dresden, Germany

Karolin Moraweck Technical University Dresden / Senckenberg Natrual History

Collections Dresden, Germany

Zhe-Kun Zhou Xishuangbanna Tropical Botanical Garden (CAS), Yunnan, PR China

Tao Su Xishuangbanna Tropical Botanical Garden (CAS), Yunnan, PR China

Sun Hang Kunming Institute of Botany (CAS), Yunnan, VR China

Organizing team

Sigrid Beutner Senckenberg, Dresden, Germany

Mina Breuer Technical University of Dresden, Germany

Franziska Ferdani Senckenberg, Dresden, Germany

Markward Fischer Senckenberg, Dresden, Germany

Denise Hennig Senckenberg, Dresden, Germany

Felix P. Herrmann Technical University of Dresden, Germany

Carola Kunzmann Senckenberg, Dresden, Germany

Lutz Kunzmann Senckenberg, Dresden, Germany

Karolin Moraweck Senckenberg, Dresden, Germany

Madeleine Streubig Senckenberg, Dresden, Germany

Susann Stiller Senckenberg, Dresden, Germany

Birgit Walker Senckenberg, Dresden, Germany


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Preliminary Project Outline and Drafts of Group Proposals

How orogenesis drives evolution: Processes, status & trends of geo-biodiversity in

the Himalayan-Yunnan Hotspot.

DFG Trans-Regio: Dresden, Frankfurt, Marburg

A. Brief Project Outline

0 Synthesis and integration

1. The physical basis

1.1 Current state of geo- and climate diversity

1.2 Uplift history and climate reconstruction

2. Evolution of Biodiversity – the fossil record

2.1 Standard section/drill core to study vegetation (pollen), climate (incl. glacial history)

using a multiproxy approach (incl. isotopes, biomarkers, etc.)

2.2 Neogene biota: paleo-biodiversity, paleoecology, biomarkers, paleoclimate,

stratigraphy, biogeography, altitudinal zonation

2.3 The hominoids

3. Evolution of Biodiversity – the genomic record

3.1 Plant groups

3.2 Invertebrate groups

3.3 Vertebrate groups

4. Present-day diversity distribution in Yunnan – a macroecology and modelling analysis

of present-day patterns

5. Future threats


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B. Drafts of Superordinated Project Proposal, Group Proposals and Abstracts of

Subprojects

How orogenesis drives evolution: Processes, status & trends of geo-biodiversity in

the Himalayan-Yunnan Hotspot.

Joerg Bendix, Volker Mosbrugger, Georg Miehe, Lars Opgenoorth

Summary

On a global scale mountains are the most important harbor of biodiversity best exemplified

by the fact that all six major diversity centers surpassing ca. 5000 vascular plant species per

10 000 km² are located in mountain areas(Barthlott et al., 1996). There are two possibilities

for this link – either mountains act as refugia for species from the surrounding area or they

are themselves species pumps. In most cases it will be a combination of both. Nevertheless,

the role of orogenesis as a driver for evolutionary processes has long been accepted in

evolutionary ecology. The drivers behind this connection are manifold and include (i)

topographic complexity which increases geographic and thus genetic barriers, as well as

area, hydrological regime and others, (ii) climatic complexity including microclimatic effects

such as exposition and slope, as well as larger scale effects such as LEE effect and MEE

effect, all of which directly increase the available niche space, (iii) geological complexity by

driving habitat structure e.g. through influencing soil conditions, and erosional processes, (iv)

orogen stability – or more generally speaking – habitat stability. In this proposal we will

summarize all these effects under the term geodiversity. The effects of geodiversity vary in

time – depending on how long orogenesis has been taken place with what specific features –

as well as in space – as the latitudinal position has significant influence on how orographic

complexity acts upon biological processes and thus biodiversity.

Geodiversity is an abstract concept that is not easily measured and even less easily

translated into biotic processes because so many different drivers are superimposed over

each other. Furthermore, the effect each of these drivers have on different biological units is

diverse as well. For example, while relatively small elevational differences can be significant

barriers for wingless ground beetles directly driving evolutionary processes and eventually

speciation, flying beetles, birds or mammals will not be affected by them. Thus topographic

complexity will have much different effects and effect-thresholds depending on the life history

traits of a given taxon. One of the overarching goals of this research proposal therefore is to

disentangle important parts of these complex mechanisms and ultimately to derive measures

for geodiversity that will allow to model ‘across taxa biodiversity’ in mountain systems of

the world and moreover, determine how biodiversity as well as evolutionary processes can

be preserved in the future. More specifically, we want to


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Derive high resolution descriptors of current geodiversity through remote sensing and

spatial modelling

Derive indices for past orographic complexity by analysing the biologically relevant

history of the orogenesis of the Himalaya-Tibet Orogen

Disentangle the specific drivers responsible for the creation of the Himalaya-Yunnan

Hotspot specifically comparing diverse evolutionary and spatial histories of diverse taxon

groups

Date when the Himalaya-Yunnan Hotspot first appeared in Earth History

Characterize the extant large scale biodiversity patterns by means of macroecology

and compare them to the historical processes.

Assess future threats to diverse groups of biodiversity and ecosystems in the

Himalaya-Yunnan Hotspot


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0. Synthesis and integration

Developing a Memorandum of Understanding / Cooperation Agreement between German

institutions, and XTBG (CAS), KIB (CAS) of PR China (Volker Mosbrugger, Zhe-Kun Zhou,

Sun Hang, Georg Miehe)

Developing a common database.

0.1. Additional overview presentations

Evolution and extinction of some plants and their response to Asia Monsoon climate.

Zhe-Kun Zhou, Xishuangbanna Tropical Botanical garden, Chinese Academy of Sciences,

Yunnan, PR China

Seasonal precipitation is the main characteristic of the Eastern Asian monsoon: almost 80

percent of the annual precipitation is concentrated during the rainy season, usually from May

to October. The Asian monsoon region experiences wet summers and very dry winters and

springs. The Asian monsoon system was established during the Late Miocene in association

with the uplift of the Qinghai-Xizang Plateau. The following scientific questions are

addressed: how does the Asian monsoon impact on biodiversity and how do the plants

respond to the dry winter and spring. We propose the following hypothesis: Due to the

seasonality of the monsoon climate, some physiological processes of plants such as

germination and growth of seedlings would suffer from dryness in winter and spring, causing

their distribution area to shrink or even leading to their extinction. In order to answer these

questions, five research goals are designed in the current project: First, to reconstruct the

paleoclimate and intensity of the monsoon using stable isotopes from sediments of middle

Miocene to Pleistocene age. Concurrently the paleoclimate will be reconstructed using plant

fossil fruits and seeds. These independent approaches will be used to cross-check one

another. Second, the fossil histories of Metasequoia and other relict plants will be studied

and their paleoclimatic signal reconstructed. Third, research into the differentiation of Pinus

yunnanensis and Pinus kesiya var. langbianensis and their response to the monsoon climate

will be carried out. Molecular analysis, wood anatomy, stable isotopes of wood and leaves

will be involved in this mission. Fourth, seed biology and experiments of controlled cultivation

of Cedrus, Metasequoia, oaks with wider distributions and limited distributions will be

undertaken. Fifth, fossil history and phenology of Quercus subgenus Cyclobalanopsis will be

carried out in order to test if extinction of some oaks such as Quercus sichouensis is caused

by the dry winter.


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1. The physical basis

1.1. Current state of geo- and climate diversity

High resolution digital elevation model“

Meteorological data of Yunnan (incl. data from MOST Project 973: Three River Gorges; and

data from China Meteorological Forcing Dataset)

Volker Mosbrugger and Joerg Bendix will develop a concept

Zhou Zhekun will find out, which data are publicly available


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1.2. Uplift history and climate reconstruction

Cenozoic climate and uplift in Yunnan – evidence from the palaeobotanical record

Torsten Utescher1, Volker Mosbrugger1, Su Tao2, Zhou Zhe-Kun2

1Senckenberg Research Institute, Senckenberganlage 25, 60325 Frankfurt am Main,

Germany

2Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun,

Mengla, Yunnan 666303, China

Climate data obtained from a total of 29 Yunnan palaeofloras (macro-remains, pollen) by

using the Coexistence Approach (29 floras, new calculations) and CLAMP (8 floras, literature

data) provide a first insight into Cenozoic temperatures in general, inferred altitudinal

changes and time resolution of the macro-records. Temperature data compiled for the

southern part of Yunnan reflect the large scale global trend of the late Neogene Cooling. The

record obtained for northern Yunnan can hardly be interpreted, most probably due to

differential uplift processes in that area. When relating Cenozoic temperatures inferred from

sites presently located at low and mid- altitudes to modern values it is shown that this

continental part was not significantly warmer compared to today, partly it might have been

even cooler. For most sites, presently located at higher altitudes pointed positive temperature

anomalies with respect to present are obtained. Thus, it has to be assumed that various

parts of Yunnan were subject to more recent uplift.

As is shown by first preliminary results the palaeobotanical record of Yunnan can contribute

substantial data on the uplift history of Yunnan while the high degree of overlapping of CA

and CLAMP-derive data testifies the coherence of the data obtained. However, there still

exist large gaps in the macrofloral record so far available comprising most of the

Palaeogene, the Middle Miocene, and the Early Pliocene. In order to study regional climate

dynamics in the context of landscape evolution at higher resolution suitable pollen records

are required and other proxies have to be included.


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2. Evolution of Biodiversity – the fossil record

2.1. Standard section/drill core to study vegetation (pollen), climate (incl. glacial

history) using a multiproxy approach (incl. isotopes, biomarkers, etc.)

Perspectives on Late Cenozoic vegetation and climate study in Yunnan based on

standard sections and cores

Wei-Ming Wang

Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing

210008, P. R. China, [email protected]

The Yunnan-Guizhou Plateau, located adjacent to the southeast of the Tibetan Plateau, has

a complicated topography, diverse climate conditions, and high plant diversity. Its geological

history has been strongly affected by the uplift of the Tibetan Plateau in the Cenozoic. Late

Cenozoic is a crucial period in the development of the local vegetation in Yunnan.

During the past decades, many studies focused on the the late Miocene deposits, such as

Xiaolongtan, Lincang and Xianfeng floras. There were severe environmental changes

occurred both on the lands and in the oceans at that time, for example, the development and

spread of C4 grasses, the aridification of the interiors of continents, the expansion of open

landscapes, and the development and evolution of the East Asian monsoon.

The high-resolution pollen study of the lake sediment in Heqing Basin is the only long core

which displays the vegetation history of the mountains around the basin since 2.78 Ma B.P. It

shows that the increase or decrease of vertical vegetational belts and the changes in the

components of vegetational belts are controlled by the tectonic uplift of mountains and the

climatic changes. Core from Lake Erhai also exhibits the local vegetaion and climate

changes since the latest Pleistocene.

Yunnan was once suggested to be one of the most possible areas for the origin of rice

agriculture. During the past decade, many Palaeolithic sites were excavated, which gives us

chances to reconstruct the vegetation and climate background for the early human beings.

Case studies were carried out on several sites. Among them, the site at Xiangbidong is the

first Palaeolithic cave relic, which is located in the Hengduan Mountains with complex

landforms and diverse natural landscapes. Pollen study reveals vegetation and environment

changes in relation with human activies.

It is anticipated that we will have a much more complete knowlege on the Late Cenozoic

vegetation and climate history in Yunnan, if we get further improved dating for these

Neogene floras, and some continued long profiles for comprehensive study with multi-

approaches.


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Improving chronostratigraphic frame by palaeomagnetic and rock magnetic

investigations

Christian Rolf, Leipniz Institute for Applied Geophysics, Hannover, Germany

Ulrich Hambach, University of Bayreuth, Germany

The potential of rock and palaeomagnetism in stratigraphic and chronometric investigations

was intensely discussed during the previous meetings in Xishuangbanna (2013) and

Kunming (2015) where we have reported jointly with our Chinese colleagues about the broad

application possibilities of rock and palaeomagnetic investigations. All sub-projects with

stratigraphic approaches in the biodiversity hotspot program need a chronostratigraphic

frame of the pre-hotspot world in Yunnan. The palaeomagnetic methods should be able to

serve this request, as our Chinese colleagues have already shown in excellent publications

concerning magnetic polarity stratigraphy (MPS) of the late Neogene in Yunnan.

Not only the changing polarity of the Earth’s Magnetic Field (EMF) but also its variation in

intensity is recorded in rocks. In lithologically homogenous sediments the so called relative

palaeointensity (RPI) is recorded. Correlating these RPI data with the quite well-known

variation of the EMF during the Upper Neogene provides an independent tool for

stratigraphic dating with a temporal resolution in the order of ten thousand to thousands of

years. Great variability in lithology and magnetic mineralogy are limiting factors. Suitable

outcrops or drill cores with homogenous lithology are mandatory.

Time series analyses of down-section/down-hole rock-magnetic proxies -like magnetic

susceptibility-can provide a temporal frame in much higher resolution than magnetic polarity

stratigraphy (e.g. Milanković cycles). Cyclostratigraphy alone provides no absolute data, but

relative time estimates and need anchor points. This approach may be applied on large

outcrop sections or drill cores with wider time windows (> 100 ka).

In addition the changing properties of magnetic minerals give information about the temporal

and spatial environmental dynamics (climate, provenance, depositional systems and

diagenesis, etc.). Rock magnetic investigations are also mandatory to back the quality of

MPS and RPI recording.

Furthermore, we think it could be worth to investigate rock magnetic parameters as proxy

indicators for volcanic crypto-tephras. They will be excellent time markers, if recorded, and if

a tephra chronology of the area under investigation can be established. Following the

statements of Chenglong up to date tephra layers were found only in few sedimentary basins

of late Cenozoic age in Yunnan. Suggested volcanic activities of late Cenozoic age in

western Yunnan, however, makes the search of volcanic tephra layers in Yunnan basins

highly promising.


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Cheng-Long Deng proposed acooperation on magnetostratigraphy (including rock

magnetism as proxies for quality of the magnetostratigraphic results) secular variation,

“palaeointensities” and the search for tephra layers. In addition we would like to focus on

younger (Plio-/Pleistocene) stratigraphic intervals in outcrops or drill cores in order to explore

the record of the Plio-/Pleistocene environmental change.

In case drilling projects will be realised we can offer the LIAG downhole logging equipment to

get geophysical borehole data in high resolution (for examples magnetic susceptibility,

natural gamma ray etc.). These data can be analysed by time series/cyclostratigraphic

methods to improve the age depth models of the sediments recovered.

Our experience in applying for funding from the DFG for new drilling projects tells us, that it is

advisable to have good quality data in advance. In case already existing and accessible

boreholes are available, our team would come on own funds to China to acquire data and

test their quality.


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2.2. Neogene biota: paleo-biodiversity, paleoecology, biomarkers, paleoclimate,

stratigraphy, biogeography, altitudinal zonation

Deciphering a “pre-hotspot” world: the palaeoenvironmental perspective on Neogene

biota

Key words: palaeobiodiversity, palaeoherbivory, molecular biomarkers, palaeoecology,

palaeoclimate fluctuations, palaeoatmospheric composition

Group members:

Chinese side: Su Tao, Huang Yong-Jiang, Xie Sanping, Shi Gongle , Deng Cheng-Long

German side: Lutz Kunzmann, Karolin Moraweck, Torsten Wappler, Eva Niedermeyer, Ulf

Linnemann, Anita Roth-Nebelsick

Outline:

- Project group proposal: key questions and summary

- Project 1: Evolution of ecophysiological “fingerprints” of Neogene vegetation in

Yunnan (China) under multicausal influence of monsoon, mountain uplift, climate

change, atmospheric composition and forming topographies. (Lutz Kunzmann, Su

Tao, Karolin Moraweck, Gongle Shi, Yong-Jiang Huang, Eva Niedermeyer, Anita

Roth-Nebelsick)

- Project 2: Structural changes of herbivory since the Neogene in response to climatic

fluctuations from Yunnan (China). (Torsten Wappler, Su Tao)

- Project 3: Neogene climate change in Yunnan (China) inferred from molecular

biomarkers. (Eva Niedermeyer/Karolin Moraweck/Lutz Kunzmann/???)

- Project 4: “Dating/stratigraphy”. (Ulf Linnemann/Cheng-Long Deng/ …)

Project group proposal: key questions and summary

The present biodiversity in Yunnan’s mountain regions and its proximities in other Chinese

provinces as well as in neighboring countries is discussed to be the result of a multicausal

evolution. Main drivers certainly were mountain uplift, global climate change, establishment

of a monsoonal regime and constitution of the landscape after/during the Pleistocene by

glacigenous processes. The first key question concerns the crucial driver of the

establishment of the Yunnan Biodiversity Hotspot.

Several results of biodiversity studies in extant plant and animal groups tentatively give rise

to the hypothesis that the biodiversity hotspot isn’t that old as one could perhaps conclude on

the age of the appearance of the monsoonal regime, i.e. at least early Miocene or late

Eocene. In fact, by studying Neogene biota and environments the “pre-hotspot” scenario


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could be deciphered as well as the underlying circumstances for the evolution of a

biodiversity hotspot.

Therefore, another key question of the project group is to test whether pre-Pleistocene floras

and herbivory data reveal any information of the rising biodiversity hotspot. With respect to

the megafloras several (late) Miocene and Pliocene floras will be involved in the studies.

Based on data on their systematic diversity and data of plant-arthropod interactions their

palaeoenvironments, palaeoclimatic conditions and palaeoatmospheric background will be

reconstructed. These reconstruction will be condacted based on published data as well as by

new studies. Molecular biomarkers are an essential tool for reconstructing ancient climate

from horizons where fossil biota are absent. The coupling of these results, i.e. proxy data

derived from several floras from two distinct time intervals, data from plant-arthropod

interactions and from fossil molecular biomarkers, is a novelty and allows the

consideration of the different factors in a multiproxy analysis.

If the hypothetical scenario is true, the project bundle can presumably explain the “pre-

hotspot world” of that region as the starting point and place of the evolution of a hotspot. In

particular we want to solve these questions:

(1) Why was this region predestined for the evolution of a hotspot?

(2) Which biotic and abiotic factors were essential for the “rise” of the hotspot?

(3) What can we learn from fossil floras and fossil herbivory about the hotspot’s starting

point and early evolution?

(4) Which climatic conditions and/or fluctuations and paleo-atmospheric conditions

favored the evolution of the hotspot?

Selected Neogene floras of Yunnan in the project

Tao Su, Xishuangbanna Tropical Botanical Garden, CAS, Yunnan, PR China

The tectonic activities in southeastern margin of the Qinghai-Tibetan Plateau created more

than 200 Neogene basins in Yunnan Province. Among them, plenty of well preserved fossil

floras have been uncovered especially during the last decade, which provide good

opportunity to study the biodiversity history under paloenvironmental changes in Yunnan.

Previous studies with focus on Neogene floras indicated that, the monsoon intensified

gradually since the Neogene in Yunnan and had profound influence on shaping modern

biodiversity. However, our understanding about the diversification pattern is far from being

enough comparing to the abundant fossil floras as well as extant high diversity here. In this

project, we will select several megafloras ranging from the early Miocene to the late Pliocene

as materials for study, i.e., the early Miocene Jinggu flora (S Yunnan), the middle Miocene

Wenshan flora (SE Yunnan) and Zhenyuan flora (SW Yunnan), the late Miocene Xiaolongtan


23

flora (SE Yunnan) and Lvhe flora (C Yunnan), the Pliocene Heqing flora (NW Yunnan) and

Weixi flora (NW Yunnan). According to the preliminary investigation, all these floras are not

only in excellent preservation condition, but also rich in plant diversity preserved as leaves,

fruits/seeds, flowers, and wood. Fossils from these floras will be collected, and their

systematic positions will be determined with morphological comparison to modern plants.

Moreover, new fossil sites in Yunnan will be explored according to this project.

Project interactions:

Coupling of morphometric, anatomical and ecophysiological plant traits: Climatic and

environmental driven changes in biodiversity patterns are often preserved in fossil floras.

Studying the composition of these plant assemblages as well as focusing on single taxa and

their adaptions in anatomy and morphology to extant climatic and environmental conditions

will help to understand the development and distribution of this hotspot region. The

morphometric reconstruction of fragmented fossil leaves delivers quantitative data about leaf

area, length, width and related data, which enable to conclude environmental data, like water

availability or temperature and heat conditions on the leaves surface. Changing climatic and

environmental conditions might have been drivers for changing compositions of plant

assemblages and perhaps led to specification of single genuses. Alongside the consideration

of morphometric traits, also anatomical adaptions can be evident for ecological and climatic

changes, as trichome density (TD) and stomata density (SD) are coupled with water use

efficiency and palaeoatmospheric conditions, which can be reconstructed using the gas

exchange model. The procedure for collecting and evaluating plant–insect associational data

is derived from an explicit classification of insect-mediated damage used by the PIs and

colleagues. For the Yunnan data, we will examine on slabs each identifiable plant fragment

greater than ca. 0.5 cm2, followed by a database record of unique specimen number, plant

identification, DTs (if any), image or line drawings of the damage (if any), repository location

and comments for each specimen. This spectrum of damage types (DTs) is based on life-

plant response to insect feeding using four diagnostic criteria. The presence or absence and

type of DT data are tabulated for each leaf per locality, together with host-plant identification,

habitat type and stratigraphic position, for the Neogene floras in Yunnan. The data will be

expressed as stratigraphic plots listing DTs, the percent of leaves damaged (frequency data),

the number of DTs per leaf (diversity data), and will be partitioned by plant-host and

organ/tissue specificity using several lines of evidence. Attention will be directed to the

temporal and spatial damage-type composition of the dominant plant hosts to assess the

evolution of herbivore partitioning of tissues.

Connecting factors to other project groups:


24

Observed variations in climate conditions probably represent intrinsic change and not shifts

resulting from tectonic uplift of the area. We expect that our findings will have major effects

on understanding interorganismic palaeoecological relationships and increase understanding

of the determinative role that past environmental changes have on those relationships, and

analogous, modern, ecological changes. The project is connected to the groups 3 and 7.

The knowledge of plants taxonomy and their composition in these fossil plant assemblages is

connected to group 5, analysing the diversity evolution and present day distribution. As

mountain uplift is connected with climate changes, the plant derived palaeoatmospheric and

palaeoclimate data can be coupled and verified using the uplift and climate data obtained by

group 3, using isotopes and other proxies.


25

Project 1: Evolution of ecophysiological “fingerprints” of Neogene vegetation in

Yunnan (China) under multicausal influence of monsoon, mountain uplift, climate

change, atmospheric composition and forming topographies

Among Neogene biota in Yunnan (China) plant megafossil assemblages occur widespread in

intracontinental sedimentary basins. Most of these floras are rather divers both in taxa and

plant parts / organs. They are therefore predestined for reconstruction of terrestrial

ecosystems through time. Main factors which influenced the vegetation are climate change

including monsoon regimes, mountain uplift, plate tectonics and constitution of the

landscape.

The project will identify an amount of morphological, morphometric, anatomical and

phytosociological data from the fossil leaf assemblages due to reveal an ecophysiological

“fingerprint” of a flora. These “fingerprints” show taxon-independent signals of environmental

and atmospheric changes through time. Two stratigraphic levels are chosen for the study,

late Miocene and Pliocene, because from here well-preserved megafloras containing a

diverse leaf component are known.

The fossil leaf remains themselves reveal climatic and atmospheric signals which could be

quantified by using several approaches. Climatic signals are calculated by CLAMP and CA,

whereas atmospheric signals, i.e. pCO2, are calculated using the mechanistic gas exchange

model. From this a scenario for changing climatic and atmospheric conditions under the

influence of mountain uplift and plate tectonics will be developed. The most important

questions are: (1) whether an increase in biodiversity detectable from the floras is visible or

not; (2) could an increase in diversity possibly lead to the current hotspot; (3) which were the

main drivers of increasing diversity; (4) when did main increases in diversity occur and if

vegetation changes did not necessarily coincide with an increase in diversity of plants (5)

what factors prevent the evolution of a hotspot.

Presumably an integrated analyses of Neogene floras lead to the conclusion that that the

main biodiversity rise appeared later than Pliocene. Thus, the evolution of the Neogene

vegetation gives evidence for a pre-hotspot diversity.


26

Project 2: Structural changes of herbivory since the Neogene in response to climatic

fluctuations from Yunnan (China)

Vascular plants together with insects contribute substantially to the Earth's rich biodiversity.

Their interactions constitute a complex and intricate trophic network in terrestrial ecosystems;

therefore, the patterns of plant-arthropod interactions in geological times are pivotal to

understand the evolution of terrestrial ecosystems. The origin of these trophic networks can

be traced back for more than 400 million years ago (Misof et al., 2014). However, research

on changes in the food web structures in ecological systems on the basis of

paleaeontological data is only at its beginnings and underlines our need to understand the

impacts of climate change on these systems. Still some progress has been reached in the

last decades and by now, the fossil compressions of dicotyledonous plant leaves represent

our main source of evidence of changes of these interactions (e.g., Labandeira, 2013). Most

analyses and results presented in previous studies have shown that the relationships are

more complex than generally anticipated (e.g., Wappler et al., 2012; Su et al., 2015), and

that most observed processes act in network structures rather than in isolation (Wappler et

al., 2015).

The study of plant-arthropod interactions faces many fundamental challenges. The

plants have to be identified from often-fragmentary material, and the arthropods interacting

with plants have to be inferred even only from the traces they had left on leaves before they

fossilized. The records of fossil insects themselves are still relatively scarce (Wappler, pers.

observ. 2015). On the contrary, fossilized leaves of Neogene age are relatively common at

many sites (e.g., Su et al., 2013; Zhang et al., 2015) and frequently large proportions of

leaves bear well-preserved traces of arthropod feeding (Wappler, pers. observ. 2015).

The work employing the plant-arthropod interactions for better reconstruction of

palaeoenvironments and for study of reactions of palaeoecosystems to climatic fluctuations

poses more constraints on our choice of our study material (e.g., sites – palaeoecosystems).

The quantitative reconstructions require large collections of fossil leaves (and also sufficient

numbers of those with arthropod damage) and the compared palaeoecosystems have to

differ only in as few characteristics as possible. This means that they should be close to each

other geographically and host as similar taxa as possible, so that the differences in plant-

arthropod associations can be attributed to the differences in climate and not to other

palaeogeographical or palaeoecological conditions. The less we can comply with these

conditions the more drastic changes in climate we have to focus on in order to attain at least

some certainty about ascribing the change in palaeoecosystem biodiversity and structure to

the effect of climate. Finally, it is also important from the viewpoint of comparison to current

ecosystems to find a geological era with climate fluctuations sufficiently similar to our current

ecosystems.


27

The Neogene seems to be very perspective from the point of the above-mentioned

conditions for study of impacts of climatic fluctuations on past and recent ecosystems. For

example, Miocene climate fluctuations were found similar in form to later Plio-Pleistocene

fluctuations resulting in cycles known as the Ice Ages. Generally speaking, the main triggers

to Miocene climate fluctuations are thought to be the emergence of the Antarctic ice cap

following the formation of Circum-Antarctic current and concurrent decline of

circumequatorial circulation due to closing of Tethys Ocean. Regionally, more important

agents were the large continental uplift due to collision of Afro-Arabian and Indian plates in to

Eurasia (Sun et al., 2015), coupled with the evolution of the Asian monsoon system (Su et

al., 2013). Overall, this was also coupled by long-term fluctuations in CO2 concentration (e.g.,

Kürschner et al., 2009). The use of Neogene plant-arthropod interactions harbours one more

important advantage – patterns of DT diversity and agent host specificity are already very

similar to recent conditions as evidenced by insect damage on an oak group (Su et al.,

2015). Due to all these reasons, Neogene represents an extremely interesting and important

period in relation to study of the way the ecosystems cope with climate fluctuations.

Despite this suitability, the studies that focus on this period are still in its very

beginnings (e.g., Prokop et al., 2010; Wappler 2010; Knor et al., 2013, 2015). Some of the

world’s best-developed deposits of Neogene sediments are those of Yunnan Province,

located at the southeastern boundary of the Qinghai-Tibetan Plateau. Until now, more than

twenty Neogene megafloras have been reported from different localities of Yunnan (Li, 1995;

Ge and Li, 1999). More importantly, the intensification of the monsoonal climate caused by

the uplift of the QTP became more prominent since the Miocene, characterizing by the drier

winter and wetter summer. Early studies indicate that this climate trend has largely

influenced the plant diversity in Yunnan (Su et al., 2013; Zhang et al., 2015). However, the

plant-insect interactions in terrestrial systems are still largely unknown there.

The study of the plant-insect associations from Neogene sediments located in the

Yunnan Province gives an opportunity to confirm some important predictions with regard to

climatic fluctuations in the light of the elevation and extent of the Himalaya–Tibetan plateau

(HTE). Together with modern samples from this biodiversity hotspot, the project aims to

understand: (1) the evolutionary pattern of insect damage on specific plant groups, e.g.,

Fagaceae, and Betulaceae; (2) the plant-insect interactions among paleofloras in respose to

paleoenvironmental changes there, such as the intensification of the monsoonal climate; and

(3) the mechanism of plant-insect interactions in structuring terrestrial ecosystems under

dramatically Neogene paleoenvironmental changes.

References:

Currano, E.D., Labandeira, C. and Wilf, P., 2010. Fossil insect folivory tracks paleotemperature for six million

years. Ecological Monographs, 80, 547-567.


28

Ge, H.-R., and Li, D.-Y., 1999. Cenozoic Coal-Bearing Basins and Coal-Forming Regularity in West Yunnan.

Yunnan Science and Technology Press, Kunming.

Knor, S., Kvaček, Z., Wappler, T. and Prokop, J., 2015. Diversity, taphonomy and palaeoecology of plant-

arthropod interactions in the lower Miocene (Burdigalian) in the Most Basin in north-western Bohemia

(Czech Republic). Review of Palaeobotany and Palynology, 219, 52-70.

Knor, S., Skuhravá, M., Wappler, T. and Prokop, J., 2013. Galls and gall makers on plant leaves from the lower

Miocene (Burdigalian) of the Czech Republic: systematic and palaeoecological implications. Review of

Palaeobotany and Palynology, 188, 38-51.

Kürschner, W.M., Kvacek, Z. and Dilcher, D.L., 2008. The impact of Miocene atmospheric carbon dioxide

fluctuations on climate and the evolution of terrestrial ecosystems. Proceedings of the National Academy of

Sciences, 105(2), 449-453.

Labandeira, C.C., 2013. A paleobiologic perspective on plant-insect interactions. Current Opinion in plant biology,

16(4), 414-421.

Li, X.-X., ed., 1995. Fossil Floras of China through the Geological Ages. Guangdong Science and Technology

Press, Guangzhou.

Misof B, Liu S, Meusemann K, Peters RS, Donath A, Mayer C, et al., 2014. Phylogenomics resolves the timing

and pattern of insect evolution. Science, 346(6210), 763-767.

Prokop, J., Wappler, T., Knor, S. and Kvaček, Z., 2010. Plant-arthropod associations from the Lower Miocene of

the Most Basin in northern Bohemia (Czech Republic): A preliminary report. Acta Geologica Sinica, 84(4),

903-914.

Su T, Adams JM, Wappler T, Huang Y-J, Jacques FMB, Liu Y-S, et al., 2015. Resilience of plant-insect

interactions in an oak lineage through Quaternary climate change. Paleobiology, 41(1), 174-186.

Su, T., Jacques, F.M.B., Ma, H.-J. and Zhou, Z.-K., 2013. Fossil fruits of Ailanthus confucii from the Upper

Miocene of Wenshan, Yunnan Province, southwestern China. Palaeoworld, 22(3–4), 153-158.

Su, T., Jacques, F.M.B., Spicer, R.A., Liu, Y.-S., Huang, Y.-J., Xing, Y.-W., and Zhou, Z.-K., 2013. Post-Pliocene

establishment of the present monsoonal climate in SW China: evidence from the late Pliocene Longmen

megaflora. Climate of the Past, 9(4): 1911-1920.

Sun, B. et al., 2015. Early Miocene elevation in northern Tibet estimated by palaeobotanical evidence. Scientific

Reports, 5, 10379.

Wappler, T., Labandeira, C.C., Engel, M.S., Zetter, R. and Grimsson, F., 2015. Specialized and Generalized

Pollen-Collection Strategies in an Ancient Bee Lineage. Current Biology, 25(23), 3092-3098.

Wappler, T., Labandeira, C.C., Rust, J., Frankenhäuser, H. and Wilde, V., 2012. Testing for the Effects and

Consequences of Mid Paleogene Climate Change on Insect Herbivory. PLoS ONE, 7(7), e40744.

Zhang, J.-W. et al., 2015. Sequoia maguanensis, a new Miocene relative of the coast redwood, Sequoia

sempervirens, from China: Implications for paleogeography and paleoclimate. American Journal of Botany,

102(1), 103-118.


29

Project 3: Neogene climate change in Yunnan (China) inferred from molecular

biomarkers

Project 3 will address climate reconstruction in Yunnan (China) during the Neogene using a

set of molecular biomarkers. On the base of Project group 5 (dating & stratigraphy) and

complementing Project group 3, the proposed study will focus on reconstructing the two most

important climate variables, i.e. temperature and rainfall. Whereas macrofossils may be

restricted to distinct stratigraphic sections, biomarkers are present throughout a stratigraphic

sequence, allowing for a high-resolution (i.e. cm-scale sampling) reconstruction of ancient

climates. They are, however, less specific than macrofossils with respect to their organismal

source. Therefore, the proposed study will be conducted in close cooperation with Project 1.

Paleo-temperatures will be reconstructed using microbial membrane lipids, so called GDGTs

(Glycerol-Diphenyl-Glycerol-Tetraethers), produced by produced by anaerobic soil bacteria.

Comparison with CA and CLAMP based temperature estimates (Project 1) will lead to a

robust paleo-temperature reconstruction.

Information on past rainfall changes will be inferred form the analysis of the stable hydrogen

and carbon isotopic composition of sedimentary higher plant waxes (δDwax and δ13Cwax,

respectively). Together with the above mentioned temperature reconstruction and the result

of macrofossil-based environmental and floral reconstruction conducted in Project 1, we will

use δDwax to derive an estimate of total annual mean rainfall as well as fluctuations of

northwest vs southeast monsoonal flow. δ13Cwax, in turn, may contain valuable information on

changes in canopy closure/vegetation density throughout the study interval. The proposed

study aims furthermore at comparing leaf wax isotopes from individual fossil cuticles with the

integrated leaf wax signal in the enclosing sediment to capture “snapshots” of monsoonal

activity at the time of individual leaf formation vs. long-term changes in monsoonal flow.

In summary, our paleoclimatic data will be interpreted together with paleo-diversity

reconstructions of Project 1 and paleo-altimetry and –topographical changes inferred from

Project Group 3 to understand the interaction of topography and climate on creating the

biodiversity hotspot in Yunnan.

Project 4: “Dating/stratigraphy/provenance analysis” (Ulf Linnemann / Cheng-Long

Deng?)


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2.3. The hominoids

Ecology and Diversity of Primate Communities in the Neogene of Yunnan Province

Christine Hertler1, Tina Lüdecke2, Ji Xueping3 and Andreas Mulch2

1 Research Center “The role of culture in early expansions of humans (ROCEEH)”,

Senckenberg Research Institute, Frankfurt am Main, Germany.

2 Senckenberg Biodiversity and Climate Research Centre, Frankfurt am Main, Germany

3 Yunnan Cultural Relics and Archaeology Institute, Kunming, Yunnan, China.

The evolution of Neogene mammal communities in the Yunnan province is well documented

by a sequence of faunas from several Miocene to Plio-Pleistocene localities. These large

mammal communities display an unusual large taxonomic diversity, which is apparently

related to changes in the geological and climatological setting. The assemblages represent

major episodes of faunal change and therefore provide a rich database for the study of

faunal responses in relation to changes in their environments.

The Neogene records both, the uplift of the Tibetian plateau and the establishment of the

modern configuration of the East Asian Monsoon System, which played a significant role in

environmental change. Previous paleoecological studies however, found a remarkable

homogeneity in the dietary spectrum of herbivores indicating the persistence of tropical

rainforest. E.g., Biasatti et al. (2012) focused on herbivore communities and noted an

increase in the diversity of the dietary signals only after 3.5 Ma. Surprisingly, primates were

not included, although the primate communities in this region represent an important source

of information particularly for forest habitats.

South China plays an important role in our present understanding of primate diversity. Some

of the localities in the Yunnan province represent type localities for enigmatic taxa like the

hominoids Lufengpithecus and Gigantopithecus. Although the taxonomy of the hominoids in

the Neogene of South China has been studied by several authors (among others Harrison et

al. 2002 and Qi Guoqin et al. 2006) and the relationship of taxonomic diversity and

environmental change is undisputed (Jablonski 2005), the ecology of the primate

communities in this region has not yet been studied in greater detail.

We will study the ecological diversity and variability of the Yunnan province primates and

associated herbivore guilds to gain insights in differential faunal responses to environmental

change. The collection of ecological diversity indices and ecomorphological data will be

complemented by analyzing stable carbon and oxygen isotopes of their tooth enamel, which

record nutrition (including food and drinking water) and foraging strategies of the individual


31

mammals, which in turn reflects environmental parameters such as vegetation, water supply

or seasonality.

References:

Biasatti, D, Wang Yang, Gao Feng, Xu Yingfeng, Flynn, Lawrence, 2012. Paleoecologies and palaeoclimates of

late Cenozoic mammals from Southwest China: Evidence from stable carbon and oxygen isotopes, Journal of

Asian Earth Sciences 44, 48-61.

Harrison, T., Ji Xueping, Su, Denise, 2001. On the systematic status of the late Neogene hominoids from Yunnan

Province, China. Journal of Human Evolution 43, 207-227.

Jablonski, N.G., 2005. Primate homeland: forests and the evolution of primates during the Tertiary and

Quaternary in Asia. Anthropological Science 113, 117-122.

Qi Guoqin, Dong Wei, Zheng Liang, Zhao Lingxia, Gao Feng, Yue Leping, Zhang Yunxiang, 2006. Taxonomy,

age and environment status of the Yuanmou hominoids. Chinese Science Bulletin 51(6), 704-712.


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3. Evolution of Biodiversity – the genomic record

3.1. Plant groups

What are the specific plant diversity patterns of the SE Himalaya – Yunnan Hotspot?

Georg Miehe

The ‘Eastern Himalaya-Yunnan Centre’ (Barthlott et al.1996) is

(1) the only biodiversity hotspot outside the Tropical Realms, in the Holarctic Floristic Realm

of the northern hemisphere. In contrast to the other hotspots it has

(2) a uniquely high geo-diversity with (i) largest vertical extent, (ii) unique mountain ranges in

parallel with deep gorges (UNESCO World Heritage Site) and a (iii) unique archipelago of

sky mountains in the southeastern periphery of the world’s largest highlands.

The region host global maxima with respect to

(3) a highly diverse humidity pattern ranging between 296 mm/a (Dry River Gorges:

Benzilan) and probably exceeding 6000 mm/a (Hponyin Razi, NW Myanmar),

(4) the tremendous floristic contrasts of the highest and lowest biodiversity (Hotspot of the

southern slopes of the Southeastern Himalaya and the Coldspot (Dickore & Miehe 2001) of

the Tibetan highlands)

(5) the hosting of world’s probably richest alpine flora, and the occurrence of a snowbed in

direct vicinity to the tropics.

The project’s principal goals are

(1) to unveil general diversity structures of the hotspot (“where the hotspot is hot and where

is it cold?”/”dark diversity”), expecting that such patterns open insights to the evolution of

species diversity/richness,

(2) to unveil diversity structures of the supposed Tethys woodlands of the Dry River Gorges

as a set of the ‘Plant Museums’ (Lopez-Pujol et al. 2011), expecting that the species set and

its genetics contribute to the question of the relief evolution of the area,

(3) to record the alpine plant communities of sky mountains along their longitudinal chain

from 21°N towards the Southeastern Himalaya, expecting that latitudinal trends in species

numbers and species sets are related to evolutionary pathways (“Out of Tibet into the

Himalayan Periphery Exile” vs “Out of the Tropical/Holarctic contact zone into the emerging

mountain biota of the Himalayas/Tibet”; Plant Cradle of Alpine Flora: Lopez-Pujol et al.

2011),

(4) to record the cryptogamic and vascular plant species set of snowbeds and snow

avalanche areas.

Suggested work packages


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1.1 Review of current knowledge (This is pre-submission work to pre-test hypothesis):

Diversity Patterns according to the regional accounts (e.g. Flora Yunnanica/Flora of

China/Seed Plants of the Alpine Subnival Belt…/Plant List Hengduan Shan/Plant List

Gaoligong Shan/ Animals??). Question: What are the distribution patterns according

to plant lists? Which biota are rich/poor, what is the share of alpine biota?

1.2 Plot-based elevational transects: where is it possible to record natural sites all along

from the river to the summit (Yunnan??/Myanmar!!, Vietnam??, Bhutan!!).

Question: How is diversity distributed (where is it rich/poor/are there humps, who

makes the humps?)

1.3 Plant communities of the Dry River Gorges.

Question: Does the similarity of species set in the 3 river gorges give insights in the

ecological stability/age of dry Tethys woodlands?

1.4 Alpine plant communities of longitudinal (21° - 28°N) Sky Mountains in the SE

Himalayan Periphery (Chin Hills/Myanmar, Gaoligong Shan, Daxue Shan, Wuliang

Shan, Ailao Shan).

Question: Cradle/Species pump of the Eurasian Alpine Flora or Exile out of

Tibet/Hengduan Shan?

1.5 Plant communities of snowbeds and snow avalanches.

Question: Is the southernmost snowbed flora the cradle of the Eurasian snowbed and

avalanche flora? How old is the snowbed flora?


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Project 1: Role of topographically based habitat richness for the future and the past of

the biodiversity hotspot of Yunnan

Contributors (alphabetical): Wilfried Konrad, Georg Miehe, Christoph Neinhuis, Anita Roth-

Nebelsick, Christiane Ritz, Stefan Wanke

Introduction

Abiotic environmental parameters, such as soil and air humidity, type of soil, insolation, mean

temperature and its annual course, form a subset within all conditions which are limiting the

successful existence of a plant taxon. Accordingly, habitat shifts are expected to be a

consequence of climate change (and already partially demonstrated for current

anthropogenically caused global changes), as is – over longer time periods – evolution.

Speciation and species extinction were repeatedly linked to changing environmental

conditions. For instance, disappearance of biotopes are considered a major extinction cause.

Important for the formation of suitable habitats is, however, not only the macroclimate which

characterizes a large area, or the mesoclimate characterizing regional conditions. In fact, a

quite large number of microhabitats exist within a local climate, as is, for example, commonly

demonstrated for species with different insolation demands but living quite next to each other

on adjacent shady or fully insolated spots. A certain macro- or mesoclimate can thus

accommodate a number of different habitats with quite different conditions.

The number of different local habitats should increase with increasing topographic

complexity. For instance, in mountainous areas, slopes face different directions (north-facing

slopes, south facing slopes etc.), with very different daily climate. If such slopes include

further topographic structuring, such as rifts or smaller elevated areas, the number of

different microhabitats on the slopes will further increase, and expectedly with increasing

topographic complexity. When being subjected to climate change, such a mosaic of

microhabitats may serve as a “habitat buffer” that allows for local habitat shifts within its

boundaries, thus preventing extinction. Moreover, microhabitats may be fragmented and

isolated in the process, depending on topography and presence of geological processes,

such as erosion, facilitating speciation processes.

Complex topographies are expected to be particularly possible with mountainous

landscapes. The combination of topographically caused mosaic patterns of microhabitats

and additional climatic diversity superimposed by altitude gradients may thus make richly

structured mountainous areas particularly suitable for accumulating species (via habitat shifts

within the microclimatic buffers) as well as for speciation events. It is suggested to test this

idea for the Yunnan hotspot with a suit of cooperation subprojects. The results of this project

will be constantly discussed and updated with respect to the work of other groups.


35

Subproject Leaf traits and function along selected altitude gradients

Leaf traits have attracted considerable attention in ecophysiology as well as in palaeobotany,

based upon numerous correlations with environment and/or leaf functions. There are three

types of leaf traits: 1) architectural leaf traits, such as leaf shape and type of margin, 2)

(eco)physiological leaf traits, such as maximum assimilation rate Amax or leaf life span (LLS),

and 3) (bio)chemical traits such as nitrogen content or leaf dry mass. In ecophysiology, the

concept of the Leaf Economic Spectrum (LES) focuses on LLS, leaf mass per area (LMarea),

nitrogen content and Amax as traits that are substantially correlated with environment and with

each other (Wright et al., 2005). The underlying reason for these interrelationships is

explained as two different economic strategies, that of “slow” and “fast” resource acquisition

and processing (Reich, 2014). These functional traits are completed by venation

pattern/density and stomatal density and arrangement which are - as representing the leaf

supply and gas exchange system – connected to productivity and the LES (Blonder et al.,

2011; Fiorin et al., 2016; Roth-Nebelsick et al., 2001).

In palaeobotany, architectural leaf traits showing correlations with environment are

extensively used in palaeoclimate reconstruction (Yang et al., 2007). Possible functional

causes for many of these correlations are still not well understood (Royer et al., 2012), but

may be at least partially related to the LES. The adaptive character of various traits may,

however, be attenuated by phylogenetic signals, i.e. various lineages “stick” to taxon-specific

traits (Little et al., 2010; Nobis et al., 2012).

It is suggested to address the following questions, in cooperation with project partners:

Are there correlations between LES and other leaf traits and selected

altitude/microclimate gradients, with respect to lineages and communities?

How do leaf traits of lineages behave, i.e. do they show adaptive plasticity or reveal

phylogenetic signals?

What are the ecophysiological profiles of the lineage representatives?

How do the results relate to palaeobotanical results?

Based on:

Monitoring of environmental conditions along transects

Vegetation along the transects, correlations with microclimate and topography

Determination of microscopic and macroscopic leaf traits, according to the LES and

other concepts

Community-based as well as taxon-based

Correlations with microclimate and topography


36

Subproject Ploidy level variation across altitudinal gradients

Polyploidy is ubiquitous phenomenon in plants and is considered as rapid mechanism for

sympatric speciation (Adams and Wendel 2005). Since Stebbins (1950) and Löve and Löve

(1967) it has been generally assumed that polyploids are more frequent in stressful

environments at high latitudes and high altitudes. This view is supported by the observation

that formation of polyploids is most frequently triggered by the production of unreduced

gametes which is in turned enhanced by environmental stress (e.g. cold, nutrition, water

stress; (Mason and Pires, 2015; Ramsey and Schemske, 1998). However, the dogma about

distribution of polyploids has becomes recently more elusive because floristic approaches did

not account for differences in biogeographical and phylogenetic histories of lineages, and

due to the fact that tropical and subtropical regions remained understudied (Weiss-

Schneeweiss et al., 2013). In contrast to Brochmann et al. (2004) who found a high incidence

of polyploids in the artic flora (60.7%), European mountain ranges and the Hengduan

mountains in Yunnan contained an unexpected low proportion of polyploids in their flora

(about 20%; Nie et al. 2005; Loureiro et al. 2013), but these polyploids were indeed mostly

restricted to high altitudes (Loureiro et al. 2013). Such studies relied mainly on bibliographic

reviews and did not systematically analyse the distribution of cytotypes in relation to altitude

or other ecological factors. However, such knowledge is very useful because ploidy variation

may lead to ecological niche differentiation (Sonnleitner et al. 2010). Polyploidy might also

indirectly influence ecological diversification because lineages containing polyploids were

taxonomically more diverse than diploid lineages (Petite and Thompson 1999; Vamosi and

Dickinson 2006). Up to now case studies in species-rich genera from Hengduan mountains

revealed controversial results some implying the polyploidy played a major role in the

evolution of some genera (Chen et al. 2014; Meng et al. 2014, Luo et al. 2011); but not in

others (Deng et al. 2011, Liu et al. 2010; Yuan et al. 2008, Zhuo et al. 2008, Jin et al. 2007).

Possible research questions

Does polyploidy have an impact on plant diversification in Hengduan mountains? Are

there differences between lineages containing polyplpids than those containing

mainly diploids (e.g. Delphinium/Rosa: endemic Rosa praelucens highest naturally

occurring ploidy level in the genus (Jian et al., 2010)

Is there any correlation between the distribution of polyploids and altitude?

Does polyploidy effect functional traits (leaf characters etc.)?


37

Subproject Phylogenetic study of lineages occurring along selected altitude gradients

- Phylogenomic analyses of plant biodiversity

Potentially hundreds of loci are needed to resolve recalcitrant phylogenetic relationships

(Leaché and Rannala 2010, Wickett et al. 2014, Prum et al. 2015). Recalcitrant relationships

are very short internodes usually on shallow level, but also found on deep level. Those ‘‘short

branched clades’’ are found all over the tree of life and as such in many plant lineages. For

plants, it has early been recognized that these nodes are among the most challenging to

resolve (e.g. Richardson et al., 2004).

Whole genome sequencing would provide all available loci of an organism and researchers

would only need to select loci matching the requirements of the respective phylogenetic

question. Despite decreasing sequencing costs, whole genome sequencing is very

expensive if applied on a broad sampling and overstates the case as only a fraction of the

genome is potentially needed (Ruane et al. 2015). Furthermore, it requires a lot of

downstream (post sequencing) processing with still a lot of custom scripts and only a glimpse

of the obtained data is usable for phylogenetic questions (Carstens et al., 2012; Lemmon and

Lemmon, 2013). Consequently, if it is not possible to resort to multiple already sequenced

genomes, other methods have to be used to obtain a sufficient number of informative loci.

One recently developed technique is “Anchored Hybrid Gene Enrichment” (AHGE). AHGE

describes an approach where, prior to sequencing, selected loci are captured and enriched

with the help of “capture probes”, oligonucleotide sequences of ~60-120 bp that hybridize to

target regions (Lemmon and Lemmon 2013). Lemmon et al. (2012) designed a vertebrate

capture probe set and obtained a fully resolved and supported species tree for amniotes and

vertebrates. More recently, the same capture probe set was applied on Caenophidian snakes

(Pyron et al. 2014), African Agama lizards (Leaché et al. 2014 a), birds (Prum et al. 2015)

and Malagasy pseudoxyhophiine snakes (Ruane et al. 2015). However, this probe set only

works for vertebrates. For plants a capture probe set has been developed (Mast et al. 2013,

2014) and was successfully tested on the species used to design the probes (Buddenhagen

et al. in prep, Systematic Biology). Furthermore qur research group has applied the probe

set using closely related species/populations (Müller et al. in prep., Molecular Phylogenetics

and Evolution) and we developed a multiplexing strategy called Plant Anchored MetaPrep

(Grandos-Mendoza et al. in prep, Methods in Ecology and Evolution) to significantly reduce

the costs for densely sampled phylogenomic studies with multiple target taxa across the

angiosperms.

Possible taxa to be studied are (eight-twelve target lineages covering the angiosperm tree of

life)

Aristolochiaceae, Magnoliids


38

Piperaceae, Magnoliids

Hydrangeaceae, Asterids

Theaceae, Asterids

Cyperaceae (traits?) monocot

One additional monocot lineages (to be discussed) Bambus Gattungen?

Pedicularis, Lamiales

Rhododndron (Ericales)

Rosa, Rosids

Fabaceae, Rosids? (Christiane?)

Caryophylaceae? Primulaceae?

Subproject Evolution of Rosa

Out of 166 wild rose species native in China, 41 species are endemic to the Yunnan province

occurring at altitudes from 1800 to 3400 m (Cuizhi and Roberston, 2003). Some of these

species were analysed in genus-wide or section-wide phylogenies (Chen et al., 2014;

Fougère-Danezan et al., 2014) (Fougere-Danezan et al. 2015; Zhu et al. 2015) but a

biogeographical analyses of the roses from the region is missing to date. Comparisons

between fossil rose leaves from the late Miocene of the Yunnan Province (R. fortutita) with

extant species of the region revealed the presence of semicraspedodromous venation – a

rather rare character state in the genus – in both fossil and many extant taxa suggesting that

the general climatic requirements for the genus did not substantially change since the late

Miocene in the region. (Su et al., 2015). Moreover, the semicraspedromous venation was

also found in Rosa lignitum from Oligocene in Europe (Kellner et al., 2012) supporting the

hypothesis that the current climatic conditions in SW China match the European tertiary

climate.

In general, combined with genetic analyses leave traits are appropriate tools to investigate

ecological adaptations and/or evolutionary relationships in Rosa.

Possible research questions

Biogeographic origin of Rosa species in Yunnan

Do leaf characters correlate with ecological characters and/or with taxonomy

Preliminary work:

Kellner, A., Benner, M., Walther, H., Kunzmann, L., Wissemann, V., Ritz, C. M.

(2012) Leaf architecture of extant species of Rosa L. and the Paleogene species

Rosa lignitum Heer (Rosaceae). Int. J. Plant. Sci. 173 (3) 239-250.


39

DFG-LIS project: Digitisation / Cataloguing of non-textual objects: Development of

new digitization standards for the large-scale assessment of leaf venation traits from

herbarium collections using microradiographic imaging (Botanik der SGN: FR und

GLM)

References

Chen GF, Sun W-G, Hong, D-Y, Zhou Z, Niu Y, Nie ZL, Sun H, Zhang JW, Li ZM (2014) Systematic significance

of cytology in Cyananthus ( Campanulaceae) endemic to the Sino-Himalayan region. J. Syst. Evol. 52:

260-270.

Cuizhi, G.; Roberston, K. R. (2003) Rosa Linnaeus. In: Zhengyi, W.; Raven, P. H.; Deyuan, H. Flora of China.

Band 9 Pittosporaceae through Connaraceae. Science Press, Beejing, China; Missouri Botanic Garden

Press, St. Louis, USA.

Deng T, Meng Y, Sun H, Nie ZL (2011) Chromosome counts and karyotypes in Chaetoseris and Stenoseris

(Asteraceae-Cichorieae) from the Hengduan Mountains of SW China. . Syst. Evol. 49: 339-446.

Fougere-Danezan M, Joly S, Bruneau A, Gao XF, Zhang LB (2015) Phylogeny and biogeography of wild roses

with specific attention to polyploids. Ann. Bot. 115: 275-291

Blonder, B., Violle, C., Bentley, L. P., and Enquist, B. J., 2011, Venation networks and the origin of the leaf

economics spectrum: Ecology Letters, v. 14, no. 2, p. 91-100.

Chen, G.-F., Sun, W.-G., Hong, D.-Y., Zhou, Z., Niu, Y., Nie, Z.-L., Sun, H., Zhang, J.-W., and Li, Z.-M., 2014,

Systematic significance of cytology in Cyananthus (Campanulaceae) endemic to the Sino-Himalayan

region: Journal of Systematics and Evolution, v. 52, no. 3, p. 260-270.

Cuizhi, G., and Roberston, K. R., 2003, Rosa Linnaeus, Beijing, St. Louis, Science Press, Missouri Botanic

Garden Press, Flora of China - Pittosporaceae through Connaraceae.

Fiorin, L., Brodribb, T. J., and Anfodillo, T., 2016, Transport efficiency through uniformity: organization of veins

and stomata in angiosperm leaves: New Phytologist, v. 209, no. 1, p. 216-227.

Fougère-Danezan, M., Joly, S., Bruneau, A., Gao, X.-F., and Zhang, L.-B., 2014, Phylogeny and biogeography of

wild roses with specific attention to polyploids: Annals of Botany.

Jian, H., Zhang, H., Tang, K., Li, S., Wang, Q., Zhang, T., Qiu, X., and Yan, H., 2010, Decaploidy in Rosa

praelucens Byhouwer (Rosaceae) Endemic to Zhongdian Plateau, Yunnan, China: Caryologia, v. 63, no.

2, p. 162-167.

Kellner, A., Benner, M., Walther, H., Kunzmann, L., Wissemann, V., and Ritz, C. M., 2012, Leaf architecture of

extant species of Rosa L. and the Paleogene species Rosa lignitum Heer (Rosaceae): International

Journal of Plant Sciences, v. 173, no. 3, p. 239-250.

Little, S. A., Kembel, S. W., and Wilf, P., 2010, Paleotemperature Proxies from Leaf Fossils Reinterpreted in Light

of Evolutionary History: PLoS ONE, v. 5, no. 12, p. e15161.

Mason, A. S., and Pires, J. C., 2015, Unreduced gametes: meiotic mishap or evolutionary mechanism?: Trends in

Genetics, v. 31, no. 1, p. 5-10.

Nobis, M. P., Traiser, C., and Roth-Nebelsick, A., 2012, Latitudinal variation in morphological traits of the genus

Pinus and its relation to environmental and phylogenetic signals: Plant Ecology & Diversity, v. 5, no. 1, p.

1-11.

Ramsey, J., and Schemske, D. W., 1998, Pathways, mechanisms, and rates of polyploid formation in flowering

plants: Annual Review of Ecology and Systematics, v. 29, p. 467-501.

Reich, P. B., 2014, The world-wide ‘fast–slow’ plant economics spectrum: a traits manifesto: Journal of Ecology,

v. 102, no. 2, p. 275-301.


40

Roth-Nebelsick, A., Uhl, D., Mosbrugger, V., and Kerp, H., 2001, Evolution and Function of Leaf Venation

Architecture: A Review: Annals of Botany, v. 87, no. 5, p. 553-566.

Royer, D. L., Peppe, D. J., Wheeler, E. A., and Niinemets, Ü., 2012, Roles of climate and functional traits in

controlling toothed vs. untoothed leaf margins: American Journal of Botany, v. 99, no. 5, p. 915-922.

Su, T., Huang, Y.-J., Meng, J., Zhang, S.-T., Huang, J., and Zhou, Z.-K., 2015, A Miocene leaf fossil record of

Rosa (R. fortuita n. sp.) from its modern diversity center in SW China: Paleoworld.

Weiss-Schneeweiss, H., Emadzade, K., Jang, T. S., and Schneeweiss, G. M., 2013, Evolutionary Consequences,

Constraints and Potential of Polyploidy in Plants: Cytogenetic and Genome Research, v. 140, no. 2-4, p.

137-150.

Wright, I. J., Reich, P. B., Cornelissen, J. H. C., Falster, D. S., Garnier, E., Hikosaka, K., Lamont, B. B., Lee, W.,

Oleksyn, J., Osada, N., Poorter, H., Villar, R., Warton, D. I., and Westoby, M., 2005, Assessing the

generality of global leaf trait relationships: New Phytologist, v. 166, p. 485-496.

Yang, J., Wang, Y. F., Spicer, R. A., Mosbrugger, V., Li, C. S., and Sun, Q. G., 2007, Climatic reconstruction at

the Miocene Shanwang basin, China, Using leaf margin analysis, CLAMP, Coexistence approach, and

overlapping distribution analysis: American Journal of Botany, v. 94, no. 4, p. 599-608.

Subproject: Research activities in Vietnam National Museum of Nature an Hoang Lien

plant diversity hotspot in Northern Vietnam

Truong Van Do

Vietnam National Museum of Nature, [email protected]

Vietnam National Museum of Nature (VNMN) has been found in 2006, is a state-owned

cultural science museum, under Vietnam Academy of Science and Technology. Over past

ten years, the collection of specimens of VNMN has over 100,000 specimens, i.e. mammals,

reptiles and amphibians, fish, insects, plants, fossils, mineral, and including 300 type

specimens of these collections. The evolutionary history exhibition room has been also built

here for training and education to pupils and students. During past ten years, scientists of

VNMN have described approximately 65 species of reptiles and amphibians, insects, and

plants as new to science.

Additionally, we also present about Hoang Lien National Park as a plant diversity hotspot in

northern Vietnam. Hoang Lien NP locates in the Hoang Lien mountain range in northern

Vietnam where is linked to the Ailao Shan, Yunnan, China, in the southeast of the Himalaya.

Most of the area is over 1000 m above sea level, with a main peak is about 3143 m and is

considered to be the “Roof of Indochina”. Within its natural area of 29,845 ha, there are

21,894 ha of forest. Of this there are 14,678 ha of natural forest which only been slightly

impacted by human activities. Up to date botanists have investigated and listed about 2346

vascular plant species of 1,020 genera and 285 families occurring in the area. Of which,

many species are sharing the distribution with the Yunnan plant diversity hotspot.


41

Project 2: Evolutionary Ecology and phylogenomics of endemic plant taxa unravel

evolutionary cues of the HTO uplift

Jian-Quan Liu, School of Life Science, Lanzhou University, Lanzhou, PR China

Lars Opgenoorth, Philipp University of Marburg, Marburg, Germany

Project Description

Based on integrating phylogenomic, fossil and biogeographic analyses, we aim to clarify the

origin and evolutionary drivers of the Yunnan plant diversity. We will utilize groups with high

diversification (endemic species), varying ecological niches, and sufficiently small genomes

in order to be able to utilize genomic data to clarify identify genes or genomic regions with

high differentiation to derive their speciation patterns and drivers. Also we will use the data to

detect and partly date underlying changes of the effective population sizes by modeling and

testing demographic scenarios of speciation. Based on these results, we will recover the key

genes accounting for species divergences and adaptive differentiations as well as their

causal relationships. We will integrate all results to make clear the origin and evolution of the

Yunnan plant diversity.


42

3.2. Invertebrate groups

Project 1: Radiation into megadiversity: Ground beetles as phylogenetic model group

can unravel evolutionary principles of the Himalaya-Yunnan Hotspot

Joachim Schmidt, University Rostock, Rostock, Germany

Lars Opgenoorth, Philipp University of Marburg, Marburg, Germany

Project Description

We will utilize ground beetle phylogenies to unravel how orogenesis and the radiation in this

megadiverse taxa are linked. Ground beetles are an ideal biological indicator to derive

evolutionary processes, timing and consistence of the Himalaya-Yunnan Hotspot. Some of

the features of ground beetles that make them uniquely suited are, that they

1. provide a large spatial resolution (many species, populations, or genotypes) in the entire

research area.

2. have strong ties to a specific ecological niche, which will enable conclusions to be drawn

from the presence of that taxon regarding important abiotic factors of the environment

(e.g., temperature/elevation and humidity/precipitation).

3. populate all altitudinal belts of the high mountain range with characteristic and specialized

species assemblages.

4. are unspecialized predator or detritivore, which is independent of the existence and the

distribution of specific prey species.

5. are weak disperser in evolutionary timescales and thus are unable to fly or disperse over

large distances via vectors such as wind, large animals or humans, and thus exhibits

strongly differentiated lineages spatially.

6. have an availability of sufficient calibration points also extendable in the framework of the

project for the molecular-clock approach to date adequately the clades, taxa, and

genotypes.

7. provide a sufficient amount of taxa to fine-tune the molecular-clock approach and include

a coherent timeframe with coherent error bars.

8. have an availability of a comprehensive sample collection that is easily extendable by

additional fieldwork and for which a good systematic revision exists.

In mountains at lower latitudes as in the Himalaya-Yunnan Hotspot, almost all of the ground

beetle lineages adapt to the high mountain habitat by the reduction of their hind wings and

flight muscles as well as an irreversible transformation of their exoskeleton (distinct

shortening of the metathoracal plates). This evolutionary stage is termed primary

winglessness (winglessness by descent, Schmidt et al. 2011) to differentiate it from the

facultative winglessness stage (species with a normal metathorax and presumably with the


43

reversible development of hind wings in at least some individuals). Facultative winglessness

in ground beetles is a common phenomenon in mountains of mid and higher latitudes. In

contrast, our long-term study in the Himalaya and in Tibet demonstrated that the vast

majority of ground beetle lineages adapted to the cloud forests and to the alpine zone of the

HTO belong to the primary wingless species (e.g., Andrewesius: ca. 45 species; Ethira clade:

76 species; Lepcha: ca. 50 species; Meganebrius: 25 species; Xestagonum: > 250 species).

In these species groups, winglessness appears to have evolved during an early stage in their

evolutionary history. Winglessness induces an extreme reduction in ground beetle vagility.

The dispersal of primary wingless species can only occur on foot. Therefore, because of the

multitude of potential barriers in high mountains (mountain crests, glaciers, deep river

gorges, dry slopes), dispersal is only possible within a very restricted geographical range.

Thus, the current range position of a lineage of wingless ground beetles is closely related to

the geomorphological and climatological development of the mountain range they inhabit.

The fossil database of ground beetles that will be used for calibrating the molecular clock is

of high-quality although it has only been partly revised. Ober & Heider (2010) published an

overview of the most useful Carabidae fossil evidence in the literature. Additional fossils of

the genus Carabus from the Oligocene and Miocene of Europe and North America are

known (reviewed by Deuve, 2004), and recently, two modern ground beetle genera

(Bembidion and Calathus) in Baltic Amber were described by Ortuño & Arillo (2009, 2010)

and of our working group (Schmidt 2015, 2016). However, if one considers the catalogues of

insect inclusions in a number of Baltic Amber collections (e.g., Klebs 1910, Larsson 1978,

Hieke & Pietrzeniuk 1984, Bachofen-Echt 1996, Weitschat & Wichard 1998), the fossil data

pool of ground beetles is much higher than one would expect from the number of existing

descriptions of fossil taxa. Indeed, many fossil examples of many modern ground beetle

tribes or genera are preserved in Baltic Amber collections (e.g., Agonum, Lebia,

Pterostichus, Syntomus, and Trechus among others). However, to date, these fossils have

not been morphologically analyzed or systematically revised, and the true systematic

positions of most of the Baltic Amber fossils remain questionable. Morphologic-systematical

studies of the fossils are of major importance in paleo-carabidology and are a key element of

this application.


44

Project 2: Freshwater Invertebrates as models for studying the origin and the causes

of species diversity in Yunnan – Project outline

Martin Brändle, Roland Brandl

Philipps-Universität Marburg, Faculty of Biology, Department of Ecology – Animal Ecology

Due to their insular character and clear physical boundaries freshwaters are good model

systems to study species communities, species propensity for dispersal but also the

evolution of species diversity. Freshwaters in general provide habitats for a large number of

invertebrate species comprising several taxonomic groups. Species within and among

taxonomic groups differ in life history traits, diet utilization and environmental requirements.

Many freshwater invertebrates are restricted to use either standing (lentic) or running (lotic)

waterbodies. Strong differences of these habitat types in predictability and environmental

conditions provide versatile opportunities to study genetic population differentiation, species

propensity for dispersal and the evolution of species diversity.

The Yunnan landscape is covered by a large number and variety of freshwater habitats.

Specific habitat characteristics of these environments and their spatial distributions may

serve as important template for the origin of the Yunnan hotspot of species diversity. In our

project we will focus on two overarching core question that are central for the main project: (i)

When did the Yunnan hotspot first appear in Earth History, and how did diversity

evolve? (ii) What are the main drivers in the evolution of diversity?

(i) To identify temporal patterns of diversification and the timing of radiations we will

consider time calibrated molecular phylogenies and lineage through time plots of

selected lentic and lotic freshwater invertebrates. For both habitat types we will

also analyze taxonomic groups that differ a priori in propensity for dispersal, for

instance wingless versus winged taxonomic groups (e.g. flatworms, mollusks,

crustaceans versus insect orders). The spatial patterns of diversification will be

analyzed by contrasting taxon age and phylogenetic relatedness with geographic

settings and distribution and subsequently compared between lineages differing in

propensity for dispersal. Here we are in particular interested how mountain uplift

influenced the diversification and distribution of species differing in phylogenetic

age. For example, we expect for poor dispersers that phylogenetic older groups

are more restricted to the bottom of the mountains while younger groups should

be found rather at higher elevation. In addition, we expect that populations of lotic

and wingless invertebrate species (flatworms, mollusks and crustaceans) are

more strongly spatially structured than lentic species and species of insect orders.


45

(ii) To identify the main drivers of diversification in Yunnan we will compare patterns

of species richness and composition of the Yunnan freshwater invertebrate fauna

with those of another mountain region, namely Central Europe and Southern

Europe. By relating species richness among higher taxonomic groups against

each other it will be possible to identify groups that show a disproportionally

higher richness in Yunnan. After having identified these groups we will determine

what are the dominant life history traits or niche requirements of the species

within these groups? For instance, the case that taxonomic groups comprising

more lotic species show a much higher species number in Yunnan than in the

Alps would indicate that the structuring of lotic freshwaters may have served as

an important driver of diversification. In a similar vein, a finding that higher

richness of typically alpine or subtropical clades in Yunnan would provide hints

that the distinct relief and the heterogeneity of the landscape have to be

considered as an important factor driving the diversification of the Yunnan

hotspot.


46

Project 3: Biodiversity of Cenozoic insects from Yunnan (China)

Sonja Wedmann, Senckenberg Research Institute Frankfurt/M., Germany

Insects are the most diverse organisms in the history of life, with more than one million

described species, and millions of species still unknown. Insects also have a rich fossil

record, but although many studies have been done and are still ongoing, the course of

evolution of the biodiversity of insects is still poorly known in large parts, even for the

Cenozoic.

During the early Cretaceous there was a major turnover in the insect fauna, probably related

to the radiation of flowering plants (Ross et al. 2000). By the late Cretaceous, about 95% of

insect families were the same as they are today, with no significant decrease across the K/T

boundary (Grimaldi & Engel 2005). For the Paleocene, fossil sites with insect remains are

scarce, but the general trend for increasing insect biodiversity during the Cenozoic is well

documented by rich insect localities from the Eocene, e.g. Baltic Amber and Messel from

Europe. The high insect biodiversity in the Paleogene was attributed to high origination rates

and low extinction rates along with a nearly worldwide tropical to temperate climate (e.g.

Grimaldi & Engel 2005). A drastic climatic cooling during the Eocene-Oligocene transition

resulted in a “lesser” mass extinction that also affected the insects. Since modern types of

tropical forests and grasslands began to establish themselves in the Oligocene, this might

also be the beginning of our modern insect fauna (Grimaldi & Engel 2005). The implications

of the climatic shifts during the Neogene are poorly studied, the Miocene is a very interesting

time span in this respect, and there is still much to do.

Similar to the evolution of insect biodiversity, the interrelationships between climate change

and biogeographic diversity patterns are still under discussion. Fossils have been shown to

be important to redefine biogeographic hypotheses for many different insect groups, which

were believed to have restricted distributions when only extant taxa were considered; see

Eskov (2002) and Grimaldi & Engel (2005).

Considering this broad context, it seems very promising to study fossil insects from the

Yunnan province in China. The knowledge on fossil insects from this region is very scarce. If

collections of fossil insects from Yunnan can be made and are available for study, their

analysis would be the base for a comparative study of the evolution of insect biodiversity in

this region, and also the base for further biogeographical and comparative studies.

From the Shangyong village, Mengla County, Yunnan, a few unique insect compression

fossils have been described (Lin et al. 2010a, b), but it is recorded that fossil insects can be

found frequently (Lin et al. 2010 a). The age of these insects is considered to be upper


47

Paleocene or lower Eocene, but the dating of the sediments was based on Conchostracans

and it is not uncontroversial. But the time is certainly very interesting, and comparison with

Messel and other fossil sites would be promising.

From Maguan County, Wenschan City, Yunnan, besides plants (Zhang et al. 2015) several

insects were found, which will be studied soon (personal comm. T. Wappler October, 2015).

These insects probably have a late Miocene age (Zhang et al. 2015), and so they represent

a sample of a fauna which is much more recent. Especially analysis of a larger collection of

these insects, which probably would have to be assembled in the course of the project,

would permit interesting comparisons over time. There are also other Miocene sites from

which plants are known, and it would be interesting to investigate whether there can perhaps

also insects be found (Ma et al. 2005, Su et al. 2013).

Eskov, K. Y. 2002. Geographical history of insects. 427-435. In: Rasnitsyn, A. & Quicke, D. L. J. (Eds.): History of

insects. Kluwer Academic Publishers, Dordrecht, Boston, London, 517 pp.

Grimaldi, D & Engel, M. S. 2005. Evolution of the insects. Cambridge University press, Cambridge, New York, etc.

755 pp

Lin Qibin, Szwedo, J., Huang Di-Ying, Stroinski, A. 2010a. Weiwoboidae fam. nov. of 'Higher' Fulgoroidea

(Hemiptera: Fulgoromorpha) from the Eocene Deposits of Yunnan, China. Acta Geologica Sinica (English Edition)

84: 751-755.

Lin Qi-Bin, Petrulevicius, J.F., Huang Di-Ying, Nel, A., and Engel, M.S., 2010. First fossil Calopterygoidea from

Southeastern Asia (Odonata: Zygoptera): A new genus and species from the Paleogene of China. Geobios 43:

349-353.

Ma , Q. W., Li , F. L. & Li, C. S. 2005: The coast redwoods ( Sequoia ,Taxodiaceae) from the Eocene Heilongjiang

and the Miocene of Yunnan, China. Review of Palaeobotany and Palynology 135 : 117 – 129.

Ross, A. J., Jarzembowski, E. A. & Brooks, S. J. 2000. The Cretaceous and Cenozoic record of insects

(Hexapoda) with regard to global change. 288-302. In: Culver, S. J. & Rawson, P. F. (Eds.): Biotic response to

global change - The last 145 million years. Cambridge University Press, 501 pp.

Su , T. , F. B. Jacques , H. J. Ma , & Z. K. Zhou. 2013. Fossil fruits of Ailanthus confucii from the upper Miocene

of Wenshan, Yunnan Province, SW China. Palaeoworld 22 : 153 – 158 .

Zhang, J. W., D’Rozario, A., Adams, J. M. , Li, Y. ,Xiao-Qing Liang, X-Q., Jacques, F. M., Su, T. & Zhou, Z.-K.

2015. Sequoia maguanensis , a new Miocene relative of the coast redwood, Sequoia sempervirens , from China:

implications for paleogeography and Paleoclimate. American Journal of Botany 102: 103 – 118.


48

3.3. Vertebrate groups

Project 1: Evolutionary history and present phylogeographic patterns of passerine

birds of the Yunnan biodiversity hotspot

Martin Päckert1, Jochen Martens2, Yue-Hua Sun3, Patrick Strutzenberger1, Swen Renner4

1Senckenberg Naturhistorische Sammlungen, Dresden, Germany

2Institut für Zoologie, Johannes-Gutenberg-Universität, Mainz, Germany

3Key Laboratory of Animal Ecology and Conservation, Institute of Zoology, CAS, Beijing, China

4Institute of Zoology, University of Natural Resources and Life Sciences, Vienna, Austria

Background: The mountain forests along the southern and south-eastern margins of the

Qinghai-Tibetan Plateau (QTP) are one of the most prominent biodiversity hotspots of the

Northern Hemisphere. Local species richness is particularly high in the Eastern Himalayas

with no less than 358 species along a local elevational gradient. The Hengduanshan harbors

a similarly rich biodiversity including a high number of endemics. Phylogeny combined with

reconstructions of biogeographic history suggested a major burst of diversification in almost

all passerine families during the mid to late Miocene. In these times the Yunnan biodiversity

hotspot was a center of origin and diversification for several groups of passerine birds such

as rosefinches and leaf-warblers. Extant distributions and phylogeographic patterns along

the eastern QTP margin were predominantly triggered by Pleistocene range contractions and

Holocene range expansions. Cryptic diversity is high in the Yunnan biodiversity hotspot and

species identification or recent descriptions of taxa new to science were mainly based on a

combination of morphological, genetic and bioacoustics traits – so-called integrative

taxonomy. Though genetic barcodes of birds are available in dense regional samplings

worldwide, the QTP region is still greatly underresearched. First results from a DFG-funded

barcode initiative on Eurasian birds confirm a considerable percentage of ‘undetected

species’ in the QTP region and stress the need of further research particularly in the

biodiversity hotspots.

Perspective: So far, our BOLD database comprises more than 600 COI barcodes of

Eurasian birds with dense samplings along the eastern margin of the QTP and in the Central

Himalayas (Nepal). The database is steadily being enlarged and can be used as the basis for

further biodiversity research in the region. Our sampling is complemented by rich material

from northern Myanmar (Hkakabo Razi) at the frontier to China (Yunnan) – from a

zoogeographic perspective this region is still part of the Yunnan Biodiversity Hotospot. Based

on our bacording initiative our future research will focus on phylogeographic studies that shall

help uncovering intraspecific differentiation and cryptic biodiversity. The genetic studies will

be flanked by bioaoustic analysis of territorial songs.


49

4. Present-day diversity distribution in Yunnan – a macroecology and modelling

analysis of present-day patterns

Understanding changing patterns of diversity in Yunnan and beyond

Alice C. Hughes1 and Matthew Forrest2

1Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences and 2Biodiversity

and Senckenberg Biodiversity and Climate Research Centre (BiK-F)

Abstract

Southeast Asia represents a complex and dynamic region both in terms of ecology and

geography. It currently maintains rich biodiversity [[references]] against a historical backdrop

of many geophysical processes and events (and the emergence of the Sunda shelf during

considerable periods of history has a lasting influence on contemporary patterns of

biodiversity) which have a had a significant effect on the landscape [[references,

references]]As a result, understanding modern patterns of diversity, and disentangling

genuine habitat requirements and physiological drivers of modern distributions from the

historical biogeographic drivers of biodiversity is challenging. Paleontological data from

across much of the region is patchy in terms of spatial and temporal resolution, and as a

result there is still dispute as to the landcover and connectivity across much of the region

during crucial portions of the region’s history, such as the last Glacial maximum. Employing

process-based models such as General Circulation Models (GCMs) and Dynamics Global

Vegetation Models (DGVMs) to study previous geological periods in combination with a

synthesis of existing paleontological data offers a means to build more complete picture of

past landscapes. Furthermore, using such models and data in tandem with present-day

biodiversity, species distribution and trait data may provide insights into both the habitat

changes in region and its effects on biodiversity. In a further extension to this methodology,

empirical models of biodiversity could be constructed using similar techniques to, or build

upon Species Distributions Models (SDMs) which are widely applied to predict present-day

distributions from a range of environmental data layers. By using consistent and mechanistic

simulations of climate and vegetation fields across different geological periods, and using

models of biodiversity calibrated using present day data and model output, we intend to

predict biodiversity patterns of the past and future, and examine the role of geophysical

processes in today’s biodiversity.

Research questions

We propose to develop a framework of interlinking research packages and novel

methodologies to tackle the following broad research questions:


50

Research question 1. How have key geophysical events and processes affected the

evolution of biodiversity in Southeast Asia and in particular the Yunnan biodiversity hotspot?

Research question 2. How has regional climate changed over time, and how has the

changing configuration of the region effected long term climatic trends

Research question 3. Which factors (including habitat change effects) best explain present

day biodiversity patterns in several keys groups (bird, bats, mammals, amphibians, insects,

trees) in Southeast Asia?

Background

Modern distributions of any given species will necessarily reflect former distributions, and

especially for dispersal limited species the legacy of former distributions is of particular

importance in understanding current patterns of diversity. Species distribution models for the

present and past and calibrated with appropriate data, have provided insights into species

distributions and useful projects of how species distributions might change under global

change scenarios.Such models can also be applied to previous time periods to estimate

species distribution, and how these may have changed over time, the choice of input data is

critical in obtaining meaningful and useful outputs. For example climatic models of this highly

diverse and heterogenous region are normally based on the interpolation of very coarse raw

climate data over the topography of the area using various splines, and without calibration

and testing for many regions (Deblauwe et al., 2016). As a result, the differences between

the climate in any two regions may actually be exceeded by the variability and uncertainty for

any given climate parameter in those regions, and such variation and error is likely to be

further compounded by extrapolation over time. Other issues, such as the changing

topography of the region (i.e the emergence of the Sunda shelf for half of the regions

geophysical history since taking it’s current configuration) will also have effected air-currents,

and thus profoundly affected the climate and even monsoon regimes across the region.

Thus, for accurate models of the distribution of species of any region, and especially one with

such heterogeneity having accurate climatic maps is essential.

Another important consideration when studying species distributions and biodiversity

patterns is the biotic, as opposed to the abiotic conditions of the landscape: specifically the

vegetation cover. This can be described by discrete land cover categorisations, such as

biome type, or as continuous variables such as woody cover fraction, net primary

productivity, phenological indices, and many others. For the present day, many of these can

be estimated using remote sensing data, however for the past and the future no such

measurements are available. Thus if spatially extensive maps describing the key features of

vegetation cover are to be used for calculating species ranges and biodiversity outside of the

present day, they must be reconstructed using models. This is of particular importance for

Southeast Asia, as for much of the region an understanding of the distribution of different


51

landcover types over important periods of history (i.e the Last Glacial Maximum) are crucial

to understanding patterns of refugia, population connectivity and thus drivers behind current

distribution.

Vegetation can of course be mapped and modeled in a variety of ways, and using

appropriate data (i.e. pollen, speleotherms, etc) it is possible to test, calibrate and verify the

accuracy of such vegetation models across time. LPJ-GUESS is an eco-system model which

also provides an alternate means of mapping regional landcover ( Forrest et al 2015).

Once vegetation models have been developed and tested for various time periods, the

combination of climatic, vegetative and topographic data can be used to run biodiversity

models (using predictive techniques) for a suite of taxa for each time period under scrutiny.

These changing distributions can then be explored using molecular approaches (where data

is available, and for select taxa only) to explore genetic connectivity, and phylogenetic

endemism, and explore the changing connectivity of these groups over time and the

accuracy of glacial refugia projected.

The intersection of various forms of data to understand past and present distributions of

vertebrate and plant species across the region represents a challenge, which without the

modern availability of data, software and higher capacity to run computationally intensive

models would have been impossible. Species distribution data for the present, and to some

extent for the past has already been obtained, though the databases are still under the

process of development (so are growing) under a separate initiative also lead by the leaders

of this proposal. Broadly we intend to build high resolution environmental and climatic data

for a number of time periods using various forms of available data, to build onto this

vegetation models for the appropriate periods, and then to hindcast modern diversity onto

these projected landscapes using species distribution modeling techniques, and calibrating

the outputs through genetic analysis (for changes since the LGM) and through fossil data for

older analysis. Thus the outputs promise new insights into patterns of diversity and their

drivers across geophysical time, and provide a basis to better conserve current species and

genetic diversity. Additionally outputs should be capable of detecting, to at least some

degree, the sensitivity of different species to change and their probable reactions to future

climatic changes.

Furthermore by examining the effects of climate stability and variability across such a

heterogeneous landscape across time, and relating it to modern diversity and endemism,

calibrated with molecular data will provide new perspectives on how species respond to such

changes over time. It is probable that areas with moderate levels of stability have the highest

diversity, whereas highly unstable regions may have a combination of highly endemic

dispersal poor taxa (which have survived through change) and highly generalist good

dispersers which have since colonized those regions. Modern patterns of distribution,


52

combined with population genetics will elucidate the changing distributions of each,

combined with the rate of movement and therefore carry useful implications for species

future responses to changing climate. The distribution of refugia across different areas of the

landscape, and the relative “velocity of climate change” related to the areas topography may

also carry important lessons for how species may respond to future change

Work Packages

WP 1. Paleoclimate modelling. Personnel ???

This depends on Johan Liakka's involvement. Feeds into WP 2 and 3. Alternate plan 1, use

data from/bring in Todd Ehlers? He and Jingmin Li are doing Middle Miocene stuff with the

explicit intention of looking of looking at East Asian monsoon. It might be easy(ish) for them

to do simulations other time periods. Alternate plan 2, Hui Tang’s simulations, and

collaborations, he seems to being similar things to Todd. But I think Johan is preferred ;-)

WP 2. Vegetation modeling. Personnel: One student in Frankfurt?

First we develop a regional parameterization for the model, also including paleo taxa. From

this we can offer information to about biomes, productivity, and other environmental data to

the other projects and, importantly, to WP3. Optionally, we can also consider starting to put

mechanism to allow trait variation within one PFT. Maybe, if this is done right, it also can be

used to explain biodiversity? Maybe then this is a hypothesis:

WP 3. Biodiversity modeling: Past, Present and future. Personnel ?? A student? In China

or Germany?

I think that this should start with mining the trait databases and biodiversity data (whilst the

other WP are getting the modelling going). The trait data can be used for a) A data driven

approach for testing hypotheses and b). Validating the trait-based improvements to the veg

model. Then, once the modelling is ready, they can use the model results and the data to

build the biodiversity model and answer some hypotheses. It is better for more informed

people to work these out, but initially I see possibilities like:

Hypotheses concerning drivers of present day biodiversity patterns


53

5. Future threats

Global Warming and changing monsoon? (Modelling/A. Hughes + M. Forrest)

Conservation strategies? (D. Cicuzza)

Fire? (Li Shu-Feng)

Land use / medicinal plant harvesting / mushroom economy (KIB, Yunnan University)

Abstracts/project outlines still required


54

Field trips

Day 1: Vereinigtes Schleenhain opencast mine, NW Saxony, Germany.

Day 2: Welzow-Süd opencast mine, S Brandenburg, Germany.


55

Field trip destinations opencast mine Vereinigtes Schleenhain and Welzow-Süd (map:

Susann Stiller).

S a

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0km


56

Practical palaeontological field work part 1

Eocene to Oligocene vegetation and climate change in the central German

Weißelster Basin (Vereinigtes Schleenhain opencast mine, Saxony)

Lutz Kunzmann & Karolin Moraweck

(text compiled from Kunzmann 2012, Kunzmann and Walther 2012, Kunzmann et al. 2016)

The Vereinigtes Schleenhain opencast mine is situated in the center of the Weißelster Basin,

which is a lignite lagerstätte south of the city of Leipzig in Saxony and Saxony-Anhalt, central

Germany (Meyer, 1950; Fig. 1). The name Weißelster is also applied to the important

palaeontological site in the strata (Walther and Kunzmann, 2008). However, in geological

and structural terms, the Weißelster Basin is not a distinct or isolated sedimentary

depositional area, but rather the southern part of the Leipzig Embayment, which pertains to

the North German Depression (Fig. 2; Standke, 2008a, 2008b). This name has therefore

gradually disappeared from the geological literature, whereas in the palaeobotanical

literature, it remains in use.

The Leipzig Embayment contains middle Eocene to middle Miocene terrestrial and marine

sedimentary sequences (Standke et al., 2010). Its southern end, the so-called Weißelster

Basin, is characterized by a series of fluvial, estuarine, and marine strata, including the

occurrence of several currently economically important lignite seams (Standke et al., 2010).

Lignite has been excavated here for several centuries and is nowadays mined by the

MIBRAG mbH in the Vereinigtes Schleenhain and in the Profen mines (Fig. 3).

The general lithostratigraphic section of the Weißelster Basin (Fig. 4) shows five main

sedimentation units, (1) the late middle to late Eocene Profen Formation with lignite seam

complex 1, (2) the late Eocene Borna Formation with lignite seam complex 2/3, (3) the latest

Eocene to earliest Oligocene Böhlen Formation with lignite seam complex 4, (4) the early

Oligocene “Rupelian” marine sediments, and (5) the late Oligocene Cottbus Formation

without lignite seams. Units (1) – (3) and (5) are predominated by clastic, non-consolidated

terrestrial sediments but (4) is brackish to fully marine. The field trip will guide to fossiliferous

horizons within the units (2) and (3). These sediments represent deposits of a shallow marine

tidal embayment and a coastal alluvial plain (Kunzmann 2012). A series of cyclic

transgressions and regressions caused a formation of large coastal mires accumulating peat

in the flat alluvial plain (Kunzmann et al. in press).


57

This area is well-known for its rich and diverse plant assemblages that cover the rather long

time span from the late middle Eocene to the earliest Miocene (Walther & Kunzmann 2008).

Here one can find a series of taphocoenoses of plant fossils, intensively studied, that touch

on the Mid-Eocene Climatic Optimum, the Eocene-Oligocene turnover as well as the late

Oligocene cooling (Zachos et al. 2008). Consecutively, these plant assemblages were used

to establish phytosociological and phytostratigraphic concepts, namely the “Florenkomplexe”

(floristic complex; Mai & Walther 1983, Mai 1995). By a definition, a floristic complex is a

temporary stage in the floristic evolution of regional vegetation independently from the

sedimentary facies type, the soil moisture and the type of plant communities (Mai and

Walther 1983, Mai 1995). Therefore, a floristic complex represents a kind of stasis in the

evolution of regional vegetation over a certain time span. Simultaneously, rich palynofloras

from the same horizons were the basis for another phytostratigraphic tool, the spore-pollen

zonation (e.g. Krutzsch 2011 and references therein). More recently, fossil floras from the

Weißelster Basin have been frequently used as an excellent database to estimate

palaeoclimatic and palaeoatmospheric changes (e. g. Mosbrugger et al. 2005; Grein et al.

2013; Roth-Nebelsick et al. 2014, Moraweck et al. 2015, Steinthorsdottir et al. 2015).

The fossiliferous horizon from unit (2) which is visited during the field trip contains floras from

the late Eocene floristic complex Zeitz (Mai and Walther 2000), and the fossiliferous horizon

from unit (3) contains a flora from the earliest Oligocene floristic complex Haselbach

(Kunzmann 2012). Between these two complexes an important floristic turnover from

predominantly notophyllous evergreen broad-leaved vegetation to mixed evergreen

deciduous vegetation could be observed (Kvaček & Walther 2001, Kvaček et al. 2014). This

change coincides with important climatic changes, i.e. temperature drop from “subtropical” to

warm-temperate (Kvaček et al. 2014), and decreasing palaeo-pCO2 values in the

atmosphere (Steinthorsdottir et al. 2016). In short, here a regional vegetation change is

recognized as a response to global change at the Eocene-Oligocene turnover (Fig. 5).

Late Eocene megafloras. Fossil floras from the late Eocene Borna Formation [unit (2)]

belong to the Zeitz floristic complex (Fig. 6; Mai & Walther 1985, 2000; Hennig & Kunzmann

2013; Ferdani 2014). This complex covers a considerable period of ca. 3 Ma from ~ 38.5 –

35 Myr (Text-fig. 1; Moraweck et al. 2015). The respective time slice is characterized by a

global cooling trend detected from the marine isotope record (Zachos et al. 2008) from nearly

‘subtropical’ to warm-temperate conditions. At the moment this trend could not be traced

adequately by various macrofloras in this area. Moraweck et al. (2015) concluded that floras

of the Zeitz complex are most likely unsuitable to detect gradual climatic changes based on

the limited species diversity in the individual taphocoenoses and high fragmentation rates of

the fossil plant material, which lowers the significance of the derived results by quantitative


58

palaeoclimate reconstruction methods. However, recent phytosociological investigations on

late Eocene macrofloras from Northern Bohemia and central Germany (Kvaček 2010;

Kvaček et al. 2014; Teodoridis and Kvaček 2015; Kunzmann et al. in press) provide hints

about gradual restructuring of late Eocene vegetation in central Europe that is most likely

caused by climate change.

Plant assemblages from about 35 floras of the Zeitz complex represent primarily riparian

vegetation of mainly azonal environments and to minor extent swamp forests of floodplains

and pioneer vegetation in highly disturbed habitats such as river banks, and mesophytic

forests. Kvaček (2010) named the vegetation type notophyllous evergreen broad-leaved

(riparian) forest with palms.

In more detail, Mai and Walther (2000) reconstructed for the Zeitz floristic complex several

azonal vegetation units, i.e. the Steinhauera-Rhodomyrtophyllum riparian forest, and the

Lauraceae-conifer swamp forests. These plant assemblages are usually predominated by

taxa such as Steinhauera subglobosa, Rhodomyrtophyllum reticulosum and Lauraceae (e.g.

Daphnogene cinnamomifolia, Laurophyllum syncarpifolium), which are associated with

evergreen Eotrigonobalanus furcinervis (Fagaceae), several Theaceae, palms and conifers.

Conifers are represented by Quasisequoia couttsiae (Cupressaceae) - a typical back-swamp

conifer element, and by Doliostrobus taxiformis (Doliostrobaceae), which is more

characteristic for riparian associations. Mai and Walther (2000) also concluded on the

presence of zonal Eotrigonobalanus-oak-Lauraceae forests with conifers, which is a matter

of interpretation.

Several plant taphocoenoses derived from the interbedding horizon between two measures

of the main lignite seam complex (Mai & Walther 2000). Some of these taphocoenoses have

been described recently (Hennig & Kunzmann 2013; Ferdani 2014). The siliciclastic

interbedding horizon consists of alluvial plain sediments (lacustrine, fluvial) and tidal

sediments (Rascher et al. 2008; pers. comm. Gerda Standke, Freiberg/Sa., Germany).

Fruits, seeds, leaves and wood are preserved as coalified remains (compressions, Ferdani

2014) and come from silts and silty clays.

Early Oligocene megaflora. The Haselbach horizon of unit (3) is assigned to the lowermost

part of the Böhlen Formation, recently named the Gröbers Member (Standke et al., 2010). In

particular, the Gröbers Member consists of brackish (tidal), fluvial, lacustrine, and palustrine

sediments (Standke et al., 2010) that in general represent a coastal plain environment during

a transgression setting. The Haselbach horizon is intercalated between a series of tidal

sands and silts and the Böhlen lignite seam complex. In its type area, the horizon consists

primarily of a 10–15-m-thick clay stratum (clay in the technological sense) that has been

mined commercially. However, the Haselbach horizon represents a series of fluvial,


59

lacustrine, and palustrine strata that were mainly deposited by anastomosing and

meandering rivers onto a heterogenous floodplain and by floodplain lakes. Abandoned

channel deposits, levee sediments, crevasse splay deposits, and floodplain mudstone are

recognized as common facies types (Kunzmann and Walther 2012). Recently, a split of the

Haselbach horizon into a lower and an upper stratum has been observed in Schleenhain.

The two units are separated from one another by a sand/silt layer. The age of the Haselbach

horizon is most likely earliest Oligocene based on pollen assemblages. Floristically, it is a

post Eocene-Oligocene turnover flora (Steinthorsdottir et al. 2016).

The megafossil plant assemblages were mostly excavated from abandoned channel deposits

at the base of the Haselbach horizon. These abandoned channel sediments are

characterized as generally planar-laminated, fine-grained sand and/or silt that alternate with

silty clay and phytoclasts (Figs. 7). Fruits, seeds, leaves and wood are preserved as coalified

remains (compressions, Kunzmann 2012). The fossil vegetation was characterized as mixed

deciduous-evergreen that were growing under warm-temperate and humid climate. In

particular, waterplant associations, riparian gallery forests, backswamp forests, and

mesophytic forests were reconstructed from the fossil assemblages (Kunzmann and Walther

2012).

In the last decades several taphonomical and phytosociological studies were used for

reconstructing the following fossil plant communities (see Kunzmann and Walther 2012).

(1) Water plant association: Lacustrine deposits of smaller floodplain lakes contain rarer

autochthonous carpo-assemblages such as a Salvinia-Azolla community including Azolla,

Salvinia, Lemna, Eichhornia, Hydrocharis, Ottelia, Stratiotes and Desembaya, and a

submersed-plant community including Aldrovanda, Ceratophyllum, Ludwigia, Potamogeton

and Vallisneria.

(2) Fern–monocot–shrubby angiosperm association from floodplain mud indicating alluviation

of still water: including the ferns Osmunda lignitum and Pronephrium stiriacum and

(deciduous) subshrubs such as Apocynophyllum neriifolium (Kunzmann and Walther 2007)

and Zingiberoideophyllum liblarense (Kunzmann 2012).

(3) Swamp forests (Fig. 8) are often dominated by a single conifer species such as

Quasisequoia couttsiae and Taxodium dubium. Myrica longifolia, several Lauraceae and

evergreen Fagaceae (Eotrigonobalanus furcinervis), palms, and Apocynophyllum neriifolium

also belong to this community that comes from organic-rich clays (Kunzmann and Walther

2012). A Taxodium-Nyssa swamp forest (Kunzmann et al. 2009) was a quite characteristic

association in the coastal lowlands consisting of Taxodium dubium, Nyssa altenburgensis,

Myrica longifolia and Sabal raphifolia.

(4) Riparian or gallery forest (Fig. 9): a softwood is comprised of Populus germanica,

Populus zaddachii, Alnus spp., Salix varians, Salix breunsdorfensis, Liquidambar europaea,


60

and Taxodium dubium, whereas Ulmus fischeri, Acer haselbachense, Carpinus grandis and

Platanus neptuni dominate the hardwood type. In more diversified taphocoenoses

Pyracantha kräuselii, Rosa lignitum, Lauraceae and the fern Lygodium kaulfussii come

along.

(5) Communities on sandy river banks have consisted of Pinus spp., Comptonia acutiloba,

Engelhardia orsbergensis, Liquidambar europaea, Pyracantha kräuselii, Pinus spp. and

palms.

(6) The zonal vegetation was represented by a Mixed Mesophytic Forest. This vegetation

unit is often dominated by evergreen species. Eotrigonobalanus furcinervis,

Trigonobalanopsis rhamnoides, Distylium fergusoni, Daphnogene cinnamomifolia,

Laurophyllum acutimontanum, L. pseudoprinceps and Illicium sp. are recognized in the leaf

component of the taphocoenoses. A divers range of deciduous plants such as Acer

haselbachense, Carpinus grandis, Engelhardia orsbergensis, Matudaea menzelii, Platanus

neptuni, and Ulmus fischeri were abundant but less frequent. Conifers including Pinus spp.

Tsuga plicata, Sequoia abietina and Cephalotaxus sp. occurred scattered. Shrubs and lianas

were recognized by Ampelopsis rotundata, Lygodium kaulfussii, Rosa lignitum,

Majanthemophyllum petiolatum.

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mitteldeutschen Raum. – Z. dt. Ges. Geowiss. 159/1, 81-103.

Standke, G., Escher, D., Fischer, J., Rascher, J. 2010. Das Tertiär Nordwestsachsens. Ein

geologischer Überblick. – 156 pp. (Sächsisches Landesamt für Umwelt, Landwirtschaft und

Geologie) Freiberg.


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Steinthorsdottir, M., Porter, A., Holohan, A., Kunzmann, L., Collinson, M. E., McElwain, J. C. (2015):

Fossil plant stomata indicate decreasing atmospheric CO2 prior to the Eocene–Oligocene

boundary. – Clim. Past 12, 439-454.

Teodoridis, V. and Kvaček, Z. 2015. Palaeoenvironmental evaluation of Cainozoic plant assemblages

from the Bohemian Massif (Czech Republic) and adjacent Germany. – Bull. Geosci. 90(3),

695-720.

Walther, H. and Kunzmann, L. 2008. Zur Geschichte der paläobotanischen Forschung im

Weißelsterbecken. – Z. dt. Ges. Geowiss. 159(1), 13-21.

Zachos, J.C., Dickens, G.R., Zeebe, R.E. 2008. An early Cenozoic perspective on greenhouse

warming and carbon-cycle dynamics. – Nature 451, 279–283.


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Figures

Fig. 1: Distribution of Paleogene sediments in central Germany – the Weißelster Basin (southern part

of the Leipzig embayment), arrow: Vereinigtes Schleenhain opencast min (from Standke 2008a).


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Fig. 2: Distribution of marine Oligocene sediments in northern central Europe, maximum extension of

the “Rupelian North Sea”, arrow: the Leipzig embayment (from Standke 2008a).


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Fig. 3: Central German mining district Weißelster Basin with two active opencast mines (Profen and

Vereinigtes Schleenhain) of the MIBRAG company (from www.mibrag.de).

http://www.mibrag.de/


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Fig. 4: General lithostratigraphic chart of the Paleogene and Neogene in central Germany, arrows

indicate the position of the fossiliferous horizons in the Vereinigtes Schleenhain opencast mine, i.e.

main interbedding horizon of lignite seam complex 2/3 and Haselbach Clay complex (from Standke

2008a).


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Fig. 5: Two leaf assemblages from the Weißelster Basin: the late Eocene assemblage Stedten of the

Zeitz floristic complex (left, from Mai 1995) and the early Oligocene assemblage Schleenhain of the

Haselbach floristic complex (right, from Kunzmann and Walther 2012).


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Fig. 6 (next page): Predominant, key and accessory taxa of the floras of the Zeitz floristic complex in

the Cenozoic of the Weißelster Basin (central Germany) and in North Bohemian basins (Czech

Republic) (from Kunzmann et al. in press).

6/1 Doliostrobus taxiformis (Sternb. 1833) Kvaček 1971, foliage shoot, Klausa, Germany, MMG

PB Kl 11.

6/2 Rhodomyrtophyllum reticulosum (Rossm. 1840) Knobloch & Kvaček in Knobloch, Kvaček &

Konzalová 1996, leaf with characteristic emarginated apex, Schleenhain 1, Germany, MMG PB

SchleOE 755 (orig. Hennig & Kunzmann 2013: pl. 1, fig. F; orig. Moraweck et al. 2015: fig. 3I).

6/3 Polyspora saxonica Walther & Kvaček in Kvaček & Walther 1984, leaf, Klausa, Germany,

MMG PB Kl 423a (orig. Mai & Walther 1985: pl. 24, fig. 6; orig. Moraweck et al. 2015: fig. 3D).

6/4 Daphnogene cinnamomifolia (Brongn. in Cuvier 1822) Unger 1850, leaf, Staré Sedlo

Formation, Czech Republic, MMG PB Ss 579.

6/5 Trigonobalanopsis rhamnoides (Rossm. 1840) Kvaček & Walther 1988, leaf, Staré Sedlo

Formation, Czech Republic, MMG PB Ss 35 (orig. Knobloch et al. 1996: pl. 23, fig. 4).

6/6 Steinhauera subglobosa C. Presl in Sternb. 1838, two infructescences (left: surface view,

right: in cross section), Schleenhain 1, Germany, MMG PB SchleOE 709:1a, b (orig. Hennig &

Kunzmann 2013: pl. 1, fig. E; orig. Moraweck et al. 2015: fig. 3J).

6/7 Platanus neptuni (Ettingsh. 1866) Bůžek, Holý & Kvaček 1967, compound leaf, Kučlín, Czech

Republic, MMG PB Kin 84 (orig. Kvaček & Manchester 2004: fig. 7b, orig. Kvaček & Teodoridis 2011:

pl. 5, fig. 10).

6/8 Eotrigonobalanus furcinervis (Rossm.1840) Walther & Kvaček in Kvaček & Walther 1989, leaf,

Schleenhain 1, Germany, MMG PB SchleOE 720e (orig. Hennig & Kunzmann 2013: pl. 1, fig. A; orig.

Moraweck et al. 2015: fig. 3A).

6/9 Sabal raphifolia (Sternb. 1822) Knobloch & Kvaček in Knobloch, Kvaček & Konzalová 1996,

leaf, Schleenhain 2, Germany, MMG PB SchleOE 1:1 (orig. Moraweck et al. 2015: fig. 3K).

6/10 Laurophyllum (“Neolitsea”) syncarpifolium (Friedrich 1883) Wilde 1989 f. gardneri Kvaček in

Knobloch, Kvaček & Konzalová 1996, leaf, Profen-Süd 1, Germany, MMG PB Pf 297g.

6/11 "Laurophyllum“ fischkandelii L. Kunzmann & Walther 2002, leaf, Profen-Süd 1, Germany,

MMG PB Pf 1379a.


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Fig. 7: Typical bedding plane with leaf compressions from an abundoned channel fill deposit from the

Haselbach Clay complex, Gröbers Member (from Kunzmann and Walther 2012).


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Fig. 8: Vegetation scheme of a typical early Oligocene Quasisequoia couttsiae swamp forest (modified

from Kunzmann and Walther 2012).

Fig. 9: Vegetation scheme and taxa of a typical early Oligocene riparian softwood and hardwood

riparian forest from the Weißelster Basin (unpublished conference contribution).


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Practical palaeontological field work part 2

Middle Miocene sealevel changes documented in plant taphocoenesis in the

2nd Lusatian lignite complex in East Germany

Franziska Ferdani & Lutz Kunzmann

During the field trip the Vattanfall Europe Mining opencast mine Welzow-Süd in the

Bundesland Brandenburg will be visited. The characteristic succession of plant

taphocoenoses within a middle Miocene lignite seam complex in relation to sealevel changes

and sedimentation processes in a coastal plane will be the focal point in the field. The

following pages contain useful diagrams and maps as an introduction to the geology,

stratigraphy and paleobotany of the East German mining area Lausitz (Lusatia).

Some important references:

Autorenkollektiv (2010): Die Geologische Entwicklung der Lausitz. – 193 pp. (Vattenfall

Europe Mining AG) Cottbus, Germany.

Dolezych, M. & Schneider, W. (2006): Inkohlte Hölzer und Cuticulae dispersae aus dem 2.

Miozänen Flözhorizont im Tagebau Welzow (Lausitz) – Taxonomie und

vergleichende feinstratigraphisch-fazielle Zuordnung. – Z. geol. Wiss., 34 (3-4): 165-

259; Stuttgart.

Dolezych, M. & Schneider, W. (2007): Taxonomie und Taphonomie von Koniferenhölzern

und Cuticulae dispersae im 2. Lausitzer Flözhorizont (Miozän) des Senftenberger

Göthel, M. (2004): Stratigraphie des Känozoikums in Brandenburg mit spezieller

Berücksichtigung des Braunkohlenreviers Lausitz. – Brandenburgische Geowiss.

Beitr., 11 (1-2): 149-168; Kleinmachnow.

Mai, D. H. (1999a): Die untermiozänen Floren aus der Spremberger Folge und dem 2.

Flözhorizont in der Lausitz. Teil I: Farnpflanzen, Koniferen und Monokotyledonen. –

Palaeontographica Abt. B, 250: 1-76; Stuttgart.

Mai, D. H. (1999b): Die untermiozänen Floren aus der Spremberger Folge und dem 2.

Flözhorizont in der Lausitz Teil II. - Palaeontographica Abt. B, 251: 1-70; Stuttgart.

Mai, D. H. (2000a): Die untermiozänen Floren aus der Spremberger Folge und dem 2.

Flözhorizont in der Lausitz. Teil IV: Fundstellen und Paläobiologie. –

Palaeontographica Abt. B, 254: 65-176; Stuttgart.

Mai, D. H. (2000b): Die mittelmiozänen und obermiozänen Floren aus der Meuroer und

Raunoer Folge in der Lausitz Teil I. – Palaeontographica Abt. B, 256: 1-68; Stuttgart.


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Mai, D. H. (2001a): Die mittelmiozänen und obermiozänen Floren aus der Meuroer und

Raunoer Folge in der Lausitz Teil II. – Palaeontographica Abt. B, 257: 35-174;

Stuttgart.

Mai, D. H. (2001b): Die mittelmiozänen und obermiozänen Floren aus der Meuroer und

Raunoer Folge in der Lausitz Teil III. – Palaeontographica Abt. B, 258: 1-85; Stuttgart.

Nowel, W., Bönisch, R., Schneider, W. & Schulze, H. (1995): Geologie des Lausitzer

Braunkohlenreviers, 2. Aufl. – 104 pp. (Lausitzer Braunkohlengesellschaft)

Senftenberg, Germany.

Roselt, G. & Schneider, W. (1969). Cuticulae dispersae, ihre Merkmale, Nomenklatur und

Klassifikation. – Paläont. Abh., B III (1): 1-128; Berlin.

Schneider, W. (1966): Beziehungen zwischen Pflanzeninhalt und petrographischer

Beschaffenheit von Weichbraunkohlen am Beispiel der miozänen Braunkohlen der

Oberlausitz. – Ber. deutsch. Ges. geol. Wiss., A Geol. Paläont., 11 (5): 615-633;

Berlin.

Schneider, W. (1992): Floral Successions in Miocene swamps and bogs of Central Europe. –

Z. geol. Wiss., 20 (5/6): 555-570; Berlin.

Schneider, W. (1995): Palaeohistological studies on Miocene brown coals of Central Europe.

– Int. J. Coal Geol., 28 (2-4): 229-248; Amsterdam.

Schneider, W. (2004): Eine blätterführende Taphocoenose im 2. Miozänen Flöz von Nochten

(Lausitz): Taxonomie, Taphonomie und Phytostratigraphie. – Palaeontographica Abt.

B, 268 (1-3): 1-74; Stuttgart.

Schneider, W. (2007): Magnolia L. in peat-forming associations of the Miocene seams in

Lower Lusatia (East Germany). – Acta Palaeobotanica, 47 (1): 217-235; Kraków.

Standke, G. (2006): Paläogeographisch-fazielle Modellierung des Unter-/Mittelmiozän-

Grenzbereiches in der Lausitz (Briesker Folge/Formation). – Schriftenreihe für

Geowissenschaften, 14: 1-130; Berlin.

Standke, G. mit einem Beitrag von P. Suhr (2008): Tertiär. – In: Pälchen, W. & Walter, H.

(Hrsg.): Geologie von Sachsen. Geologischer Bau und Entwicklungsgeschichte. –

358-419 (E. Schweizerbart’sche Verlagsbuchhandlung) Stuttgart.

Standke, G., Rascher, J. & Strauss, C. (1993): Relative sea-level fluctuations and brown coal

formation around the Early-Middle Miocene boundary in the Lusatian Brown Coal

District. – Geol. Rundschau, 82: 295-305; Stuttgart.


74

Fig. 10: General lithostratigraphic chart of the Paleogene and Neogene in central Germany, arrows

indicate the position of the fossiliferous horizons in the Vereinigtes Schleenhain opencast mine, i.e.

main interbedding horizon of lignite seam complex 2/3 and Haselbach Clay complex (from Standke

2008a).


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Fig. 11: East German mining area Lausitz (Lusatia) of the Vattenfall Europe Mining AG.


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Fig. 12: Distribution of the lower Miocene sediments of the Drebkau Member (Brieske Formation), blue

coloured = distribution of marine sediments, brown coloured = distribution of the complex of

underlaying sediments (from Autorenkollektiv 2010).


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Fig. 13: Figure above: Distribution of the middle Miocene sediments of the Welzow Member (Brieske

Formation) included the 2nd

Miocene lignite complex; Figure below: Distribution of regional

interburdens from the 2nd

Miocene lignite complex (from Autorenkollektiv 2010).


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Fig. 14: Distribution of the middle Miocene sediments of the Greifenhain Member (Meuro Formation),

blue coloured = distribution of mainly marine sediments, brown coloured = distribution of the complex

of overlaying sediment, yellow coloured = supposed distribution of the “Sands of Seese” (from

Autorenkollektiv 2010).


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Fig 15. Lithostratigraphic correlation chart of the Lausitzer (Lusatian) Paleogene and Neogene. Global

sea level changes and local sediment records (from Autorenkollektiv 2010).


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Fig. 16: Model of successions of plant taphocoenosis wothin the middle Miocene 2nd

Lignite Seam

Complex of the Lausitz, east Germany (Schneider in Nowell et al. 1995).


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Useful information


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