Molekulare Studien zur Evolution der Myrmekophytie in der Gattung Macaranga (Euphorbiaceae)

Im Rahmen der vorliegenden Arbeit wurde eine detaillierte phylogenetische Analyse der Ameisenpflanzen aus der Gattung Macaranga (Euphorbiaceae) und ihres verwandtschaftlichen Umfelds mit Hilfe von AFLP-Fingerprinting („amplified fragment length polymorphisms“) sowie vergleichender Analyse von mehreren nichtkodierenden Chloroplasten-DNA-Loci vorgenommen. Anhand dieser Untersuchungen sollten im Wesentlichen die folgenden Fragen geklärt werden: (1) Wie stellen sich die Verwandtschaftsverhältnisse zwischen den myrmekophytischen Macaranga-Sektionen Pachystemon, Winklerianae und Pruinosae dar? (2) Wie sind die einzelnen Arten dieser Sektionen miteinander verwandt? (3) Wie oft ist die Lebensweise ”Myrmekophytie” unabhängig voneinander entstanden? Gibt es Hinweise auf Reversionen? (4) Wo liegt genealogisch und auch geographisch der Ursprung der Symbiose zwischen den myrmekophytischen Macaranga-Arten und ihren Partnerameisen? (5) Welche Bedeutung spielen koevolutive Entwicklungen für das Macaranga-Crematogaster-Symbiosesystem? Ist Myrmekophytie im Sinne einer Schlüsselinnovation (Givnish, 1997) als Stimulus für eine adaptive Radiation zu betrachten? (1) Für die AFLP-Analyse wurden 108 Proben aus 43 Macaranga-Arten und 5 unbeschriebenen Morphospezies in die phylogenetische Untersuchung einbezogen. Auf der Basis von 426 Merkmalen wurden Phänogramme sowie Kladogramme rekonstruiert. Zur statistischen Absicherung wurden Bootstrap-Analysen durchgeführt und im Falle der Kladogramme darüber hinaus der „consistency“-Index bestimmt. Die AFLP-Datensätze wurden zusätzlich einer Hauptkomponentenanalyse unterzogen. Mit Hilfe der verschiedenen Untersuchungsmethoden konnten weitgehend übereinstimmende Gruppierungen bzw. evolutive Linien identifiziert werden. Die Sektionen Pachystemon und Pruinosae bilden eine jeweils gut gestützte monophyletische Gruppe. Beide sind vermutlich Schwestergruppen und damit gleich alt. Für die Monophylie der nur aus zwei Arten bestehenden Sektion Winklerianae ergab sich keine Unterstützung. Die Arten der Sektion Pruinosae sind im AFLP-Baum gut aufgelöst. Die nicht myrmekophytische M. gigantea sitzt dabei an der Basis und ist Schwestergruppe zu den myrmekophytischen Arten. Innerhalb der Sektion Pachystemon wurden mit Hilfe der AFLP-Analyse vier gut gestützte Gruppen identifiziert. Für die puncticulata-Gruppe konnte hier erstmals auf molekularer Ebene eine Zugehörigkeit zur Sekt. Pachystemon nachgewiesen werden. Der von Davies (2001) vorgenommene Ausschluss von M. recurvata aus der Sekt. Pachystemon konnte bestätigt werden. Die Verwandtschaftsbeziehungen einzelner Arten zueinander sind in den AFLP-Bäumen nicht aufgelöst. (2) Für die vergleichende Chloroplasten-Sequenzierung wurden nach Maßgabe der Sequenzvariabilität in Testsequenzierungen die Bereiche atpB-rbcL und psbI-trnS für die phylogenetische Untersuchung ausgewählt. Für die Chloroplasten-Phylogenie wurden für jeden Locus mehr als 100 Sequenzen analysiert. Neben 29 Pachystemon-Arten inkl. vier unbekannter Morphospezies, acht Pruinosae-Arten inkl. eines möglichen Hybriden und den beiden Arten der Sekt. Winklerianae wurden 22 weitere Macaranga- und 10 Mallotus-Arten in die Untersuchung einbezogen. Zwischen den südostasiatischen Arten bestanden nur geringe Sequenzunterschiede. Maximum-Parsimonie-Kladogramme wurden rekonstruiert und die Sequenzen der beiden Loci wurden sowohl einzeln, als auch kombiniert ausgewertet. Indels wurden kodiert und als separate Merkmalsmatrix an die Sequenzdaten angehangen. Innerhalb von Macaranga konnten nur wenige abgesicherte Gruppen identifiziert werden. Deutlich war die Zusammengehörigkeit der afrikanischen Arten und ihr Entstehung aus den südostasiatischen Arten. Die von Davies (2001) der Sektion Pruinosae zugeordnete M. siamensis steht deutlich außerhalb dieser Sektion. Die Arten der Sektionen Pruinosae, Pachystemon und Winklerianae bilden keine statistisch gesicherten monophyletischen Gruppen. Während der Pilotstudien stellte sich heraus, dass die Chloroplastensequenzen nahe verwandter Arten der Sektion Pachystemon weniger nach den Artgrenzen, sondern vielmehr nach geographischen Kriterien gruppierten. (3) Es wurde daher zusätzlich eine phylogeographische Analyse der Chloroplasten-Sequenzen auf der Basis eines Parsimonie-Netzwerks durchgeführt. Neben dem atpB-rbcL-Spacer und einer Teilsequenze des psbI-trnS-Locus (ccmp2) wurde dafür zusätzlich der ccmp6-Locus (ein Abschnitt des ycf3-Introns) sequenziert. Die phylogeographische Untersuchung wurde mit 144 Proben aus 41 Macaranga-Arten durchgeführt. Darin enthalten waren 29 Arten (inkl. vier Morphospezies) mit 112 Proben der Sektion Pachystemon, sieben 7 Arten (inkl. eines potentiellen Hybriden) mit 22 Proben der Sekt. Pruinosae und zwei Arten mit 5 Proben der Sekt. Winklerianae. Das voll aufgelöste statistische Parsimonie-Netzwerk umfasste 88 Haplotypen. Die Sektionen Pachystemon und Pruinosae bilden jeweils eine monophyletische Gruppe. Das geographische Arrangement der Haplotypen unabhängig von der Artzugehörigkeit könnte durch Introgression und/oder „lineage sorting“ bedingt sein. Mit Hilfe der im Rahmen dieser Arbeit gewonnenen Ergebnisse kann man davon ausgehen, dass eine enge Ameisen-Pflanzen-Symbiose innerhalb der Gattung Macaranga mindestens drei-, möglicherweise viermal unabhängig voneinander entstanden ist Eine Reversion hat mindestens einmal, möglicherweise häufiger in der bancana-Gruppe stattgefunden. Ob sich die Symbiose dabei in Westmalaysia oder in Borneo entwickelt hat, kann man nicht sicher sagen; Ob und inwieweit die große Artenzahl in der bancana-Gruppe als eine Folge der Myrmekophytie anzusehen ist, bleibt zunächst offen. Wesentliche Teile der vorliegenden Arbeit liegen bereits in publizierter Form vor (AFLP-Analyse: Bänfer et al. 2004; Chloroplasten-Analyse: Vogel et al. 2003; Bänfer et al. 2006).

Compound specific stable isotopes, BIT, pollen and spores of Miocene to Pliocene sediments of ODP Hole 175-1085A

Pollen and stable carbon (d13C) and hydrogen (dD) isotope ratios of terrestrial plant wax from the South Atlantic sediment core, ODP Site 1085, is used to reconstruct Miocene to Pliocene changes of vegetation and rainfall regime of western southern Africa. Our results reveal changes in the relative amount of precipitation and indicate a shift of the main moisture source from the Atlantic to the Indian Ocean during the onset of a major aridification 8 Ma ago. We emphasise the importance of declining precipitation during the expansion of C4 and CAM (mainly succulent) vegetation in South Africa. We suggest that the C4 plant expansion resulted from an increased equator-pole temperature gradient caused by the initiation of strong Atlantic Meridional Overturning Circulation following the shoaling of the Central American Seaway during the Late Miocene.

Palynological record and radiocarbon dates for sediment core T89-16, Congo River mouth

We present a high-resolution reconstruction of tropical palaeoenvironmental changes for the last deglacial transition (18 to 9 cal. kyr BP) based on integrated oceanic and terrestrial proxies from a Congo fan core. Pollen, grass cuticle, Pediastrum and dinoflagellate cyst fluxes, sedimentation rates and planktonic foraminiferal d18O ratios, uK37 sea-surface temperature and alkane/alkenone ratio data highlight a series of abrupt changes in Congo River palaeodischarge. A major discharge pulse is registered at around 13.0 cal. kyr BP which we attribute to latitudinal migration of the Intertropical Convergence Zone (ITCZ) during deglaciation. The data indicate abrupt and short-lived changes in the equatorial precipitation regime within a system of monsoonal dynamics forced by precessional cycles. The phases of enhanced Congo discharge stimulated river-induced upwelling and enhanced productivity in the adjacent ocean.

AMS radiocarbon dates and pollen analysis of ODP Hole 175-1078C

ODP Site 1078 situated under the coast of Angola provides the first record of the vegetation history for Angola. The upper 11 m of the core covers the past 30 thousand years, which has been analysed palynologically in decadal to centennial resolution. Alkenone sea surface temperature estimates were analysed in centennial resolution. We studied sea surface temperatures and vegetation development during full glacial, deglacial, and interglacial conditions. During the glacial the vegetation in Angola was very open consisting of grass and heath lands, deserts and semi-deserts, which suggests a cool and dry climate. A change to warmer and more humid conditions is indicated by forest expansion starting in step with the earliest temperature rise in Antarctica, 22 thousand years ago. We infer that around the period of Heinrich Event 1, a northward excursion of the Angola Benguela Front and the Congo Air Boundary resulted in cool sea surface temperatures but rain forest remained present in the northern lowlands of Angola. Rain forest and dry forest area increase 15 thousand years ago. During the Holocene, dry forests and Miombo woodlands expanded. Also in Angola globally recognised climate changes at 8 thousand and 4 thousand years ago had an impact on the vegetation. During the past 2 thousand years, savannah vegetation became dominant.

Early - Late Miocene palaeoclimate data and palaefloras of 48 samples from Southern China

Southern China, especially Yunnan, has undergone high tectonic activity caused by the uplift of Himalayan Mountains during the Neogene, which led to a fast changing palaeogeography. Previous study shows that Southern China has been influenced by the Asian Monsoon since at least the Early Miocene. However, it is yet not well understood how intense the Miocene monsoon system was. In the present study, 63 fossil floras of 16 localities from Southern China are compiled and evaluated for obtaining available information concerning floristic composition, stratigraphic age, sedimentology, etc. Based on such reliable information, selected mega- and micro-floras have been analysed with the coexistence approach to obtain quantitative palaeoclimate data. Visualization of climate results in maps shows a distinct spatial differentiation in Southern China during the Miocene. Higher seasonalities of temperature and precipitation occur in the north and south parts of Southern China, respectively. During the Miocene, most regions of Southern China and Europe were both warm and humid. Central Eurasia was likely to be an arid center, which gradually spread westward and eastward. Our data provide information about Miocene climate patterns in Southern China and about the evolution of these patterns throughout the Miocene, and is also crucial to unravel and understand the climatic signals of global cooling and tectonic uplift.

Distribution of pollen and spores in sediments of ODP Hole 175-1078C

The distribution of pollen in marine sediments is used to record vegetation change on the continent. Generally, a good latitudinal correspondence exists between the distribution patterns of pollen in the marine surface sediments and the occurrence of the source plants on the adjacent continent. To investigate land-sea interactions during deglaciation, we compare proxies for continental (pollen assemblages) and marine conditions (alkenone-derived sea surface temperatures) of two high-resolution, radiocarbon-dated sedimentary records from the tropical southeast Atlantic. The southern site is located West of the Cunene River mouth; the northern site is located West of the Angolan Huambe Mountains. It is inferred that the vegetation in Angola developed from Afroalpine and open savannah during the last Glacial maximum (LGM) via Afromontane Podocarpus forest during Heinrich Event 1 (H1), to an early increase of lowland forest after 14.5 ka. The vegetation record indicates dry and cold conditions during the LGM, cool and wet conditions during H1 and a gradual rise in temperature starting well before the Younger Dryas (YD) period. Terrestrial and oceanic climate developments seem largely running parallel, in contrast to the situation ca. 5° further South, where marine and terrestrial developments diverge during the YD. The cool and wet conditions in tropical West Africa, South of the equator, during H1 suggest that low-latitude insolation variation is more important than the slowdown of the thermohaline circulation for the climate in tropical Africa.

Palynology of marine sediment cores from the West African coast

Seven sediment cores from the cruises of the "Meteor" and "Valdivia" were examined palynologically. The cores were retrieved from the lower continental slope in the area of between 33.5° N and 8° N, off the West African coast. Most of the cores contain sediments from the last Glacial and Interglacial period. In some cases, the Holocene sediments are missing. Some individual cores contain sediments also from earlier Glacial and Interglacial periods. The main reason for making this palynological study was to find out the differences between the vegetation of Glacial and Interglacial periods in those parts of West Africa which at present belong to the Mediterranean zone, the Sahara and the zones of the savannas and tropical forests. In today's Mediterranean vegetation zone at core 33.5° N, forests and deciduous forests in particular, are missing during Glacial conditions. Semi-deserts are found instead of these. In the early isotope stage 1, there is a very significant development of forests which contain evergreen oaks; this is the Mediterranean type of vegestation development. The Sahara type of vegetation development is shown in four cores from between 27° N and 19° N. The differences between Glacial and Interglacial periods are very small. It must be assumed therefore that in this latitudes, both Glacial and Interglacial conditions gave rise to desert generally. The results are in favour of a slightly more arid climate during Glacial and more humid one during Interglacial periods. The southern boundary of the Sahara and the adjacent savannas with grassland and tropical woods were situated more to the south during the Glacial periods than they were during the Interglacial ones. In front of today's savanna belt, it can be seen from the palynological results that there are considerable differences between the vegetation of Glacial and Interglacial periods. The woods are more important in Interglacial periods. During the Glacial periods these are replaced from north to south decreasingly by grassland (savanna and rainforest type of vegetation development). The southern limit of the Sahara during stage 2 was somewhat between 12° N and 8° N which is between 1.5 and 5 degrees in latitude further south than it i s today. Not only do these differences in climate and vegetation apply to the maximum of the last Glacial and for the Holocene, but they apparently apply also to the older Glacial and Interglacial periods, where they have been found in the profiles. The North African deset belt can be said to have expanded during Glacial times both towards the north and towards the south. All the available evidence of this study indicates that the grass land or the semi-desert of the Southern Europe cam einto connection with those of the N Africa; there could not have been any forest zone between them. The present study was also a good opportunity for investigating some of the basic marine palynological problems. The very well known overrepresentation of pollen grains of the genus Pinus in marine sediments can be traced as fa as 21° N. The present southern limit for the genus Pinus is on the Canaries and on the African continent as approximately 31° N. Highest values of Ephedra pollen grains even occur south of the main area of the present distribution of that genus. These does not seem to be any satisfactory explanation for this. In general, it would appear that the transport of pollen grains from the north is more important than transport from the south. The results so far, indicate strongly that further palynological studies are necessary. These should concentrate particularly on cores from between 33° N and 27° N as well as between 17° N and 10° N. It would also be useful to have a more detailed examination of sediments from the last Intergalcial period (substage 5 e). Absolute pollen counts and more general examination of surface samples would be desirable. Surface samples should be taken from the shelf down to the bottom of the continental slope in different latitudes.

Palynology of ODP Site 108-658

Ocean Drilling Program Site 658 at 21°N off northwest Africa has a high sedimentation rate and a high concentration of pollen grains and is thus very suitable for detailed pollen analysis. The time scale for the upper 100 m (the last 670 k.y.) of Site 658 is based on biostratigraphic data and isotope stratigraphy. The pollen record has been divided into 34 zones. These are classified into 7 zone types covering a range from very arid to rather humid conditions. The sequence shows a long-term climatic decline: strong glacial stages were found only after 480 k.y. and strong interglacial stages only before 280 k.y. The Site 658 record correlates well with a terrestrial sequence from northern Greece, although both records differ in their response to global climatic change. Spectral analysis shows a 100- and a 42-k.y. period in the curves of pollen brought in by the northwest trade winds and only a 42-k.y. period in the curves of pollen mostly transported by the African Easterly Jet. A 31-k.y. period is found in the curves for Ephedra and Chenopodiaceae-Amaranthaceae. In addition, Ephedra shows a 54-k.y. period.

Pollen analysis of sediment cores from the mouth of the river Congo to Walvis Bay and Lüderitz

The distribution of pollen in marine sediments is used to record vegetation changes over the past 30,000 years on the adjacent continent. A transect of marine pollen sequences from the mouth of the river Congo (~5°S) to Walvis Bay and Lüderitz (~25°S) shows vegetation changes in Congo, Angola and Namibia from the last glacial period into the Holocene. The comparison of pollen records from different latitudes provides information about the latitudinal shift of open forest and savannahs (Poaceae pollen), the extension of lowland forest (rain forest pollen) and Afromontane forest (Podocarpus pollen), and the position of the desert fringe (pollen of Caryophyllaceae, Chenopodiaceae and Amaranthaceae). High Cyperaceae pollen percentages in sediments from the last glacial period off the mouth of the river Congo suggest the presence of open swamps rather than savannah vegetation in the Congo Basin. Pollen from Restionaceae in combination with Stoebe-type pollen (probably from Elytropappus) indicates a possible northwards extension of winter rain vegetation during the last glacial period. The record of Rhizophora (mangrove) pollen is linked to erosion of the continental shelf and sea-level rise. Pollen influx is highest off river mouths (10-2000 grains year**-1 cm**-2), close to the coast (300-6000 grains year**-1 cm**-2), but is an order of magnitude lower at sites situated far from the continent (<10 grains year**-1 cm**-2).

Pollen analysis and age model of sediment core GIK-15856-2

Pollen and spores from a deep-sea core located west of the Niger Delta record an uninterrupted area of lowland rain forest in West Africa from Guinea to Cameroon during the last Interglacial and the early Holocene. During other periods of the last 150 ka, a savanna corridor between the western - Guinean - and the eastern - Congolian - part of the African lowland rain forest existed. This so-called Dahomey Gap had its largest extension during Glacial Stages 6, 4, 3, and 2. Reduced surface salinity in the eastern Gulf of Guinea as recorded by dinoflagellate cysts indicates sufficient precipitation for extensive forest growth during Stages 5 and 1. The large modern extension of dry forest and savanna in West Africa cannot be solely explained by climatic factors. Mangrove expansion in and west of the Niger Delta was largest during the phases of sea-level rise of Stages 5 and 1. During Stages 6, 4, 3, and 2, shelf areas were exposed and the area of the mangrove swamps was minimal.

Pollen analysis of surface sediment off northwest Africa

Over 100 samples of recent surface sediments from the bottomn of the Atlantic Ocean offshore NW Africa between 34° and 6° N have been analysed palynologically. The objective of this study was to reveal the relation between source areas, transport systems, and resulting distribution patterns of pollen and spores in marine sediments off NW Africa, in order to lay a sound foundation for the interpretation of pollen records of marine cores from this area. The clear zonation of the NW-African vegetation (due to the distinct climatic gradient) is helpful in determining main source areas, and the presence of some major wind belts facilitates the registration of the average course of wind trajectories. The present circulation pattern is driven by the intertropical front (ITCZ) which shifts over the continent between c. 22° N (summer position) and c. 4° N (winter position) in the course of the year. Determination of the period of main pollen release and the average atmospheric circulation pattern effective at that time of the years is of prime importance. The distribution patterns in recent marine sediments of pollen of a series of genera and families appear to record climatological/ecological variables, such as the trajectory of the NE trade, January trades, African Easterly Jet (Saharan Air Layer), the northernmost and southernmost position of the intertropical convergence zone, and the extent and latitudinal situation of the NW-African vegetation belt. Pollen analysis of a series of dated deep-sea cores taken between c. 35° and the equator off NW African enable the construction of paleo-distribution maps for time slices of the past, forming a register of paleoclimatological/paleoecological information.