Cyclamen: time, sea and speciation biogeography using a temporally calibrated phylogeny

Aim The Mediterranean region is a species-rich area with a complex geographical history. Geographical barriers have been removed and restored due to sea level changes and local climatic change. Such barriers have been proposed as a plausible mechanism driving the high levels of speciation and endemism in the Mediterranean basin. This raises the fundamental question: is allopatric isolation the mechanism by which speciation occurs? This study explores the potential driving influence of palaeo-geographical events on the speciation of Cyclamen (Myrsinaceae), a group with most species endemic to the Mediterranean region. Cyclamen species have been shown experimentally to have few genetic barriers to hybridization. Location The Mediterranean region, including northern Africa, extending eastwards to the Black Sea coast. Methods A generic level molecular phylogeny of Myrsinaceae and Primulaceae is constructed, using Bayesian approximation, to produce a secondary age estimate for the stem lineage of Cyclamen. This estimate is used to calibrate temporally an infrageneric phylogeny of Cyclamen, built with nrDNA ITS, cpDNA trnL-F and cpDNA rps16 sequences. A biogeographical analysis of Cyclamen is performed using dispersal-vicariance analysis. Results The emergence of the Cyclamen stem lineage is estimated at 30.1-29.2 Ma, and the crown divergence at 12.9-12.2 Ma. The average age of Cyclamen species is 3.7 Myr. Every pair of sister species have mutually exclusive, allopatric distributions relative to each other. This pattern appears typical of divergence events throughout the evolutionary history of the genus. Main conclusions Geographical barriers, such as the varying levels of the Mediterranean Sea, are the most plausible explanation for speciation events throughout the phylogenetic history of Cyclamen. The genus demonstrates distributional patterns congruent with the temporally reticulate palaeogeography of the Mediterranean region.

Lake Son Kol geochemistry, µXRF, biogeochemistry and pollen record

In general, a moderate drying trend is observed in mid-latitude arid Central Asia since the Mid-Holocene, attributed to the progressively weakening influence of the mid-latitude Westerlies on regional climate. However, as the spatio-temporal pattern of this development and the underlying climatic mechanisms are yet not fully understood, new high-resolution paleoclimate records from this region are needed. Within this study, a sediment core from Lake Son Kol (Central Kyrgyzstan) was investigated using sedimentological, (bio)geochemical, isotopic, and palynological analyses, aiming at reconstructing regional climate development during the last 6000 years. Biogeochemical data, mainly reflecting summer moisture conditions, indicate predominantly wet conditions until 4950 cal. yr BP, succeeded by a pronounced dry interval between 4950 and 3900 cal. yr BP. In the following, a return to wet conditions and a subsequent moderate drying trend until present times are observed. This is consistent with other regional paleoclimate records and likely reflects the gradual Late Holocene diminishment of the amount of summer moisture provided by the mid-latitude Westerlies. However, climate impact of the Westerlies was apparently not only restricted to the summer season but also significant during winter as indicated by recurrent episodes of enhanced allochthonous input through snowmelt, occurring before 6000 cal. yr BP and at 5100-4350, 3450-2850, and 1900-1500 cal. yr BP. The distinct ~1500-year periodicity of these episodes of increased winter precipitation in Central Kyrgyzstan resembles similar cyclicities observed in paleoclimate records around the North Atlantic, likely indicating a hemispheric-scale climatic teleconnection and an impact of North Atlantic Oscillation (NAO) variability in Central Asia.

Palynology and radiocarbon dating on sediment profile PG1756 from Lake Billyakh, NE Siberia, Russia

In this study a radiocarbon-dated pollen record from Lake Billyakh (65°17'N, 126°47'E; 340 m a.s.l.) in the Verkhoyansk Mountains was used to reconstruct vegetation and climate change since about 15 kyr BP (1 kyr=1000 cal. yr). The pollen record and pollen-based biome reconstruction suggest that open cool steppe and grass and sedge tundra communities with Poaceae, Cyperaceae, Artemisia, Chenopodiaceae, Caryophyllaceae and Selaginella rupestris dominated the area from 15 to 13.5 kyr BP. On the other hand, the constant presence of Larix pollen in quantities comparable to today's values points to the constant presence of boreal deciduous conifer trees in the regional vegetation during the last glaciation. A major spread of shrub tundra communities, including birch (Betula sect. Nanae), alder (Duschekia fruticosa) and willow (Salix) species, is dated to 13.5-12.7 kyr BP, indicating a noticeable increase in precipitation toward the end of the last glaciation, particularly during the Allerød Interstadial. Between 12.7 and 11.4 kyr BP pollen percentages of herbaceous taxa rapidly increased, whereas shrub taxa percentages decreased, suggesting strengthening of the steppe communities associated with the relatively cold and dry Younger Dryas Stadial. However, the pollen data in hand indicate that Younger Dryas climate was less severe than the climate during the earlier interval from 15 to 13.5 kyr BP. The onset of the Holocene is marked in the pollen record by the highest values of shrub and lowest values of herbaceous taxa, suggesting a return of warmer and wetter conditions after 11.4 kyr BP. Percentages of tree taxa increase gradually and reach maximum values after 7 kyr BP, reflecting the spread of boreal cold deciduous and taiga forests in the region. An interval between 7 and 2 kyr BP is noticeable for the highest percentages of Scots spine (Pinus subgen. Diploxylon), spruce (Picea) and fir (Abies) pollen, indicating mid-Holocene spread of boreal forest communities in response to climate amelioration and degradation of the permafrost layer.

Pollen record and age determinations on four sites from Taymyr Peninsula, Russia

Pollen data from a Levinson-Lessing Lake sediment core (74°28'N, 98°38'E) and Cape Sabler, Taymyr Lake permafrost sequences (74°33'N, 100°32'E) reveal substantial environmental changes on the northern Taymyr Peninsula during the last c. 32 000 14C years. The continuous records confirm that a scarce steppe-like vegetation with Poaceae, Artemisia and Cyperaceae dominated c. 32 000-10 300 14C yr BP, while tundra-like vegetation with Oxyria, Ranunculaceae and Caryophyllaceae grew in wetter areas. The coldest interval occurred c. 18 000 yr BP. Lateglacial pollen data show several warming events followed by a climate deterioration c. 10 500 14C yr BP, which may correspond with the Younger Dryas. The Late Pleistocene/Holocene transition, c. 10 300-10 000 14C yr BP, is characterized by a change from the herb-dominated vegetation to shrubby tundra with Betula sect. Nanae and Salix. Alnus fruticosa arrived locally c. 9000-8500 14C yr BP and disappeared c. 4000-3500 14C yr BP. Communities of Betula sect. Nanae, broadly distributed at c. 10 000-3500 14C yr BP, almost disappeared when vegetation became similar to the modern herb tundra after 3500-3000 14C yr BP. Quantitative climate reconstructions show Last Glacial Maximum summer temperature about 4°C below the present and Preboreal (c. 10 000 14C yr BP) temperature 2-4°C above the present. Maximum summer temperature occurred between 10 000 and 5500 14C yr BP; later summers were similar to present or slightly warmer.

Pollen record from Kleiner Mochowsee, Germany

A Holocene pollen diagram from Kleiner Mochowsee (northern Niederlausitz, East Germany) shows pine as an important constituent of the woodland south of the Schwielochsee. Oak woodland was widespread since the Atlantic. Betula lost its importance at the end of the Preboreal. Fagus is represented continuously in the pollen record since the Atlantic, Carpinus since the Subboreal. However, the two latter tree species remain without great importance throughout the whole pollen record. The poor sandy soils are furthermore reflected by the low values of Corylus during the Boreal, comparable to other records from Berlin and its surrounding area. The 'classical' elm decline could be shown for the Niederlausitz, radiocarbon dates assume a contemporaneous age for this event with other records from northern Germany. Only small-scaled human impact is indicated in prehistoric times, during the migration period it seems to have ceased completely. Later, in the Medieval, deforestation and tillage can be shown. Secale was cultivated since the early Medieval; an accompanying weed flora appeared at the same time. Cultivation of Fagopyrum and Linum usitatissimum could be shown for the late Medieval times.

Palynology on sediment profile Tso Kar in Ladakh

Palynological investigation of a 410 cm long core section from Tso Kar (33°10'N, 78°E, 4527 m a.s.l.), an alpine lake situated in the arid Ladakh area of NW India at the limit of the present-day Indian summer monsoon, was performed in order to reconstruct post-glacial regional vegetation and climate dynamics. The area was covered with alpine desert vegetation from ca. 15.2 to 14 kyr BP (1 kyr=1000 cal. years), reflecting dry and cold conditions. High influx values of long-distance transported Pinus sylvestris type pollen suggest prevailing air flow from the west and northwest. The spread of alpine meadow communities and local aquatic vegetation is a weak sign of climate amelioration after ca. 14 kyr BP. Pollen data (e.g. influx values of Pinus roxburghii type and Quercus) suggest that this was due to a strengthening of the summer monsoon and the reduced activity of westerly winds. The further spread of Artemisia and species-rich meadows occurred in response to improved moisture conditions between ca. 12.9 and 12.5 kyr BP. The subsequent change towards drier desert-steppe vegetation likely indicates more frequent westerly disturbances and associated snowfalls, which favoured the persistence of alpine meadows on edaphically moist sites. The spread of Chenopodiaceae-dominated vegetation associated with an extremely weak monsoon occurred at ca. 12.2-11.8 kyr BP during the Younger Dryas interstadial. A major increase in humidity is inferred from the development of Artemisia-dominated steppe and wet alpine meadows with Gentianaceae after the late glacial/early Holocene transition in response to the strengthening of the summer monsoon. Monsoonal influence reached maximum activity in the Tso Kar region between ca. 10.9 and 9.2 kyr BP. The subsequent development of the alpine meadow, steppe and desert-steppe vegetation points to a moderate reduction in the moisture supply, which can be linked to the weaker summer monsoon and the accompanying enhancement of the winter westerly flow from ca. 9.2 to 4.8 kyr BP. The highest water levels of Tso Kar around 8 kyr BP probably reflect combined effect of both monsoonal and westerly influence in the region. An abrupt shift towards aridity in the Tso Kar region occurred after ca. 4.8 kyr BP, as evidenced by an expansion of Chenopodiaceae-dominated desert-steppe. Low pollen influx values registered ca. 2.8-1.3 kyr BP suggest scarce vegetation cover and unfavourable growing conditions likely associated with a further weakening of the Indian Monsoon.

Pollen records and lithology of profiles from Lobsigensee, Switzerland

Lobsigensee is a small kettle hole lake 15 km north-west of Bern on the Swiss Plateau, at an altitude of 514 m asl. Its surface is 2ha today, its maximum depth 2.7 m; it has no inlet and the overflow functions mainly during snow melting. The area was covered by Rhone ice during the Last Glaciation (map in Fig.2). Local geology, climate and vegetation are summarized in Figure 3A-C, the history of settlement in Figures 5-7. In order to reconstruct the vegetational and environmental history of the lake and its surroundings pollen analysis and other bio- and isotope stratigraphies were applied to twelve profiles cored across the basin with modified Livingstone corers (Fig.3 D). (1) The standard diagram: The central core LQ-90 is described as the standard pollen diagram (Chapter 3) with 10 local pollen assemblage zones of the Late-Glacial (local PAZ Ll to Ll0, from about 16'000(7) to 10'000 years BP) and 20 PAZ of the Holocene (local PAZ L11 to L30), see Figs. 8-10 and 20-24. Local PAZ L 1 to L3 are in the Late-Glacial clay and record the vegetational development after the ice retreat: L1 shows very low pollen concentration and high Pinus percentages due to long-distance transport and reworking; the latter mechanism is corroborated by the findings of thermophilous and pre-Quaternary taxa. Local PAZ L2 has a high di versi ty of non-arboreal pollen (NAP) and reflects the Late-Glacial steppe rich in heliophilous species. Local PAZ L3 is similar but additionally rich in Betula nana and Sal1x, thus reflecting a "shrub tundra". The PAZ L1 to L3 belong to the Oldest Dryas biozone. Local PAZ L4 to L 10 are found in the gyttja of the profundal or in the lake marl of the littoral and record the Late-Glacial forests. L4 is the shrub phase of reforestation with very high Junlperus and rapidly increasing Betula percentages. L5 is the PAZ with a first, L7 with a second dominance of tree-birches, separated by L6 showing a depression in the Betula curve. L4 to L7 can be assigned to the Balling biozone. Possible correlation of the Betula depression to the Older Dryas biozone is discussed. In local PAZ L8 Plnus immigrates and expands. L9 shows a facies difference in that Plnus dominates over Betula in littoral but not in profundal spectra. L8 and L9 belong to the Allerod biozone. In its youngest part the volcanic ash from Laach/Eifel is regularly found (11,000 BP). The local PAZ Ll0 corresponds to the Younger Dryas blozone. The merely slight increase of the NAP indicates that the pine forests of the lowland were not strongly affected by a cooler climate. In order to evaluate the significance of the littoral accumulation of coniferous pollen the littoral profile LQ-150 is compared to the profundal. Radiocarbon stratigraphies derived from different materials are presented in Figures 13 and 14 and in Tables 2 and 3. The hard-water errors in the gyttja samples and the carbonate samples are similar. The samples of terrestrial plant macrofossils are not affected by hard-water errors. Two plateaux of constant age appear in the age-depth relationship; their consequence for biostratigraphy as well as pollen concentration and influx diagrams are discussed. Radiocarbon ages of the Late-Glacial pollen zones are shown in Table 10. The Holocene vegetational history is recorded in the local PAZ L 11 to L30. After a Preboreal (PAZ L11) dominated by pine and birch the expansions of Corylus, Ulmus and Quercus are very rapid. Among these taxa Corylus dominates dur ing the Boreal (PAZ L 12 and L 1 3), whereas the components of the mixed oak forest dominate in the Older Atlantic (PAZ L14 to L16). In the Younger Atlantic (PAZ L 17 to L 19) Fagus and Alnus play an increasing, the mixed oak forest a decreasing role. During the period of local PAZ L19 Neolithic settlers lived on the shore of Lobsigensee. During the Subboreal (PAZ L20 and L21) and the Older Subatlantic (L22 to L25) strong fluctuations of Fagus and often antagonistic peaks of NAP, Alnus, Betula and Corylus can be interpreted as signs of human impact on vegetation. L23 is characterized not only by high values of NAP (especially apophytes and anthropochorous species) but also by the appearance of Juglans, Castanea and Secale which point to the Roman colonization of the area. For a certain period during the Younger Subatlantic (PAZ L26 to L30) the lake was used for retting hemp (Cannabis). Later the dominance of Quercus pollen indicates the importance of wood pastures. The youngest sediments reflect the wide-spread agricultural grass lands and the plantation of Pinus and Picea. Radiocarbon dates for the Holocene are given in Figure 23 and Table 4, the extrapolated ages of the Holocene pollen zones in Table 15. (2) The cross sections: Figures 25 and 26 give a summary of the litho- and palynostratigraphy of the two cross sections. Based on 11 Late-Glacial and 9 Holocene pollen diagrams (in addition to the standard ones), the consistency of the criteria for the definition of the pollen zones is examined in Tables 7 and 8 for the Late-Glacial and in Tables 11 to 14 for the Holocene. Sediment thicknesses across the basin for each pollen zone are presented in these tables as well as in Figures 43 to 45 for the Late-Glacial and in Figures 59 to 65 for the Holocene. Sediment focusing can explain differences between the gyttja cores of the profundal. Focusing is more than compensated for through "stretching" by carbonate precipitation on the littoral terrace. Pollen influx to the cross section are discussed (Chapters 4.1.5. and 4.2.3.). (3) The regional pollen zones: Based on some selected sites between Lake Geneva and Lake Constance regional pollen zones are proposed (Table 16, 17 and 19). (4) Paleoecology: Climatic change in the Late-Glacial can be inferred from Coleoptera, Trichoptera, Chironomidae and d18O of carbonates: a distinct warming is recorded around 12' 600 BP and around 10' 000 BP. The Younger Dryas biozone (10'700-10'000 BP) was the only cooling found in the Late-Glacial. The Betula depression often correlated wi th the Older Dryas biozone was possibl not colder but dryer than the previous period. During the Holocene the lowland site is not very sensitive to the minor climatic changes. Table 22 summarizes climatic and trophic changes before 8'000 BP as deduced from various biostratigraphies studied by a number of authors. Ostracods, Chironomids and fossil pigments indicate that anoxic conditions prevailed during the BoIling (possibly meromixis). Changes in the lake level are illustrated in Figure 74. A first lake-level lowering occurred in the early Holocene (10'000 to 9'000 BP), a second during the Atlantic (about 6'800 to 5'200 BP). The first "shrinking" of the lake volume resulted in a eutrophication recorded by laminations in the profundal and by pigments of Cyanophyceae. The second fall in water level corresponds to an increase of Nymphaeaceae. Human impact can be inferred in three ways: eutrophication of the lake (since the Neolithic), changes of terrestrial vegetation by deforestations (cyclicity of Fagus, see Figures 78 to 80), and enhanced erosion (increasing sedimentation rates by inwashed clay, particularly since the Roman Colonization, see Figures 49 and 81). Summary: This paper was planned as the final report on Lobsigensee. However, a number of issues are not answered but can only be asked more precisely, for example: (1) For the two periods with the highest rates of change, Le. the Bolling and the Preboreal biozones, pollen influx may reflect vegetation dynamics. Detailed investigations of these periods in annually laminated sediments are planned. (2) Biostratigraphies other than palynostratigraphy are needed to estimate the degree of linkage or independence in the development of terrestrial and lacustrine ecosystems. Often our sampling intervals were not identical, thus influencing our temporal resolution. (3) 6180- and 14C-stratigraPhies with high resolution will elucidate the leads and lags of these dynamic periods. Plateaux of constant age in the age-depth relationship have a strong bearing on both biological and geophysical understanding of Late-Glacial and early Holocene developments. (4) Numerical methods applied to the pollen diagrams of the cross section will help to quantify the significance of similari ties and dissimilarities across a single basin (with Prof. Birks). (5) Numerical methods applied to different sites on the Swiss Plateau and on the transect across the Alps will be helpful in evaluating the influence of different environmental factors (with Prof. Birks). (6) A new map 1: 1000 with 50cm-contour lines prov ided by Prof. Zurbuchen will be combined with a grid of cores sampling the transition from lake marl to peat enabling us to calculate paleo-volumes of the lake. This is interesting for the two "shrinking periods" (in Fig. 74A numbers 2-6 and 7-10), both accompanied by eutrophication. The pal eo-volume during the Neoli thic set tlement of the Cortaillod culture linked wi th an est l.mate of trophic change derived from diatoms (Prof. Smol in prep.) could possibly give an indication of the size of the human population of this period. (7) For the period with the antagonism between Fagus peaks and ABC-peaks close collaboration between palynologists, geochemists and archeologists should enable us to determine the influence of prehistoric and historic people on vegetation (collaboration with Prof. Stockli and Prof. Herzig). (8) The core LL-75 taken with a "cold letter box" will be analysed for major and trace elements by Dr. Sturm for 210pb and 137Cs by Prof.von Gunten and for pollen. We will see if our local PAZ L30 really corresponds to the surface sediment and if the small seepage lake reflects modern pollution.

Palynology on profile Rotmoss at Obergurgl in Austria

Die pollenanalytische Untersuchung des Rotmooses in Verbindung mit C-14 Daten hat ergeben, daß die organogenen Sedimente nachwärmezeitliche Bildungen sind. Ein Gletschervorstoß um 2500 v. Chr. konnte mit Hilfe der C-14 Daten eingegrenzt und mit anderen Fundstellen parallelisiert werden. Weitere pollenanalytisch festgestellte Gletscher und auch Waldgrenzschwankungen konnten festgestellt, müssen aber noch genau datiert und parallelisiert werden.

Pollen record and dating of a sediment profile from Lake Voulkaria, Greece

A palynological investigation of a Holocene profile from Lake Voulkaria, western Greece, was carried out as a contribution to the environmental history of the coastal area of northwestern Acarnania and the Classical city of Palairos. It shows that deciduous oaks dominated the natural vegetation of the area throughout the Holocene. Until ca. 7000 B.C. Pistacia occurred abundantly, while other evergreen woody taxa were rare. At ca. 6300 B.C. an expansion of Carpinus orientalis/Ostrya can be observed. Around ca. 5300 B.C. spreading of Erica indicates a change to a drier climate and/or first human impact. Since ca. 3500 B.C. an increase of evergreen shrubs now clearly indicates land-use. The foundation of the Classical city of Palairos led to a temporary expansion of Phillyrea maquis. Within this period, molluscs of brackish water indicate the use of the lake as a harbour after the construction of a connection to the sea. The deciduous Quercus woodland recovered when human impact decreased in the area, and lasted until modern times.