Palynology of the Quaternary in Temperate and Tropical Areas: Chronostratigraphy, Palaeoclimatology and Vegetal Palaeoenvironment of Fossil Man

Home , Biber (geology), Grotte du Vallonnet, Gunz (geology), Pollen zone

297

PALYNOLOGY OF THE QUATERNARY IN TEMPERATE AND TROPICAL AREAS: CHRONOSTRATIGRAPHY, PALAEOCLIMATOLOGY AND VEGETAL PALAEOENVIRONMENT OF FOSSIL MAN

JOSETTE RENAULT-MISKOVSKY" and ANNE-MARIE SEMAH*

+ Unité de Palynologie du Laboratoire de Préhistoire du MNHN, Institut de Paleontologie humaine, 1, Rue René Panhard 75013, Paris, France # Uniié de Palynologie du Laboratoire de Préhistoire du MNHN, Institute de Paleontologie humaine, Orstom, Nouméa, B.P. A5, Nouvelle-Calédonia.

I. PALYNOLOGY Palynology is the study of the shape and meaning of spores and pollen grains, which provide knowledge of fossil plants and reconstruction of their history. Plants produce thousands of spores and pollen grains each year, most of which are lost for reproduction ydare carried at ground level through different vehicles: wind, water, insects, mammals, and even men. Therefore, when the palaeobotanist takes sediments in a site, he collects, at the same time, whatever pollen amount is contained in them. The matters propitious to pollen analysis are mainly the deposits that ensure a good preservation of pollen grains, such as peatbogs, lacustrine and marine deposits. Archaeological deposits are also

particularly useful to palaeoclimatic study; their place in time can be I identified out of human remains, manufactured objects and the degree of evolution of the faunae involved. Additional precision is given to the chronological scale by palaeomagnetic studies and the different radiometric methods of absolute datation. Samplings in aquatic or sub-aquatic media (peats, lacustrine muds or conchitic sands) are done in continuous series through boring and coring. However, prehistorical stratigraphies, most often established in dry milieu involve samplings in light deposits, such as clays, sands, gravel, I

.I . - __ ------Traductions : Jacqueline GAUDEY and Elise HUFFER. I 298 / Current Concepts in Pollen-Spore and Biopollution Research or even deposits consolidated into brecciae along vertical sections cut 'during excavation (Girard, 1975). In the case of indurated deposits (stalagmitic floors], sampling is made by local sawing of blocks and columns, which are more finely cut in the laboratory afterwards. These conventional techniques have been recently completed by the techniques of deep coring throughout the deposits not yet excavated. It may be necessary to take samples in such special material as burials, vases for offerings, coating, or content of mummies, so as to relate a plant deposit, if any, to cultural and even ritual practice. Whatever the gathering, it must Be done quite properly to avoid any contamination. Sample preparation depends on the mineralogical composition of the deposit, but the principle of the most current method, called classical chemical method, consists in sieving oÜt, then dissolving in acids and bases, the mineral and organic matters in which spores and pollen grains are enclosed. As archeological sediments are rarely very fossiliferous, they have to be treated with a special concentration process, based on separating the grains from the rest of the gangue through floatation in dense solution, zinc chloride or Thoulet's liquor (Girard and Renault- Miskovsky, 1969).' The extraordinary properties of the membrane surrounding the grains, or exine, allow them to resist all the chemical corrosioh specific to preparation methods, and also to last almost indefinitelyI sheltered from oxidation through geological times; the grain structure, size, shape, the arrangement, type and number of germinal apertures, and also the exine sculpture, lead to identification of a plant family, genus or even species under the microscope. As a matter of fact, each plant species is represented by one type of grain, which is dete&ed by comparison to one modem grain, as spores and pollen grains have not evolved since modem florae settled on the earth. / Fossil grains, after being determined under the photon microscope, ' are counted. On the basis of statistical calculations, the percentages of the different categories can be referred to the total number of grains collected for a certain weight of the sample (20 to Sog]. This is pollen i analysis per level. Each spectrum per level is connected to its neighbour, and thus a pollen diagram is drawn that groups together all the evolution curves of the different species throughout the sequence, including a 'global curve that enables assessment, at each level, of the proportion of I arboreal cover (AP : Arboreal Pollen) to herbaceous stratum (NAP : Non- Arboreal Pollen). The pollen diagram, based on the, principle that the general curve of the proportions of arboreal pollen to non-arboreal pollen indicates the importance of forests compared to open spaces in European ldw and . middle-altitude areas, reveals a sequence of landscapes subjected to climate evolution. Palynology of the Quaternary in Temperate and Tropical Areas /299 II. QUATERNARY AND PREHISTORY IN TEMPERATE AREAS: FOR EXAMPLE IN EUROPA 1. Chronostratigraphy, Climatology and Palaeoenvironment of Fossil Man : Generality The term Quaternary was created in 1829 by the geologist Jules Desnoyers, and once the first book concerning the Geology of Quaternary was published by Henri Reboul in 1883, a Quatemary era divided into two periods was accepted : the Pleistocene, which represents the entire period called ”glacial”, as opposed to the Holocene, which refers to the sub-present or “post-glacial” period. This new era was not only characterized by climatic variations connected with glaciation phenomena, but also, in Europe, by the appearance and development of prehistoric man. Study of the Quaternary applies the experience attained by geology, archaeozoology, anthropology and palaeobotany. The relative datations of the Quaternary are based on. the results of these different sciences: their correlation with methods of absolute datation contributes to better determining the chronological and palaeoclimatic table, which can be paralleled to that of the great phases of Prehistory (Fig. 20.1). Therefore, using data resulting from the multidisciplinary studies carried out in the geological, palaeontological or archaeological Quaternary deposits, we,’can try to reconstruct the environment of the sites that have, or not, kept man’s traces and to lay out our ancestors’ landscape during their walk through Europe for nearly two million years. As plant species are pdcularly sensitive to temperature variations and hygrometric changes, it is certainly palaeobotany that can most accurately verify climatic evolution, to which the landscapes, reconstructed out of spores, pollen grains, various macroremains (grains, fruit, bark, coals ...) and prints, are subjected. The Pleistocene is divided into three major parts, according to its climatic history, which has been reconstructed mainly in the light of palynological data. Indeed, we can observe that the chronological terminology of the different periods of the Quaternary bears most of the time on the denomination of eponymous geological or archaeological sites, which are real stratdgpes that have yielded pollen spectra of plant associations corresponding to precise climatic phases. Let us add rhat these major climatic phases have been defined by palynologists, as a function of the plant covers already established: - the “Glacials”, which are periods sufficiently long and cold for the whole forest vegetation to disappear and be replaced by a stratum o€ steppe Herbaceae; - the “Interglacials”, which are not humid climatic periods contributing to completely restoring arboreal vegetation; * I 300 / Current Concepts in Pollen-Spore and Biopollution Research

5 6 7 8

9 10 li I2 l3 I4 l5 16 17 18

E"II0NIF.N

Fig. 20.1 : Quaternary and Prehistory in Europa (after J. Renault-Miskovsky). - the "Stadials",which represent the cold periods of the glacial phases, interrupted by "interstadials", which are minor climatic oscillations that entail partial and botanically incomplete takeover of steppe by forest.

2. Change Over Pliocene-lower, Pleistocene and Lower Pleistocene The climatic history of the early Quaternary is indissociable from that one of the upper Pliocene, which was determined in Holland by Zagwijn (1963, 1974), as pollen analysis of the-clay deposits in Reuver and Tegelen has enabled three climatic phases to be distinguished: the Reuverian, the pre-Tiglian and the Tiglian (Fig. 20.2). Such a history of plants in the late Tertiary was also reconstructed in Italy through pollen analysis of Leffe clays (Lona, 1950).

Y Palynology ofthe Quaternary in Temperate and Tropical Areas/301 In France, the latest palynological works on this climatic tuming- point are concerned with the Mediterranean Basin (Suc, 1980), Normandy (Clet-Pellerin, 1983), the Massif Central (Brun, 1971; Ablin, 1985) and the Bresse region (Farjanel, 1985). A correlation between the different climatic events that affected the Mediterranean Plio-Pleistocene from southern France to southern Italy has just been determined by Combourieu-Nebout (1987).

3. The Lower Pleistocene The Lower Pleistocene, which represents a still unsteady climatic period, from the late Pliocene to the progressive settling of the first glacial colds, has lasted from 1,800,000 to 700,000 years. The climate, at first temperate-hot and humid (north-European Tiglian), degraded during the first major climatic pejoration, materialized by an advance of the alpine moraines of the “GÜd$” In Europe, the thermophil flora took refuge in sheltered areas, but tQe slightest improvements in climate enabled new forests to develop out of small surviving clusters of vegetation_ Man was already present in Europe. He was a robust hunter who used archaic tools happed from pebbles and tracked massive animals within a vegetal scenery which was to evolve from luxuriant forest to arid steppe. The earliest climaw degradation to have been pollinically individualized (by Zagwijn Tin Holland) followed the Tiglian; ‘-and can be divided into three phases, accessively cold (Eburonian), warm (Waalian) and cold (Menapian). The Menapian was still followed’by three episodes,

individualized by the pollen spectra from Bave1 [Zagwijn and DeJong, I 19841, hence the name of Bavelian given to this complex fmthe late lower Pleistocene, whose forest phase, called “Interglacial”, was contemporaneous with the magnetic episode of Jaramillo, and which preceded tlie major interglacial (Cromerian), serving as a basis for the Middle Pleistocene, at the palaeomagnetic li& Brunhes-Matuyama (0.73 MA) (Fig. 20.2). For south-eastern France, the pollen sequence derived from studying the archaeological filling of the Vallonnet cave testifies to the cold and especially the dryness contemporaneous with the first hominids’ settlement on the shoreline, and to the following climatic improvement, which was synchronous with the palaeomagnetic event of Jaramillo, and was measured at this level of filling. At t5.e top. of the diagram, palynology can record, in a stalagmitic floor, part-of the following interglacial warming, in the late Lower Pleistocene and in’the early Middle Pleistocene (Renault- Miskovsky and Girard, 1978 and 1988) (Fig. 20.3). w E O C LI MATIQ U Roppel de lo Chronologie N CLIMAT VCGCTATI ON Alpine \ Essois de corr4lbtion .------

chaud développement des forêts à CROMEKIEN et GUNZ-MINDEI éléments thermophiles 52 humide 8 'Dm tempéré chaud BAVELIEN reprise des forêts et humide ------chute des arbres et développement MENAPIEN froid GUNZ ------des steppes ------GUNZ tempéré chaud WAALIEN reprise des taxons thermophiles DONAU-GUNZ et humide ------chute des arbres : DONAU EBURONIEN froid . régression des élemenes tertiaires et de certains taxons quaternaires thermophiles

tempéré chaud flore mixte à CIGments TIGLIEN . et DONAU-GUN2 BIBER-DONA1 humide tertiaires et quaternaires

chute des arbres : extinction de certaines Taxodiacées, PRETIGLIEN froid 'DONAU BIBER disparition ou régression d'autres genres thermophiles flore tertiaire associée à des REUVERIEN haud et humidr JIBER-DONAU PRE-BIBER taxons persistant au Quaternaire BIBER

Fig. 20.2 : Plio-Quaternary boundary in Europa (after Renault-Miskovsky, Petzold, 1989-1992). ovshy h M Girard) I HYPOTHESES D'EVALUATIOKCLIMI FlPRh

.Présence de taxons méditerranéens

et de

*Pr€ssence de taons médlterranéens et dee

0 Pr&ence de rare5 taxons médlterranéens

IEGETATlON DE TWE "QUE

Platanes Rksence de Ptwcarvi Prkence de &&

Fig. 20.3 : Palynological data of Vallonnet Cave : vegetation, climate, chronology and palaeobotany(after Renault-Miskovskyand Girard, 1978-1988). 304 / Current Concepts in Pollen-Spore and Biopollution Research 4. The Middle Pleistocene The Middle Pleistocene, as a matter of fact, began ca 700,000 years; it .was a warming period which enabled the forests destroyed by the preceding glaciation to settle down again this interglacial period, lasting about 50,000 years, is little known, and was mainly revealed through pollen analysis of north-European continental peat deposits; as the first pollen sequence related to the English site of Cromer, the term “Cromerian”is often preferred to the alpine denomination “Giinz-Mindel” (Fig. 20.2). From ca 650,000, the Elster glaciation developed during nearly 350,000 years, it carried the north European ice cap as far as northern Germany and entailed moraine thrust of mindelian alpine glaciers. This cooling was responsible for the disappearance of the last villafranchian animals, replaced by Lhe first elements of a cold fauna, but palynological malyses never recorded a complete destruction of re- gional plants, whether in central Europe or on the Mediterranean coast. At the end of this ‘“indelian” period, pollen spectra indicate the next warming of the Mindel-Riss Interglacial, which was characterized by temperatures that permitted not only reinstallment of big forests, but also the growth of very thermophil taxons. The glaciation following the end of the middle Pleistocene, which probably lasted 100,000 years, was the (alpine) Riss or the (Germanic) Saale. Cold periods seemed to be more marked in it than during the preceding glaciation, but its maximum stages were interrupted by warming Interstadials. Pollen diagrams reveal developing Birches, which might even give way to herbaceous steppes in plains, whereas Coniferae from the lower Alps stretched dum almost to the Provence shoreline. Meanwhile, man had settled in Europe. He would control fire and keep up hearths : thus the charcoal found in archaeological sites, when sthdied, ’fit quite well with the fossil pollen. This new inhabitant would fashion bifaces\ called “acheulean”and set up elaborate camps by building waterproof huts. He would venture into European northyn areas, mainly during warm interglacial periods, while easily taking refuge in southern areas during glacial periods. Hence the abundance of prehistoric sites now inventorised on the Mediterranean coast and hinterland, amoiíg which it seems worth mentioning the Caune of Arago in the Corbiéres, the open-air site of Terraknata and the Lazaret cave in Nick, which marked out our ancestors way between about 700,000 and 1\50,000 years and which are the subject of numerous pollinic studies.

5. The Upper Pleistocene The upper Pleistocene began with a strong warming period, which provoked melting of ice caps and retreating‘of alpine glaciers. This interglacial period, which took ,place between 130,000 and 100,000 years, is named F+s-Würm or Eemian, afteLthe river Eem and the marine!deposit due to transaression at the mouth of this river: for EuroDe, it was characterized Palynology of the Quaternary in Temperate and Tropical Areas /305

by a complete re-settling of arboreal plants, which were initially revealed , through pollen analyses of the more or less peaty lacustrine deposits overlying the rissian moraine, at Averbergen, south of Hamburg (northern Germany) (Selle, 1957). A recent palynological study of the Vosgian peat bog of the Grande Pile shows in fact, during this period, several warming phases, the latest of which may have insensibly intermingled with the "start" of the following Würmian sequence (Woillard, 1979). The diagram (Fig. 20.4) reveals different climatic episodes most often expressed in local names. - Linexert materialized the end the next to last Vosgian glaciation, with a low arboreal cover on behalf open spaces filled with Graminae and steppe Herbaceae. - Three forest phases followed: L.niian, St-Germain I and St-Germain II, interrupted by arboreal episodes with Birches and Junipers: Melisey I and Melisey II.

I Eemian, the first phase, involving not only development of the mixed Oak grove fringed with Hazels, but also representation of the Yew, Box and Hombeam, was probably the warmest; it is dated from 125,000 years BP, and corresponded certainly to the maximum temperature of

the interglacial period. B Both the following, in which the Birch and Pine were abundant, were no doubt more temperate or cooler and presaged a return to the more or less severe climatic conditions individualized in the next part of the diagram: it seems they can be superimposed on the first two temperate oscillations of the folrowing Würmian gIaciation. Actually, the climate slowly degraded again, and a new glacial period (Würm or Weichsel) settled, first cold and humid for around 20,000 years, then cold and dry. The arboreal landscapes that had been maintained by the help of a few major temperate oscillations, three of which were identified in northern Europe through pollen analyses (Amersfoot, Brorüp and Odderade) (Zagwijn, 1961),were replaced grassland or steppes. The stratigraphies, drawn up in south-western and south-eastenp France, are based on sedimentological and archaeological criteria. The4 early Würmian was a cold period, at first humid (Würmian I), then (Würmian II), which corresponded to fillings that have yielded + mousterian industry and remains of Neanderthalian fossil hominids: as for the recent Würmian, it embraced all upper Palaeolithic sequences. Towards 40,000 years, at the end of the early Würmim, the cold prevailed throughout Europe. , The scission between the early and the later Würmian, around 35,000 years ago was due to a change iq civilization rather than to a geologicdl event. Nevertheless, it seems that in some stratigpaphies, the changeoyer between the two WÜrmian periods is marked by several oscillations thdt! 306 / Current Concepts in Pollen-Spore and Biopollution Research N O LOG PA E CLIMATS CHRONOLOGIE LY I 20 40 60 80°/o 123459

n 3 ,ln Z-

c -0, > ZEO

3- 35 .-0, L 2 .-c

Herbades

Fig. 20.4 : Palynological diagram of la Grande Pile (after Woillard, 19791, Palynology of the Quaternary in Temperate and Tropical Areas /307 are recorded either by soil formation or by pollens from arboreal covers (c$ Hengelo, les Cottés). But in the early upper Palaeolithic, and for about 15,000 years, the climate was hard; the cold was interrupted by three slight climatic fluctuations only, towards 29,000 years B.C. (Arcy), 26,000 B.C. (Kesselt)and 21,500 B.C. (Tursac),which gave rise to sporadic plant growth. From 18,000 years B.C. onwards, the end of the latest glaciation was regularly interrupted by temperate oscillations which grew more frequent and definite until the post-Glacial [Leroi-Gourhan and Renault-Miskovsky, 1977; Renault-Miskovsky and Leroi-Gourhan, 1981). It is worth stressing that they were mainly revealed by the palynology of archaeological deposits, which was often disputed, notably by palynologists of peat bogs who did not notice all such oscillations. The main temperate oscillation took place at the time of magdalenians' occupation of the Lascaux cave (Magdalenian II); it is well two , dated by I4C isotopic measurements from 15,240 B.C. and 14,150 j B.C., and can be found in many other pollen analyses of caves and peatbogs. In general, archaeo-pollinic spectra supply chronological details that prove sometimes more precise than typological studies alone; being often correlated with absolute datations, they indicate certain climatic and cultural contemporaneities and thus demonstrate that changes in industry are not specifically connected with an even less subjected to changes in climate (Fig. 20.5). After the warming recorded in the deposits of the Lascaux cave around 15,000 ELC. down to the late Pleistocene-climatic dynamics accelerated. Three successive temperate pulsations (pre-Bölling, Bölling and Alleröd), occumngl at near intervals, no doubt improved the comfort of the latest Palaeolithic men, Magdalenian and Azilian, who suffered anyway the last three sharp intermediary pejorations of the Dryas (Dryas I, II, and III); this period of junction was that of the Lateglacial. It presaged permanent post-glacial warming which grew stronger from 8200 years B.C. onwards.

6. The Psst-Glacial Consequently, the Post-GZaciaZ succeeded the Dryas III, which was the last cold wave at the end of the WürmiarI glaciation, towards 8200 years B.C. Geological data are numerous, but initially it was the poll'en chronology established by two Scandinavian botanists, Blytt and Semander 11876) , which permitted five typical climatic phases to be distinguished for this period in northern Europe; today this chronology 4s extended to the entire continent. the pre-Boreal (from 10,000 to 9000 BP), which was a phase of transition between the last lateglacial colds and the first post- glacial warming; it was characterized by development of the Rne, followed modestly by that of the Hazel, Oak and Elm; 308- / Current Concepts in Pollen-Spore and Biopollution Research DATES STADES COURBE CLIMATIOUE C 14 INDUSTRIES PALEOLITHIQljES B.F!

-10000

- 12000

- 14000

-16000

-18000

.20000

-22000

-24000

- 26000

-28000

' + ' - 30000 e +y&------' Arcy

-32000

-34000

Lea CottCo -36000 II1 - 38000 Hengelo I Fig. 20.5 : Palynology, prehistory and datations of th5 Upper-Palaeolithicin Europa (after Leroi-Gourhan, in Renault-Miskovsky,1991). of Palynology the Quaternary in Temperate and Tropical Areas /309 - the Boreal (from 9000 to 8000 BP), where warming strengthened, with moderate dryness helping the expansion of Hazels and all leafy trees: - the Atlantic [from 8000 to 5000 BPI, whose humid temperate climate enabled an expansion of the mixed Oak grove (with Limes prevailing), particularly in the second half of the phase; - the sub-Boreal (from 5000 to 2500 BPI, cooler and dryer, which corresponded to a recession of the mixed Oak grove and an extension of the Beech, Fir and Picea at a high altitude: - the sub-Atlantic [from 2500 BP to modern times), whose individualisation is due to a large extension of the Beech and Hornbeam. The changeover between the late Pleistocene and the middle Holocene [from the temperate oscillation of the Alleröd, towards 9800 B.C., to the early Atlantic) was lived by Epipalaeolithic civilizations (Azilian and Mesolithic men), who were still predatory, but already productive: hunting, fishing, snail collecting, and gathering were everyday activities in their increasingly sedentary life, which was dependent on seasonal rhythm, but still fitting into a naturd iauclscape. From the end of the Boreal, however, in the middle A-tlantic period, early Neolithic men (from ca 6000 lo 3800 B.C.) made the first attempts at breeding and agriculture, which were to further develop during the middle and recent Neolithic (down to 2000 B.C.). These agricultural practices entailed not only sedentarisation, accumulation of producer’s goods involving herbs and crops, but also intensive deforestation, which is demonstrated in pollen diagrams by a drop in the percentage of forest plant pollen, together with an increase in the ratio of Herbaceae, especially ruderal species, and a sporadic appearance of pollen from cultivated plants such -as cereals and Leguminosae. Deforestation intensified proportionately as soils were worked out because of mass utilization of wood in the furnaces meant for metal melting, from the early Bronze Age, towards 1800 B.C. to the Téne in the recent Iron Age.

III. QUATERNARY AND PREHISTORY IN TROPICAL AREAS: FOR EXAMPLE IN SOUTH-EAST ASIA 1. Quaternary and Prehistory in Indonesia: Generality The main repercussion of the Quaternary in tropical areas is the lowering of the level of the sea and the formation of land bridges enabling fauna and hominids to colonize newly formed lands. However, modifications in climate, evidence of which may be brought to Light by the study and evolution of the fauna and flora, are also going to take place. As early as 1949, Bermmelen’s work established the, main basis of 31O / Current Concepts in Pollen-Spore and Biopollution Research geology in Indonesia. As for the field of prehistory, numerous studies have been carried out by Indonesian and foreign researchers amongst which are certain synthetic works such as that of Bellwood in 1985, The Quaternary of Indonesia, and more specifically of Java, begins in fact at the time of possible arrival of the first hominids (2 million years ago) at a period when, in central Java, the sea was receding, uncovering new territories towards the East. This retreat occurred thanks to the conjugated action of eustatic readjustments and of volcanic and tectonic activity. Indonesia is situated in the zone of interaction of four twtonic plates: Eurasia, Australia, Indian Ocean, Philippines and Pacific. If one excludes the western part of Java, the island is affected by a succession of ridges following an Eakt-West axis. Amongst these structures, three are of particular interest: - in the south, the volcanic reliefs of Miocene calcareous formations; - in the north, an anticlinal ridge, the formation of which dates from the Miocene and final Pliocene, i.e.,the hills of Kendeng; - between the two, the great depression of Bandung-Solo, occupied by Quaternary alluvia and large and still active volcanoes. Parallel to the forming of the oriental and central parts of Java, the quaternary glaciation of the high and middle latitudes have for more than two million years induced a repetitive lowering of the level of the sea, which has favoured formation of land bridges on the Sunda shelf. Serving as a link between the islands and the Asian continent, these land bridges allowed the passage towards Java of continental mammals and hominids who took over the habitats, newly emerged as a result of tectonic raises and eustatic lowerings.

2. Ancient Quaternary in Indonesia In Java, the study of mammal and 1iorr;nid sites and of the environment of the Pithecanthropus have led to greater knowledqe of the Quaternary. It is at Bumiayu (Fig. 20.6),which has noi sunendered my hominid fossils, that one finds fauna considered to be ancient-the fauna of Kali- Glagah. Following the marine series;-the synorogenic series of Kali-Glagah begins at around 2 million years. It is made up of a succession of sandy, conglomeratic or clayey fossil-bearing layers. P. lower horizon contains mainly Mastodons and Hippopotamm while higher in the stratigraphy appear Fklidés and Stegtbdons. In the marine sediments, one finds the retinue of the mangrove, indicating that the littoral is fairly close by. The mangrove takes great importance at the top of these layers at zpproximately 2.3 million years, thereby localizing the shore line and the recession of the sea, a phenomenon contemporary with the surrection of the nearby mountains. Palynology of the Quaternary in Temperate and Tropical Aread311

Java Sea

Indian Ocean 100km,

1.Situ Bayongbong 2 .Bumiayu 3.Simo (Onto) 4.ltaliuter 5.Gemolong 6.Sangiran 7.SOlO 8. Sambungpacan 9.Trini1, Ngandong 10.Pacitan ,

Fig. 20.6 : Sites of Java (after Semah).

1. Danau Padang 2. Mahakam delta 3. MT Jaya 4. MT Wilhelm

Fig. 20.7 : Sites of Indonesia (P.N.G.). In the first continental levels, the mangrove which is still represented albeit faintly, attests that the emersion occurs progressively and that of the first fossil-bearing levels of Kali-Glagah are very close to the sea. On the coast, behind the mangrove, a boggy and herbaceous vegetation develops. On higher land, the tropical rain forest dominates. During this humid period, the abundance of Podocarpus imbricatus, which subsequently decreases abrupily, indicates a temperature lower than of today. As for the higher manne levels, the vegetation of the hinterland presents a more open and less humid aspect linked perhaps in part to a volcanic phase. In the first continental levels, the mangrove regresses,

I 312 /Current Concepts in Pollen-Spore and Biopollution Research the swampy and littoral formations develop while in the background, a tropical rain forest grows ih frequent imbalance under the influence of volcanic activity. The passage of the Plio-Pleistocene can also be observed in the Solo depression where the Quaternary volcanoes have crushed the thick sedimentary series and brought about the formation of domes at .their base. The late Pleistocene represented by blue marine clays is known here as the formation of Kalibeng. It ends with coastal limestone and volcanic’breccias. It is followed by three classical formations of the ea-rly, mid and late Pleistocene, known in this region as Pucangan, Kabuh and Notopuro. They appear more or less completely in the domes of Sangiran, Onto and Gemolong as well as at KaliLiter situated on the southem flmk of the Kendeng (Fig. 20.6). At Gemolong, the Kalibeng formation is made up of yellow marine mark with Globigerines topped with blue clays which are still of marine origin that also ar^ found at Sangiran, Simo and Kaliuter until around 1.8 to 2 million years. During this period, transition or directly continental facie appear, namely, Bulunus or Corbicula limestone and deltaic clay and conglomeratic deposits. The first vertebrate fossils, cervidae, crocodile, rhinoceros, Stegodon and bovidae, have been found in these transition facies. The overjacent levels present severd fossil-bearing horizons for which the chronology is still not well established. The next levels of Pucangan are made up of black clays, paludal of Sangiran and blue clays of Kaliuter. The most archaic hominids of the dome of Sangiran, Pithecanthropus IV, Sangifan 31, come from the layers of Pucangan. I At the summit of the Pliocene, a swamp forest developed behind the mangrove forest, and a rain forest developed in the higher areas. At around 1.7 millnon years, the sea retires, the lagoon fills up and frequent volcanic eruptions repetitively destroy the vegetation. The mangrove tends to disappear, the drier climate favours the development of grassy expanses. It is following these eveots that the ancient hominids were able to colonize new territories. In the early Pleistocene, the mangrove appears only sporadically and the vegetation progressively becomes exclusively continental. It is a wit- ness to climatic oscillations and presents an altei-nation between the humid interglacial phase rain forest and a more open forest during drier periods. Later, in the dome of Sangiran, one finds at Gound 700,000 to 800,000 years, the series of fluviatile alluvia, of Tufa and Kabuh clays. The alluvia series are often marked on their base by a calcareous con- glomerate named “Grembank.These formations relinquished the great- est part of human remains : the Pithecanthrophs IV, VIL VIII, the classi-

. Palynology of the Quaternary in Temperate and Tropical Areas /313 cal type of Pithecanthropus in addition to the fossil of Trinil, for example, discovered by Eugene Dubois in 1891. These Homo ereclus bear numerous resemblances to the African and European Homo ereclus who are their contemporaries. The layers of Kabuh are covered over in uncomfortable strata by the breccias and lahars of Notopuro which are the result of very important volcanic eruptions. The sandy alluvia contain more fossils. The Late Pleistocene has surrendered evolved Pithecanthropus fossils, in particular at Ngandong where more than ten skulls were extracted in 1931 from alluvia of the Solo River. The skull of Sambungmacan discovered in 1973 may also be linked to this group. For the three large groups of Pithecanthropus, we know of only few tools. In 1935, R. von Koenigswald discovered a paleolithic industry in the South of Java at Pacitan (Fig. 20.l), the study of which was followed by G.J. Bartstra. This Patjitanian industry is not dated with precision and cannot be linked, a priori, with the Pithecanthropus. In the same way, the superior levels of the Sangiran dome have revealed an industry of flakes of small size which appears to be more recent- than the Pithecanthropus series. On the other hand, at Sambungmacan, a chopper and a touched-up flake, discovered in layers contemporary with the skull, are the work of the Pithecanthropus. Excavations begun in 1990 at Ngebung in the dome of Sangiran have allowed to bring to light, in mid-Pleistocene levels, an existing set of tools as well as a structure of occupation (excavation of the Quaternary and Prehistory Mission in Indonesia; Sémah, Djubiantono) .

3. Recent Quaternary in New-Guinea and in Indonesia For more recent periods, much information has been provided by cores carried out in peat bogs or swamps of Indonesia. For the most part, these cores are undertaken in altitude. They thereby allow to keep track of the shifting of the altitudinal zonation of the vegetation and to deduce climatic variation-warming up or cooling of the climate. Where cores have taken place in New Guinea (Hope & Peterson, 1975), one notes the retreat of the glaciers around 15,000 B.P., followed by a return after 13,000 and 11,000 B.P. A neo-glacial period appears in the first 2400 years. In New Guinea, research in palynology and archaeology has shown that man has been present in Highlands since 25,000 B.P. (Gorecki, 1986). Fires and degradation of the forest are also shown to have occurred as early as 26,000 B.P. in Irian Jaya. An important amount of anthropic perturbation of the vegetation begins at around 7000 B.P. (Haberle et aZ., 1991). Numerous examdes of the influence of man on the vegetation have 314 / Current Concepts in Pollen-Spore and Biopollution Research . been brought to light by traces of horticulture at 9000 B.P. in the High- lands (Golson, 1982), by fires and destruction of the forest at 10,500 and

J by use of caves as hunting shelters at 5000 B.P. at 3600 metres of altitude on the Kemabu plateau of Mount Jaya (Hope & Peterson, 1976) (Fig. 20.7). In Sumatra, a core carried out in Lake Padang at 950 metres of altjtude (Morley, 1982), shows a change of vegetation between 10,000 and 8600 B.P., representing an improvement of climatic conditions. At around 4000 B.P., the trees of the primary forest give way ta Trema and Macaranga-witnesses of the reconquest of the soil &er an anthropic disturbance. From several pollen studies in altitude of the Highlands, Flenley has reconstituted the history of the vegetation as well as the shifting of the higher limits of the forest over 30,000 years (Fig. 20.8). One notes a climatic improvement at around 28,000 B.P. as well as starting around 12,000 B.P. In western Java, a core carried out at Situ Bayongbong, tracking in particular the evolution of Podocarpus imbricatus, brought evidence of a colder period between 17,000 and 12,400 B.P. followed by a clear climatic improvement at around 7900 B.P. (Stuijts et al., 1988). Studies have also been led in low altitude. A core over 600 metres deep was undertaken in the delta of the Mahakam located in the East of Kalimantan (Caratini and Tissot, 1988). The base of this core was dated to the late Pliocene. The authors brought ' to light the permanent nature of the deltaic conditions. The local Vegetation, completely covering that of the hinterland, has remained fairly constant. An indication of the climatic variations has been given by the pollen of Poaceae (Gramineae) the abundance of which at several periods and particularly around 18,000 B.P. has been interpreted as a phase of development of the savanna, linked to cooling of the climate. These phases correspond to low marine levels accompanied by can increase of detrital deposits. In central Java, on the Rawa Pening site, two cores carried out respectively at 10 metres of depth (4000 R.P.) and at 40 metres (16,000 B.P.) have shown variations in the vegetation of climatic but also of anthropic origin (Sémah et al., 1992). One observes the passage from a cool and relatively dry period to a warmer and humid period at around 12,000 B.P. The first clearings of land are shown in the pollinic diagrams to Dccur around 1500 B.P. and the beginning of placing the region under cultivation at around 400 B.P. (Fig. 20.9). In this part of the world, the data still remain scattered but it is &arting to show a certain homogeneity and confirms that the impact Of the great ,climatic changes characteristic of high and middle latitudes are I I' of Palynology the Quaternary in Temperate and Tropical Areas /315

.. . 4000m

3000m

ALTITUDE

2OOOn

3doOO 20600 ld000

Fig. 20.8 : Altitudinal changes of upper-forest limit in Papua, New Guinea (after Flenley, 1979). A. P. SPORA

of Fig. 20;9 : Ambarawa, evidence older grubbing periods (after Semah et ab, 1992). 316 / Current Concepts in Pollen-Spore and Biopollution Msearch clearly noticeable in tropic+ areas. Here also, prehistoric man has encountered more or less hostile surroundings, chosen his sites 'for habitat and, when conditions were favourable, reshaped his environment.

REFERENCES ABLlN, D., 1985. AnalGse pollinique de dépôts lacustres de Ceyssac (Plio-Pléistocène du Velay. Massif Central français); Flore, Végétation et Climat, Thèse de Doctorat de 3è cycle (mention : Géologie du Quaternaire). Museum National d'Historie Naturelle et Université Pierre etMarie Curie, Paris VI, 121 p. BELLWOOD, P. 1985. Prehistory of the Indo-&IalaysianArchepelago. Academic Press, éditeur, Sydney.. BERMMELEN, R.W. van. 1949. 2dme édition en 19701 :The geology of Indonesia and adjacent archipelagoes. Martinus Nijhoff éditeur, Den haag. BLYTT, A. 1876. Essay on the immigration of the Norwegian flora during the alternating rainy and dry periods. Christiana. BRUN, A. 1971. Les applications de la palynologie à l'étude géodynamique du massifvolcanique du Mont-Dore. Thèse de Doctorat d'Etat-es Sciences Naturelles. Faculté des Sciences de Paris A.O. 5422, 244 p., 31 fig., tabl. et cartes h.t. CARATINI, C. and TISSOT, C. 1988. Palaeogeographical evolution, of the Mahakani delta in Kalimantan. Indonesia during the Quatemary and the late Pliocene. Review of Palaeobotany and Palynology, 55 : 2 17-228. Elsevier Science Publishers B.V.; Amsterdam. CLET-PELLERIN, M. 1983. Le Plio-Pléistocène en Normandie. Apports de la klynologie. Thèse de Doctorat de 36 cycle (mention : Géologie). Université de Caen., 135 p. COMBOURIEU-NEBOUT, N. 1987. Les premiers cycles glaciaire-interlaciarie en région méditerranéenne d'après l'analyse palynologique de la série Plio-Pléistocène de Crotone (Italie méridionale). Thèse de Doctorat (spécialité. Géologie), Université des Sciences et . Techniques du languedoc, 161 p., 34 fig. FPJUANEL, G. 1984. La flore et le climat du Néogène.et du Pléistocène de Bresse (France) d'après.l'analyse ,polliniquezImplications chronostratigraphiques. Thèse, diplôme de Doctorat (mention : Paléontologie-Stratigraphie). Université de Dijon, 193 p. . GIRÁRD, M. 1975. Pkevements d'échanitillôns en &&es et stations de-terrain sec en vue de l'analyse pollinique, Bull. Soc. Prehist. Française, t. 72, CRSM, n"5, 158-160. GIRARD, M., and RENAULT-MISKOVSKY, J. 1969. Nouvelles techniques de préparation en palynologie appliqées a trois sédiments du Quatemaire final de l'Abri Comille (Istres, 4 , Bouches-du-Rhône). Bull. de l'A.F.E.Q., : 275-284. GOLSON, J. 1982. The ipomoeqn revolution revisite,d : society and the sweet potato in the dpper Waghi Valley. Inequalitg in the New Guinea Highlands societies. (ed. by A. Strathem) 7 109-136, Cambridge University Press. GORECKI, P. 1986. Human occupation and agricultural development in' the Papua New Gdinea Highlands. Mountain Research and Development 6 : 159-166. 1 HABERLE, S.G., HOPE, G.S. and DEFRETES, Y. 1991. Environmental change in the Baliem Valley, montane Irian Jaya, Republic of Indonesia. Journal ofBiogeography 18 : 95-40. HOPE, G.S.and PETERSON, J.A. 1976. Palaeoenvironments. The equatorial glaciers of New Guinea. (ed. by G.S. Hope et al.), 173-207.' Balkema, Rotterdam. LEROI-GOURHAN Arl. and RENAULT-MISKOVSKY, J. 1977. La Palynologie appiiquée a ' l'Archéologie. Méthodes, limites et résultats. In Approche écologique de l'Homme fossile. ' Supp. au Bull. de L'A.F.E.Q., n"47, 35-49, 6 fig., nbx. diagr. I LONA, F. 1950, Contributi alla storia della vegetazione e del clima nella sal padana. Analisi pollinica del Giacimento villafranchiano ,di Leffe (Bergamo). Atti della Soc. Ithl. di Sc. , Natur. LXXXI :123-180. MORL,,Y, R.J. 1982. A palaeological interpretation of a 10,000 year pollp' record from Danau Padang, Central Sumatra, Indonesia. Journal ofBiogeography 9 : 151-190. of Palynology the Quaternary in Temperate and Tropical Areas /317 RENAULT-MISKOVSKY, J. 199 1. L'environnement au temps de la Préhistorie. Méthodes et modeles. Masson, 2éme édition, Paris, 200 p., 68 p. RENAULT-MISKOVSKY, J., and GIRARD, M. 1978. Analyse polinique du remplissage pléistocéne inférieur et moyen de la grotte du Vallonnet (Roquebrune-Cap-Martin, Alpes- Maritimes). Géologie méditerranéenne, t. V, n"4, 385-402, 6 fig.. 1diagr., 4 table., 4 pl. phot. RENAULT-MISKOVSKY, J., and GIRARD, M. 1988. Palynologie du remplissage de la grotte du Vallonnet [Roquebrune-Cap-Martin,Alpes-Maritimes). Nouvelles données chronologiques et paléoclimatiques. L'Anthropologie, 92, n"2, 437-448. RENAULT-MISKOVSKY, J., and LEROI-GOURHAN Arl. 1981, Palynologie et Archéologie. .Nouveaux résultats du Paléolithique supérieur au Mésolithique. Bull. de L'A.F.E.Q. 3-4 : 121-128. SELLE, W, 1957. Das letzte lnteglazial in Niedersachsen. Ber. Natur. Ges. Hannover, 77-89. SEMAH, A.M., SEMAH, F., GUILLOT, C., DJUBIANTONO, T. and FOURNIER, M. 1992. Etude de la sedimentation pollinique durant les quatre derniers millénaires dans le bassin d'Ambarawa (Java Central, Indonésie). Mise en évidence de premiers défrichements. C.A. Acad. Sci. Paris, t. 315, serie II, 903-908. STUIJTS, I., NEWSOME, J.C. and FLENLEY, J.R. 1988. Evidence for Late Quaternary vegetational change in the Sumatran and Javan Highlands, Review of Palaeobotany and Palynology 55 : 207-216. SUC, J.P. 1980. Contribution à la connaissance du Pliocène et du Pléistocéne inférieur des régions méditerranéennes d'Europe occidentale par l'analyse palynologique des dépôts du Languedoc-Roussillon (Sud de la France) et de la Catalogne (Nord-Est de l'Espagne). Thése de Doctorat &Etat [mention : Sciences). Université des Sciences et Techniques du Languedoc, 198 p. WOILLARD, G. 1979. The last interglacial-glacial at Grande Pile in northeastem France. Bull. Soc. belge de Géologie 88, fasc. 1, 51-69, Bruxelles. t. *t ZAGWIJN, W.H. 1961. Vegetation, climate and radio-carbon datings in the Late Pleistocene in the Netherlands, Part I : Eemian and Early Wechselian. Meded. Géol. Sticht, N.S. 14, 15-45,Maastricht. ZAGWIJN, W.H. 1963. Pleistocene stratigraphy in the Netherlands based on changes in vegetation and climate. Verh. K.N.G.M.G. Géol., sér. 21, 2 : 173-196. ZAGWIJN, W.H. 1974. The plio-Pleistocene boundary in Western and Southern Europe. Boreas 3 : 75-97. ZAGWIJN, W.H. and DE JONG, J. 1984. Die interglaziale von Bave1 und Leerdam und ihre stratigraphische Stellung in Niederlkdischen Früh-pleistozän. Meded. Rijkcs. Goel. Dienst. 37, 3 : 155-169. I

CONTENTS OF THE VOLUME,

Foreword / v 12. The Pollen Morphology of Bauhinia pottsii Preface / vi¡ G. Don (Leguminosae: Caesalpiniodeae) Editors' Note / ix I.K. Ferguson and Hannah Banks 1207 Messages 11 13. Xerocline and Hygrocline Phases in Brief Biographical Profile I35 Oberharz Mires During the Se!tling Time Some Precious Pictures I39 (Pollen Zone X) Monika Ulmann 1215 Contributors of the Felicitation Volume I49 14. Stain Reversal in Pollen Exines John R. List of selected publications of Prof. S. Rowley and Joanne S. Rowley I 223 Chanda (upto October 1994) / 55 15. Airspora and Respiratory Symptoms, Synopses of the Ph.D Theses made under the Adelaide, South Australia W.E. Boyd, supervision of Prof. S. Chanda 169 M.B. Howland, A.E. Galeand K.L. List of Ph.D, Theses submitted under Prof. S. Thomson1233 Chanda as Co-SupervisorlAdvisor/ 91 16. The New Dimensions of Palynological Names of Scholars, who obtained Ph.D. Sciences P.K.K. Nair 1251 degree exclusively under the supervision 17. The Original Position of the Generative of Prof. S. Chanda I95 Cell in Angiosperms with Heteropolar Pollen Kim-Lang Huynh I259 1. Oak Pollen Influx in the Sangre de Cristo 18. Responses of Higher Plants to Air Range, New Mexico, USA Stephen A. Pollution in the Metropolis of Calcutta : Hall I99 Light & Electron Microscopy on Leaf and 2. Some Abnormal Pollen Grains of Two Epidermal Architecture S.C. Datta and Indian Firs, Abies pindrowand A. S. Sinha-Ray / 271 spectabilis and their Phylogenetic 19. Protein Patterns of Pollen and Ovules of Significance P.N. Mehra / 105 Cucurbits and their Taxonomic Signifi- 3. Pollen Grains and other Pauci-Micronic cance M.K. Pasha and S.P. Sen I287 Particles as Airborne Allergen Carriers F. 20. Palynology of the Quaternary in Temper- Th. M. Spieksma and A.H. Nikkels I 113 ate and Tropical Areas : 4. Problems in studying Pollen Dispersal Chronostratigraphy, Palaeoclimatology into and within Scots Pine Seed Orchards and Vegetal Palaeoenvironmentof Fossil A. Rantio-Lehtimtiki and P. Pulkkinen I Man Josette Renault-Miskovskyand 121 Anne-Marie Semah 1297 5. Aerobiology in Southern Africa Ann 21. Airborne and Allergenic Tree Pollen of the Cadman 1131 Texas Gulf Coast (USA) W.H. Lewis, 6. Yearly fluctuations in atmospheric A.B. Dixit, H.J. Wedner and W.A. Ragweed Pollen Crozier 1319 Concentrations in Tulsa, Oklahoma 22. A Staining Method for Detection of Estelle Levetin I141 Protein in Pollen and other Particles in the 7. Sampling Airborne Biopollutants J. Lacey Air Ruth M. Leuschner and G. Boehm I and H.A. McCartney I149 337 8. Aspergillus fumigatus-Immunological 23. Pollen Morphology of the Indian Palm ' and Clinical Problems A.W. Frankland I Genus Bentinckia (Iguanurinae : 165 Areceae) Madeline M. Harley I343 , 9. Azadirachtin A Affects GAB+ Pools in 24. Consequence of Earth Expansion on Life Neuroendocrine Organs of Locusts S. : A Hypothesis Gerhard O.W. Kremp I Banerjee, WfOberthur and H. Rembold 353 I173 25. Aeroallergen Surveillance : A Critical 10. Air Dispersal of Pollen in the Neotropics: Review of Sampling Techniques M.K. Its Relevance to Allergic Disease I. Agarwal and Shashi Jhamb I357 Hurtado, L. Cariani and H. Ramos / 179 Subject lndex I 379 11. Epidemiology and Plant Disease -Authorlndex I 385 Forecasting R.P. Purkayastha, P.K. Ghosh and S. Bera I 18Z Our other Publications ENVIRONMENT AND AEROBIOLOGY Edited by Ashok K Jain Foreword by Prof. Sunirmal Chanda

The book presents a compendiium of knowledge on general environment and aerobiology having 29 research articles classified under five sections, i.e., General Aerobiology havingfive papers, Aeropalynology having four papers, Aeromycology having seven papers, Epidemiology and Forecast having six papers and Allergy and Immunology having seven papers. These articles have been contributed by over 50. research workers. It is a! book which will act as an updated reference for the workers of various fields of Aerobiology, Allergology, Microbiology, Plant Pathology and other allied disciplines.

PEST MANAGEMENT IN VEGETABLES K P Srivastavd & Dhamo K Butani Foreword by Prof. M S Swaminathan

r \ Published by RESEARCH PERIODICALS & BOOK PUBLISHING HOUSE P.O. Box 720728, Houston, Texas 77272 USA Tele. : (713) 779-2999, Fax : (713) 779-2992 Tollfree (800).521-0061,E mail : [email protected]. c L Oflprintffom

CURRENT CONCEPTS IN POLLEN-SPORE

AND .I BIOPOLLUTION RESEARCH (Professor Sunirmal Chanda 60th Birth Anniversary Felicitation Volume,)

Editors NARENDRA MOHAN DUTTA SWAT1 GUFTA-BHA'ITACHARYA SUDHENDU MANDAL - KASHINATH BHA'ACHARYA

Foreword by A.K. SHARMA Centre of Advanced Study (Ce¿¿and Chromosome Research) Department of Botany University of Calcutta, Calcutta

LI 1998 CI xii+395 pages ISBN 0-9656038-1-4 O Size 25cm

RESE&CH PE~ODICALS8r BOOK PUBLISHING HOUSE USA UK INDIA . TAIWAN , ABOUT THE VOLUME

Palynology in a wider sense is the study of various microscopic tiny living of fossil entities including pollen, spores, algal frag- ments, insect scales, mites, etc. This subject complex is now considered as an important combination of applied and fun- damental multidisciplinary approach encompassing the wide spectrum of life, earth and biomedical scien/ces including dis- ' cipline's like Taxonomy, Pathology, Paleobotany, Palaeontology, Medicine, Cytology, Genetics, etc. Due to exponential growth of knowledge in various fields of Palynology, most of the available reference books either lack information about some of the most recent developments or else overemphasize on Aerobiology thus ignoring classical Pa- lynology which has been proved to be equally important all the world over. The present volume, however, lays balanced emphass on classical and applied Palynology thus covering various facets of the subject. The books is divided into 25 chapters dealing with all as- pects of Palynology highlighting current researches. The grater part of the book is devoted to classical aspect of palynological research embracing pollen morphology in relation to plant taxonomy dealing with critical and disputed taxonomic problems. The mysteries have been unravelled using latest techniques of SEM and TEM contributed by internationally reputed scien- tists. The other major important aspect, i.e. the topic dealing with the biomedical approach, e.g. pollen-spore allergy, is known to be one of the leading health problems. In addition there are contributions made by leading experts in the fields of biochemical and immunological disciplines. The tremendous development in the field of pollen analysis in relation to paleaoecology, phytogeography and vegetational history have also been given due recognition. The book is well illustrated, possesses comprehensive accounts for researches and schol- qrs of palynology, aerobiology and allied subjects. Above all, this volume has got its own character because it 'is dedicated in the name of such a person who devoted his - whole life for cultivation of the subject-complex of Palynology which resulted into the establishment of leading school of pa- ' lynological and aerobiological research in India acclaimed by global palynological community.