Contemporary pollen assemblages from the Vosges (France)

Review of Palaeobotany and Palynology, 33 (1981): 183--313

183

Elsevier Scientific Publishing Company, Amsterdam - - P r i n t e d in The Netherlands

CONTEMPORARY (FRANCE)

POLLEN ASSEMBLAGES FROM THE VOSGES

C.R. JANSSEN

Laboratory of Palaeobotany and Palynology, Utrecht (The Netherlands) (Received November 3, 1980; revised version accepted February 12, 1981)

ABSTRACT Janssen, C.R., 1981. Contemporary pollen assemblages from the Vosges (France). Rev. Palaeobot. Palynol., 33: 183--313. This paper comprises a comparative study of the recent relative regional pollen deposition and the forest vegetation in a 70 km long transect across the Vosges. The forest vegetation was mapped with the aid of the qualitative data from individual forest lots, available from the "Office National des For~ts". Eight landscape regions were established and defined on the basis of forest vegetation, geology, topography and precipitation. Pollen was analysed from surface samples and the ranges, frequencies and median values of the individual pollen types were calculated for each of the landscape regions and compared with the distribution of forest vegetation types and the relative abundancies of A bies alba, Castanea sativa, Fagus sylvatica, Larix decidua, Picea abies, Pinus sylvestris, Pseudotsuga menziesii, and Quercus sp. Next to tree pollen types, all the pollen types of herbs and shrubs were fully accounted for in order to establish the relation between the pollen values and the regional and local occurrence of the relevant taxa. In general the major vegetation sequence of the forest dominants, from west to east across the mountains, viz. Quercus--Pinus, Fagus--Abies--Picea, Fagus and again Fagus-Abies--Pieea and Quercus--Pinus is well reflected in the pollen deposition. It is concluded that the landscape regions can be characterized by their regional pollen assemblages, mainly by the values of the tree pollen types; to a lesser extent by those of herbs and shrubs. Herb pollen types that characterize landscape regions include Genista, Pteridium and Dryopteris type. The degree of aforestation and cultivation is reflected in the pollen values of Anthemis type, Chenopodiaceae, Plantago lanceolata, Cerealia and by Corylus, Betula and Alnus. Three additional landscape regions were recognized on the basis of vegetation only. SOMMAIRE Cette ~tude a trait ~ la relation entre la pluie pollinique et la v~g~tation actuelle dans un transect, dirig~ ouest-est d ' u n e longueur de 70 k m et traversant la cr~te des Vosges. Les types forestiers sont cartographi~ en fonction des donn~es quantitatives des essences principales se trouvant sur des parcelles, disponible ~ l'office national des for~ts. Huit r~gions de paysage ont ~t~ ~tablies et d~finies par des critbres relevant de la v~g4tation foresti~re, de la g~ologie, de la topographie et de la pr4cipitation. L'analyse pollinique s'est effectu~e sur des ~chantillons de surface. Pour chaque type de pollen et pour chaque r~gion, la variation dans les pourcentages, la fr~quence et la 0034--6667/81/0000--0000/$2.50 © 1981 Elsevier Scientific Publishing Company

184 value m 6 d i a n e o n t 6t4 calcul6es e t c o m p a r 6 e s avec la d i s t r i b u t i o n actuelle des t y p e s de f3r~ts et avec l ' a b o n d a n c e relative des esp~ces s u i v a n t e s : Castanea sativa, Fagus sylvatica, Larix decidua, Picea abies, Pinus sylvestris, Pseudotsuga menziesii et Quercus sp. Les t y p e s d e p o l l e n s o n t trait6s 6 g a l e m e n t p a r t y p e ainsi q u e p a r r6gion de paysage. C'est de telle m a n i 6 r e q u e les r61ations e n t r e les p o u r c e n t a g e s de p o l l e n et la d i s t r i b u t i o n r6gionale et locale des essences n o n - a r b o r 6 e n n e s o n t 6t6 d6termin6s. E n general, la s e q u e n c e p r i n c i p a l e d e la v 6 g 6 t a t i o n t r a v e r s a n t la cr~te des Vosges, c ' e s t ~ dire Quercus--Pinus, Fagus--A bies--Picea, Fagus et puis e n c o r e Fagus--A bies--Picea et Quercus--Pinus d a n s l'est du t r a n s e c t est b i e n refl6t~ d a n s la pluie pollinique. O n p e u t c o n s t a t e r qu'il est possible de caract6riser les r6gions de paysages s u r t o u t p a r la pluie p o l l i n i q u e des essences foresti~res et m o i n s p a r celle des essences n o n - a r b o r 6 e n n e s , p a r e x a m p l e p a r Genista, Pteridium et Dryopteris. La densit~ de la for~t et le degr6 de la c u l t u r e est refl6t6 p a r les p o l l e n de Anthemis, C h e n o p o d i a c e a e , Plantago lanceolata, Cerealia et p a r Corylus, Betula et Alnus. I1 y a trois r6gions a d d i t i o n e l l e s , caract6ris6 par le crit~re de la v~g~tation. INTRODUCTION

The present study is part of a larger project that aims at providing insights into the development of the vegetation along a west--east transect across the Vosges, including the analysis o f recent vegetation, and pollen, spores and macrofossils in cores from peatbogs and lakes (Janssen et al., 1974). In order to enhance the interpretation of past pollen assemblages, data on the present-day relationships b e t w e e n pollen and vegetation are useful, both on a regional and on a local scale. In this paper the forest vegetation on a regional scale, the regional pollen deposition, and their relationships, will be determined in order to gain insight into the complex nature of the source of the various pollen types, and to determine to what extent the values of the various pollen types in the regional pollen deposition change with the changing patterns of vegetation. Earlier studies have shown that regional pollen assemblages can be interpreted in terms of plant formations. In Europe, Aario (1940) in Finland and Grichuk (1941) in Russia showed the relationship between extensive latitudinal vegetation zones and pollen deposition. This was confirmed in modern lake sediments by Prentice (1978). In North America, studies on such a large scale have been made, for example, b y Wenner (1947), Lichti-Federovich and Ritchie (1966), Mott (1969), McAndrews and Wright (1969), Richard (1976), and Webb (1974). In India a similar kind of study was made by Singh et al. (1973}. Also, on a smaller scale, in areas of over 10 km 2 but not more than a few hundred kilometers across, recent pollen-assemblages can be interpreted in terms of formations. [McAndrews (1966), in the prairie/ forest transition of the Upper Midwest in the U.S.A.; Ritchie (1974) in the forest/tundra transition in the Mackenzie River Delta Region.] In the Near East formations on such a scale were recognized in Iran by Wright et al. (1967), and in southern Turkey b y Van Zeist et al. {1970). In Scotland Birks (1973) established a relationship between alpine, subalpine and woodland formations anc~ the regional pollen deposition. Altitudinal vegetation zones, usually on a smaller scale, also can be

185 recognized in pollen assemblages. Examples in North America are in the Olympic Peninsula (Heusser, 1969), in Colorado (Maher, 1963), and in Arizona (King and Sigleo, 1973). In South America Salgado-Labouriau (1979) has demonstrated similar relationships. In establishing the relationships between regional pollen deposition and vegetation, the application of a number of criteria, that are not always clearly observed in the literature, is useful: (1) The influence of small local stands in the pollen deposition must be minimized. From this it follows that a vegetation map showing the location of the surface samples and the distribution of vegetation types must be provided. (2) The use of moss polsters for the determination of the regional pollen deposition has the advantage that only aerial transport of pollen is involved. The origin of pollen in lake sediments, especially in lakes with an inlet, can be at a considerable distance (Peck, 1973; Bonny, 1978). In lakes with an outlet, part of the pollen assemblages in suspension may be carried out during the summer season when the lake is stratified and the pollen input for the current year is suspended in the upper layers of the lake. Even in lakes without an inlet or outlet, pollen is washed into the lake from the surrounding slopes (West, 1973), especially during phases of deforestation (Pennington, 1979). Thus, transport by water may lead to incorporation into lake sediments of pollen which is not part of the regional aerial pollen assemblage. (3) The vegetation types must be spatially well-defined. Lichti-Federovich and Ritchie (1968) have established the concept of the landform vegetation unit, namely an area with a constellation of vegetation types defined by landforms and other features in the landscape. Likewise in Mott's (1969) study, the physiography of the area considered is an important criterion. (4) In order to assess transport of pollen across vegetation boundaries, surface samples should be either on a grid or else along a transect. (5) Pollen studies are quantitative and therefore some quantification of the vegetation is useful. One of the purposes of this paper is to determine whether, on the relative small scale of a west--east transect across the crest of a mountain chain (70 km), landscape regions (Ritchie's landform vegetation units, but on a much smaller scale), defined by vegetation and abiotic features, can be characterized in terms of regional pollen assemblages. If so, then this could be used to interpret past pollen assemblages, at least for the not too distant past. Although vegetation has certainly changed much during the last few thousand years, it may be reasonable to assume that the physiography of an area is not essentially different from that of today. If this is true, then landscape areas of similar extent should be recognisable in the regional pollen deposition o f t h e past. A second aim of this study is to determine the local presence or absence of a taxon from its pollen values.

186

The study of local pollen assemblages, rather than regional assemblages combined with plant macrofossil analyses, constitutes the best approach to the tracing of the nature of the plant community. The approach is especially valuable in ecotones, in regions of different topography, and during times of migration of plant taxa. The many kinds of local pollen deposition, and the regional pollen deposition of a landscape region are complementary features. Insight into the present pollen values under conditions where the pollen-producing taxon does not grow at the site itself is important in deciding whether past pollen values indicate local presence or absence of a taxon. In this way the recognition of the lateral extent of the regional pollen assemblages becomes a tool in the understanding of the various local pollen assemblages. Throughout this paper the concept of local and regional pollen deposition will be used sensu Janssen (1966, 1973). LOCATION OF THE T R A N S E C T AND C R I T E R I A FO R THE RECOGNITION OF LANDSCAPE REGIONS

The 70 km long transect was located in the central Vosges at 48°N lat. and 7 ° 25'E long. Obviously, the pollen/vegetation relationship will be simple when the vegetation patterns are simple; for this reason the transect was not straight. The western part of the transect was NW--SE-oriented and the eastern was W--E-oriented. In this way the transect traverses a number of regions that can be clearly defined and delimited in terms of parent material, landforms, vegetation etc. The following factors were considered in the definition of the landscape regions:

(1) Geology Information on the geology of the area was obtained from Carte Gdol0gique de France 1:80,000: Epinal (85) 1939, 1:50,000: Colmar-Artolsheim XXXVII-18, Munster XXXVI-19; and from Dubois et al. (1955), Hameurt (1967), Blanalt (1969--1971), and Eller {1976). In Fig.1 the number of rock types has been simplified into units that, on account of the concentration of minerals in the weathered products, bear the closest relationship with soil development and vegetation. In this way five units remain: (1) areas with Quaternary sediments; (2) greywacke; (3) granite/gneiss; (4) sandstone; and (5) limestone. The locations of the surface samples are also shown in Fig.1. The transect starts in a limestone area, traverses a belt of sandstone, a large central region of granite/gneiss, an area of sandstone/limestone and ends in the Upper Rhine Plain, where Quaternary sediments prevail.

pp.187-194.

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pp.195-202.

VEGETATION TO DION AN MAP ACCORDING o FORESTRY VEGETATION TYPES DATA /t..' ;

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203

(2) Topography For details of the topography the following Cartes de France 1:25.000 were consulted: from west to east: Rambervillers 5--6, Bruybres 3--4, 7--8, Munster 1--2, 3--4, Gerardmer 5--6, 7--8, Neuf Brisach 1--2, 5--6 and Colmar 5--6. In the most western part of the transect altitudes are below 400 m. They increase eastwards up to 1300 m in the crest region of the Vosges, then decline to ca.200 m in the Upper Rhine Plain.

(3) Climate Precipitation (Issler, 1942; Lecolazet, 1950; Dubois et al., 1955; Lecarpentier and Shamsi, 1972). Precipitation is ca.1000 mm/year in the west and increases gradually eastwards up to 2200 mm/year. In the rainshadow of the mountains the precipitation drops to 1000 mm/year over a distance of only 5--6 km and reaches less than 500 mm/year around Colmar.

Temperature (Rempp, 1937; Issler, 1942; Paul, 1977). The average annual temperature is 9°C at 400 m, 5--6°C at 1000 m, 5°C at 1100 m and 4°C at 1200 m altitude. The increased precipitation and decreased temperature at higher altitudes result in an increase in the number of days with mist, and especially above 1000 m altitude mist is often present (Issler, 1942; Carbiener, 1963, 1966). (4) Forest vegetation The vegetation of this part of the Vosges has been the object of many studies by pedologists, foresters and botanists. It is not the intention to review this literature here; it suffices to mention work by Rol (1954), B~rard (1958), Duchaufour and Jacamon {1959), Polge (1963), Jacamon and Timbal (1974) from the western part of the transect; and by Issler (1942), Rastetter (1959), Carbiener (1963, 1966), Claudel (1963), Boudot (1976) from the summit area and eastern part of the transect. The only forest map covering the entire transect is from Dion {1970), based on the studies listed above. It is drawn on a rather large scale although parts of the transect have been mapped or described in a more detailed way. The western part of the transect is included in the Carte de V6g~tation de France 1:200,000, feuille Nancy (Jacamon and Timbal, 1974) and the eastern part of the transect in Dubois et al. (1955). Part of the Fecht Valley has been described by Dion (1965), the forests just west of the Hohneck-Kastelberg massif have been studied in detail and mapped by Kalis (in prep.) and those of the eastern side of the Kastelberg by De Valk (in prep.). All these studies are, however, typological and qualitative in nature. Palynological data are quantitative and comparison of these data with quantitative vegetation data, such as has been done by McAndrews (1966) and Janssen

204 (1966), is therefore useful. In the Vosges, Ulrich (1965) has listed quantitative data of the forest vegetation from all districts of the Vosges. Polge (1963) and Claudel (1963) listed the same for some districts on the western and eastern slopes of the Vosges. These data are, however, too general or are confined to only small areas in the transect. In view of the scale of this study we need the more detailed data of the separate forest lots (parcelles) in the districts present along the transect. These data were obtained from the "Offices National des For~ts" who manage the national forests (For~ts dominiales) as well as the communal forests (For~ts communales). Four administrative centres were involved: the centre of Colmar, centre of Saint-Di6, subdivision of G6rardmer, centre of Epinal-Sud, subdivision of Bruy~res and the centre of Guebwiller. National and communal forests are divided into forest lots for which, for management purposes, quantitative data of the various tree species, age classes, etc., are kept. In the western part of the transect, part of the forest area is privately owned and for these areas no quantitative data were available (cf. Fig.2). The forest vegetation is shown in two ways: (I) By maps displaying the relative abundance of pollen and vegetation values for each species in each forest lot (Figs.6--14). The available information was reduced to the following abundancy classes: 1. < 5% 2. 5--15% 3. 15--25% 4. 25--35%

5. 35--55% 6. 55--75% 7. 75--100%

The following species have been plotted: Picea abies, Fagus sylvatica, Abies alba, Quercus spp., Pinus sylvestris, Larix decidua, Pseudotsuga menziesii, Castanea sativa and Robinia pseudo-acacia. The information from the administrative centres was rather uneven: (a) Most of the data were biased towards the economically important forest trees, generally conifers. Quantitative data were almost always available for Picea, Pinus, Abies, Fagus, and Quercus, although particularly at the western end of the transect Quercus and Fagus were often n o t separated in the forestry notes and for the For~t Communal de Beaum~nil even Picea and A bies were taken together. (b) In most cases the frequency of a certain forest tree was listed as a percentage of the total a m o u n t of trees. However, in the western part o f the transect the numbers of stems were often listed. This parameter is better suited for a comparison with pollen values since the size of the plot is incorporated into these values. Moreover, the pollen values in surface samples are also expressed in percentages and therefore the numbers of stems were converted into percentages. (c) Tree composition within the various lots was n o t always homogeneous because of differences in forest management or altitude within that forest lot. These differences have not been taken into account in the maps for the

205 individual f o r e s t t r e e species (Figs.6--13). H o w e v e r , c o r r e c t i o n s o f this k i n d w e r e i n c l u d e d in t h e v e g e t a t i o n m a p , w h e r e k n o w n f r o m e x t e n s i v e field e x p e r i e n c e (Fig.2). In T a b l e I t h e n a t i o n a l a n d c o m m u n a l f o r e s t s are listed f r o m w e s t t o east, t o g e t h e r w i t h t h e n a t u r e o f t h e available i n f o r m a t i o n . (II) B y using a v e g e t a t i o n m a p t o display t h e overall t r e e c o m p o s i t i o n o f each f o r e s t l o t a n d t o s h o w t h e r e l a t i o n s h i p b e t w e e n pollen a s s e m b l a g e / v e g e t a t i o n a s s e m b l a g e (Fig.2), an a t t e m p t was m a d e to c o n v e r t t h e inform a t i o n f r o m t h e a d m i n i s t r a t i v e c e n t r e s i n t o v e g e t a t i o n u n i t s t h a t were c o m p a r a b l e w i t h u n i t s f r o m existing v e g e t a t i o n m a p s o f t h e Vosges. T h e m a p o f D i o n ( 1 9 7 0 ) has b e e n t h e basis o f Fig.2 b e c a u s e this m a p covers t h e entire t r a n s e c t . D i o n ' s f o r e s t t y p e s were t e n t a t i v e l y d e f i n e d in t e r m s o f t h e q u a n t i t a t i v e d a t a o f t h e f o r e s t r y n o t e s , b u t b e c a u s e o f d i f f e r e n c e s in scale a n d l a y - o u t it was n o t a l w a y s possible t o d e t e r m i n e t h e areal e x t e n t and c o m p o s i t i o n o f D i o n ' s f o r e s t t y p e s in a m o r e d e t a i l e d w a y . M o r e o v e r , c o m p a r i s o n w i t h D i o n ' s f o r e s t t y p e s is difficult b e c a u s e t h e f o r e s t s h a v e b e e n m u c h d i s t u r b e d b y m a n . This is least t r u e f o r t h e s u b a l p i n e a n d m o n t a n e f o r e s t t y p e s . In t h e colline a n d s u b m o n t a n e z o n e , t h e p a t t e r n o f the f o r e s t s is o f t e n d e t e r m i n e d b y f o r e s t r y m a n a g e m e n t a n d it o f t e n p r o v e d i m p o s s i b l e t o d e t e r m i n e t h e f o r e s t t y p e t o w h i c h a n y individual f o r e s t p l o t belonged. Overall t e n m a j o r f o r e s t t y p e s w e r e r e c o g n i z e d f r o m t h e f o r e s t r y n o t e s o f w h i c h m a n y c o u l d be e q u a t e d with t y p e s r e c o g n i z e d b y D i o n . T h e f o l l o w i n g t e r m i n o l o g y o f t h e m a i n a l t i t u d i n a l v e g e t a t i o n z o n e s is a c c o r d i n g to H a e u p l e r ( 1 9 7 9 ) : A. Subalpine

1. Fagus forest and Fagus krummholz ("H~traie d'attitude" of Dion): Fagus sylvatica is dominant, accompanied by Acer pseudoplatanus and Sorbus aucuparia. B. Montane

2. Abies--Fagus forest ("H~traie--Sapini~re" of Dion): It appears from the forestry

notes that plots in the area classified by Dion as "H~traie--Sapini~re" may have an important admixture of spruce, but usually not over 25%. At lower altitudes pine may be present. 3. Abies--Picea forest: The forestry notes indicate that stands in the area classified by Dion as "~picea" usually have more than 25% spruce. Spruce may even be the dominant tree species, but more often Abies is the most important tree. Fagus is usually present, but it remains subordinate to Abies and Picea. Again Pinus may be present at lower altitudes. 4. Abies forest ("Sapini~re de basse altitude" of Dion): In the area of Dion's "Sapini~re de basse altitude", there is usually more than 75% Abies. The remainder is Picea. C. Colline--submontane

5. Fagus sylvatica--Quercus petraea forest: On the western side of the main chain of

the Vosges Dion distinguishes between the "H~traie---Ch~ne rouvre" type with Fagus sylvatica dominant and a "Ch~naie rouvre--H~tre" type with oak dominant.

In this study no distinction between these types has been made because the forestry notes do not distinguish between deciduous elements (Feuillus) in a number of

Laveline Herpelmont Chapelle-devant Bruy~res Jussarupt Aumontzey

x

x

X x x

x

x x x

x

x

x

x

x x

x

x

x

x

FC H e r p e l m o n t 1 - 6 FC Le T h o l y FD Housseramont

x

x

x

x

x

X

x x x

x x

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Pieea

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X

Acer

X

x

X

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X

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x x

× x

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x

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N

F C Jussarupt 1 - 1 0 F C Li~zey FC Champdray 1-10

Centre de gestion de Gdrardmer

F C G r a n g e s sur V o l o g n e

FC C h a m p d r a y

FC

FC

FC FC

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FC F i r e , n i l FC Beaum~nil

F C Vervezelle

FC Prey

FC Champqe

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Centre de gestion de Bruydres

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x

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X

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x x

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x

x ×

x x x

x

x

x

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x

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x x

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x

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x

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Quereus

Other

Tree t a x a o f w h i c h d a t a o f t h e f o r e s t l o t s were available for t h e F o r ~ t s D o m i n i a l e s ( F D ) a n d t h e F o r ~ t s C o m m u n a l e s ( F C ) (C = Castanea; L = Larix; P = Pseudotsuga; N = n u m b e r o f s t e m s )

TABLE I

t~ 0 O~

Rehaupal Vologne 3 Vologne 1,2 G~rardmer G~rardmer La Bresse 1948--69

FC FC FC FC FC FC FC FC FC FC FC FC FC FC FC FC FC FC FC FC FC FD FC FC FC FC

Soulzeren Stosswihr 1962--68 Miinster Muhlbach Metzeral Mittlach Sondernach Breitenbach Liittenbach Hohrod 1957--76 Eschbach Giinsbach Griesbach Soulzbach Wasserbourg 1970 Wihr-au-Val Walbach Tiirckheim Zimmerbach Colmar Winzenheim Wilsbach Wettolsheim Equisheim Husseren-le Chateau Voegtlingshoffen

Centre de gestion de Colmar

FC FD FD FD FC FC

X

X X X X

X

X X X

C. C. C. C. C.

C.

C.

Co

C.

tO O

FC FC FC FC FC FC FC FC

Hattstatt 1978 Gueberswihr 1966 Rouffach 1974 Pfaffenheim 1968 Westhalten 1961 Osenbach Soulzmatt 1976 Orschwihr 1978

Centre de gestion de Guebwiller

T A B L E I (continued)

x X x x x x x x

%

N

x

x

Abies/ Picea

Acer

x X x x x x x x

Picea

AR x AR x x x AR x

Pinus

x x x x x x x x

Abies

x X x x x x x x

Fagus

x X x x x x x x

Quercus

P,L P,L P,L P,L P,L P,L P P,C

Other

b~ O Oo

209 c o m m u n a l forest areas. A c c o r d i n g to Dion's map, beech is the d o m i n a n t tree species in m o s t of the forests in the western part o f the transect. 6. Quercus robur, Q. petraea--Pinus sylvestris--Abies alba forest: Mostly on sandstone, but also on granite, an oak forest t y p e could be recognized with a strong e l e m e n t of pine and, to a lesser extent, with Abies alba. A t lower altitudes Abies usually does n o t reach into the u p p e r storey of the forest. This t y p e could also be recognized in the western and eastern part of the transect, although it was n o t m a p p e d by Dion. F r o m the forestry notes an almost similar t y p e could be recognized with s o m e Fagus sylvatica. It is listed as t y p e 6b. 7. Quercus petraea, Q. pubescens--Carpinus betulus forest: This t y p e is essentially Dion's " C h i n e rouvre---Charme" forest t y p e along the eastern foothills of the Vosges. It is best developed o n calcareous soils. Pinus sylvestris may be absent ( t y p e 7a) or present (type 7b) in which t y p e it m a y be an i m p o r t a n t species. Finally, the forestry notes f r o m the c o l l i n e - - s u b m o n t a n e zone list p r e d o m i n a n t l y coniferous forest types w i t h o u t Quercus robur or Q. petraea and are difficult to place in one of the main forest types. The d o m i n a n t tree species is either Pinus or Abies with an a d m i x t u r e of Picea or Fagus. F r o m the fact that Quercus is absent, one m a y be inclined to place these types in the group o f m o n t a n e forest types. On the o t h e r hand, these types are m o s t o f t e n located in region I, II, IIIa or IV (see Fig. 2), thus generally in t h e colline-s u b m o n t a n e z o n e on non-calcareous soils. If placed here the m o s t logical group w o u l d be group 6, despite the absence of Quercus. Three types belong to this group : ( 1 ) an A bies--Pinus--Picea forest, (2) an A bies--Pinus forest and (3) a Pinus~Fagus--Abies forest. T h e y are provisionally listed as t y p e 6a. 8. Castanea ( " C h a t a i g n i e r " of Dion) D. Planar

9. Quercus robur--Carpinus betulus--Fraxinus excelsior--Ulmus--Acer campestris forest: This forest t y p e is present along the rivers Ill and Thur, with Quercus-Carpinus on drier ground and Fraxinus--Ulmus and Alnus on w e t t e r sites - - it is similar to D i o n ' s " C h i n e p~doncul~---Charme--Fr~ne--Orme--]~rable" type. E. Plantations 10. Pinus sylvestris p l a n t a t i o n ("Pin sylvestre", "Pin n o i r " of Dion), Pinus over 75%. 11. Vineyards. LANDSCAPE REGIONS

As stated in the introduction (p.185) one of the aims of this study was to determine whether landscape regions within the transect could be recognized by means of the regional pollen deposition. The characteristics of a number of landscape regions will be described below on the basis of the criteria listed above, including the vegetation. However, for the delimitation of the various regions, vegetation is of secondary importance. Although pollen deposition is a direct result of this existence of plant cover only, the vegetation is often patterned according to features that delimitate non-biotic characteristics of the landscape, for example parent material and topography. Because these parameters are, for the time period considered in Upper Quaternary studies, stable features in the landscape, the vegetation patterns in the past may have followed these physical boundaries which can be recognized at the present day even when the vegetation itself may have

210

differed considerably from that o f today. For this reason the delimitation of the following landscape regions is primarily on the basis of parent materials and of topography as follows: Region

Parent material

Altitude (m)

I II IIIa IIIb IIIc IIId IV V

limestone sandstone granite/gneiss granite/gneiss granite/gneiss granite[gneiss limestone/sandstone Quaternary sediments

below 400 400--600 600--1200 800--1000 600--1350--500 600--1000 250--600 below 250

Girecourt Bruy~res Champdray/G4rardmer T~te de Cerf Massif Kastelberg/Hohneck Massif Pt. Ballon Osenbach Upper Rhine Plain

The following landscape regions are described from west to east in more detail:

I. Grandviller--Girecourt region This is part of the larger area of the plain and hills of Lorraine from Nancy up to the western foothills of the Vosges. Parent material: calcareous soils of Liassic and Triassic (Muschelkalk and Keuper) age. Altitudes: less than 400 m. Precipitation: ca.1000 mm/year. Region much cultivated. Woods in well-drained areas with Quercus robur, Ulmus carpinifolia, Carpinus betulus and Tilia cordata (Querceto-Carpinetum); in less well-drained areas with Quercus robur and Fraxinus excelsior (AlnoPadion). Another forest t y p e is the Luzulo-Fagetum with Fagus sylvatica and Quercus petraea on rather dry and acid soils. Often Quercus has been favoured at the expense o f Fagus and the typical herb species associated with Fagus (Luzula luzuloides, Prenanthes purpurea) have been replaced by Avenella flexuosa, Vaccinium myrtillus and Calluna vulgaris. In these locations Pinus sylvestris has often been planted.

II. Bruydres region Parent material: sandstone of Triassic and Permian age (Gr~s Vosgien). Altitudes: 400--600 m above sea level. Precipitation: 1000--1200 mm / year. Area almost completely forested. According to the official vegetation map 1:200,000 this region belongs partly to submontane Quercus petraea-Fagussylvatica forest and partly to montane Fagus--Abies forest, the LuzuloFagion and Asperulo-Fagion, respectively. This is also apparent from the forestry notes: in the northeastern part (Bois de Fouchon) and the southern part of the For~t de Fai'te A bies is more abundant and Quercus less abundant than in the northern part of the For~t de Fai'te. Prior to the early nineteenth century, heath covered large areas (Dion, 1970; Guillet, 1971) especially on

pp.211-212

MAIN SURFACE SAMPLE TRANSECT QUERCUS FIELDS

MAIN VEGETA!7OV TYPES REGIONS

8

I f----lL

IIIa2

III at

•

~_.

9

••

10

••

n

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I

11

12

13

IIIc2 11. 15

I

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I

32

17

33 34 35

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16

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rnJ t-=-----'ll-------'ill.-------'L~~-llJh

n __

FAGUS ABIES

AIBIES-FAGUS

PINUS

I

LOCALITIES Tilia Acer

CENTRAL VOSGES

- ~- -- -ll

=

nl

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I

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Fig.3. Surface pollen diagram of upland trees and shrubs, main transect Vosges.

..

r II

I I j

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C R ionsW! 1970

pp. 213-214

CENTRAL VOSGES

MAIN SURFACE SAMPLE TRANSECT QUERCUS FIELDS

MAIN VEGETATION TYPES REGIONS

8 H

LDCALfTlES

..lC_ Ii

ffi ::t:

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-

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~

15 :::t

C,rslum Plantago media Lychnls Lathyrus Rhlnanthus type Campanula Trifolium repens Trifolium pratense Veronica Melandnum Botrychlum Viola undlff

Material Local

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pp.215-216.

MAIN SURFACE SAMPLE TRANSECT MAIN VEGETATION TYPES

I

QUERCUS FIELDS

REGIONS

I

I

PINUS

I 3 4 I I

LOCAUTIES

CENTRAL VOSGES

II

IIlat

S 6 7 I II

I

8

'()

Orosera intermedia Metampyrum Viola palustris Menyanthes SUCCISO Valeriano dlolea Bidens type Mentha type ,

~

Caltha Fllipendula LYSlmachla Saxifraga stellarls Polygonum blstorta Rumex eu Rumex

I

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III bt UII b2lmc1

III a2

9 II

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FAGUS ABIES

ABIES-FAGUS

22

n

I I -------jl--------~----+---t__~~---+--"------

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X

Fig.5. Surface pollen diagram of lowland herbs and indetenninate pollen types, main transect Vosges.

-

X

X

X

lL

217 poor degraded soils (podzols humo-ferrugineuse). Since that time pine has expanded considerably especially in the eastern and southern parts of this region.

III. Hercynian massif This forms the large middle part of the transect. Parent material: granite/ gneiss.

IIIa. Champdray--Gdrardmer region Consisting of a strip 24 km long from the Vologne river near Bruy~res to the Chajoux river, this region ranges in altitude from 400 m in the valleys gradually increasing to 600--1000 m on the ridges and plateaux in the southeast. Parent material: granite/gneiss, b u t in the most northwestern part on top of the hills rather large areas of sandstone are present, these decrease in size towards the main chain of the Vosges. Generally the valleys are cultivated and the hills are forested, although the plateau of Champdray is free of forest due to a thin layer of Quaternary sediments, thus permitting cultivation. Precipitation: from 1200 m m / y e a r in the northwest to 1500 m m / y e a r in the southeast. Forest vegetation: hills covered generally by montane A bies--Fagus forest, in the northwest with some Quercus and especially on sandstone Pinus sylvestris and in the southeast with Picea abies. The plateau of Champdray is an important plant-geographical boundary: towards the southeast within the transect no oaks and almost no pines are present west of the main chain of the Vosges. In the easternmost part of this region Fagus sylvatica becomes more important in the upper altitudinal reaches. However, it never becomes the sole constituent of the tree layer in the forest.

IIIb. Chain o f the Tdte de Cerf A ca.7 km long part of the transect from the Chajoux river to the Moselotte river. Precipitation: 1500 m m / y e a r up to 2000 mm/year. Altitudes range from 700 m (valleys) up to 1200 m above sea level. Above 1000--1100 m the number of misty days is considerably larger than elsewhere. The valleys are cultivated, except for the upper Chajoux valley, which is covered by a rather extensive Picea abies stand, the grande Basse. Forest vegetation: the difference from the adjoining part of region IIIa is mainly due to the presence of upper subalpine Fagus forest above 1100 m above sea level. A bies is absent above that level and the forests consist of Fagus sylvatica with Sorbus aucuparia and Acer pseudoplatanus (AceriFagion).

218

IIIc. Kastelberg--Hohneck massif The difference from region IIIb is that in the area of the crest region

Fagus becomes stunted (krummholz) towards the top of the mountains and gives way to extensive subalpine meadows (Hautes Chaumes). These treeless areas are dominated by a mixture of Vaccinium myrtillus, Vaccinium vitis-

idaea, Festuca rubra, Nardus stricta, Agrostis vulgaris, Calluna vulgaris, Potentilla erecta and Genista pilosa. The main plant communities here are, according to Carbiener (1966), the Pulsatillae-albae-Vaccinietum uliginosi and the Sorbo-Calamagrostidetum. Altitudes increase gradually on the west side up to 1360 m. To the east of the main chain altitudes drop within a distance of 3 km to ca.500 m; this area is characterized by a number of cirques and canyons. Precipitation is 1500--2000 mm/year. Again as in region IIIb the mountains above 1000 m are often envelopped by mist. Forest vegetation: On the western side of the main chain well-developed Abies alba--Fagus sylvatica forest is present below 1100 m. Above that level Abies is absent and pure Fagus forests (Aceri-Fagetum) are present. At 1100 m Fagus is ca.15--20 m tall, decreasing to ca.8 m at an altitude of 1200 m. Above that altitude Fagus becomes a tortuous shrub. These forest types have been described in detail by Carbiener (1963--1966), Kalis (in prep.) and De Valk (in prep.). On the eastern side the altitudinal zonation is roughly similar to that on the western side. Especially in canyons Ulmus scabra, Tilia cordata, Fraxinus excelsior and Acer pseudoplatanus are abundant (Tilio-Acerion). Around 500--600 m above sea level thermophilous elements such as Carpinus betulus, Juglans regia, Castanea sativa are present as well as Quercus.

IIId. Petit Ballon region Again altitudes rise, up to 1250 m. On top of this massif extensive meadows are present. Below these open areas no clear Aceri-Fagetum exists. The role of Fagus is very subordinate; instead forests dominated by A bies (up to 1200 m) and Picea prevail as well as, at lower altitude, Pinus. East of the valley of the Krebsbach (For~t de Rouffach), still on granite and at altitudes between 400--1000 m Fagus--Abies forests again prevail.

IV. Osenbach region This is a region of sandstone and limestone between 300 and 600 m altitude in an intricate pattern defined by faults along the Upper Rhine Plain. Precipitation: 600--1000 mm/year. Forest vegetation: on sandstone Quercus robur, Q. petraea, A bies alba and Pinus sylvestris dominate. On limestone Quercus pubescens is a b u n d a n t together with other Quercus spp. (Quercion pubescentis p.p.).

219

A large tract of this area is cultivated and exotics like Larix decidua and Pseudotsuga menziesii are often planted together with Pinus and Picea. Castanea is often present in the understorey of the forests and is very abundant just behind the vineyards on the lower slopes of the foothills; a result of its utilization in the vineyards as a " s u p p o r t e r " of Vitis. Also the non-native tree Robinia pseudoacacia is occasionally present, sometimes quite abundantly.

V. Upper Rhine Plain A region of Tertiary and Quaternary deposits which are largely cultivated. The east-facing foothills are covered by vineyards that also occur over a distance of some 10 km on south-facing slopes in the lower Fecht valley. In the Upper Rhine Plain itself cereals are grown except along some of the smaller rivers where patches of wet forests with mainly Quercus petraea, Fraxinus excelsior, Ulmus campestris, Tilia cordata, Alnus glutinosa and Acer campestre (Fraxino-Ulmetum) occur. In some of these forests Juglans nigra, Carya alba and Carya amara (Toussaint and Toussaint, 1969) and quite a number of archaeo- and n~ophytes like Aesculus hippocastanum, Robinia pseudoacacia and Acer negundo (Carbiener, 1970) are present. Further to the east (outside Fig.2) extensive forests of Quercus robur, Quercus pubescens and Carpinus betulus (Dion's "Charme-Ch~ne rouvre" type) are present. Precipitation is 500--600 mm/year. PALYNOLOGY

Selection o f sites The recent pollen deposition was studied by means of pollen analysis of surface samples. For these both Sphagnum or other bryophytes were used. Each sample consisted of 5--10 subsamples from within an area of 4 m 2 in order to obtain average values and to minimize some of the strongest local effects. Samples from 42 sites were obtained of which 27 were in pairs (subsites a and b) to gain insight into the variation of the pollen values from different localities of similar sites. The moss polsters were taken from a variety of localities such as bogs, ditches, walls, etc., situated in open non-wooded areas. The local herb flora for each locality is therefore quite different. This is shown in the greatly fluctuating values for herb pollen, between sites. In Table II the sites are listed together with data on the nature of the moss polsters, local vegetation and altitude. Local surface sample studies (Janssen, 1966, 1973; Heim, 1970; TamboerVan den Heuvel and Janssen, 1976) and theoretical considerations on pollen dispersal (Tauber, 1965) have shown that for trees the local effect is felt over relatively short distances from the forest edge. It is usually negligible at distances b e y o n d 150 m from the pollen source. For herb pollen this distance

220 TABLE II Sites, altitudes, materials and local vegetation of the surface samples Site

Altitude (m)

Moss

Local vegetation

meadow fen meadow meadow meadow meadow r o a d s i d e in o p e n i n g p i n e meadow s h a l l o w d i t c h in m e a d o w roadside a. m e a d o w b. w e t f e n a. r e m n a n t r a i s e d b o g b. w e t f e n remnant bog raised b o g raised bog raised b o g

1. S e r c o e u r 2. VaudEville 3. V i m ~nil 4. G r a n d v i l l e r , l ' ~ t a n g l ' a b b ~ 5. G r a n d v i U e r - B r u y ~ r e s , le m i n e d e s e a u x (5) l ' E c r e v i s s e 6. B r u y ~ r e s , les b a r a q u e s 7. B r u y ~ r e s , la b e r g e r i e 8. H a u t d e T h i ~ m o n t 9. H e r p e l m o n t lO.Champdray, l'epine

330 345 370 375 405 415 475 460 475 500 705

Bryales Bryales Bryales Bryales Sphagnum Bryales Leucobryum Sphagnum Sphagnum B r y ales Sphagnum

11. Marixose

760

Sphagnum

12. 13. 14. 15.

595 620 875 840

Sphagnum Sphagnum Sphagnum a. S p h a g n u m b. B r y a l e s Sphagnum Sphagnum

C l e u r i e , BeiUard Ronfaing H a u t de Merelle, l ' e t a n g Col de Sapois-Anc. Tourbi~res

16. G r a n d ' ~ t a n g 17. G r o s s e Pierre 18. 19. 20. 21. 22. 23.

La Cure, source Moyenmont, pr~ Jaquot Hotel du Corbeau Les Planches Lispach G r a n d e Basse

24. F e i g n e s s o u s V o l o g n e s 25. F e i g n e Pt. A r t i m o n t 26. F e i g n e t~te Pt. A r t i m o n t 27. K a s t e l b e r g 28. P e t i t H o h n e c k 29. F r a n k e n t h a l , c i r q u e 30. F r a n k e n t h a l , f a r m h o u s e 31. R o t h r i e d 32. G a s n e y , a u b e r g e , c i r q u e 33. M ~ l l b a c h , R u u z 34. Mt~hlbach, A l t m a t t k o p f 35. M i t t l a c h 36. A s c h b a c h 37. H a u t R i e d 38. S t e i n b e r g 39. D o r s c h b a c h 40. O s e n b a c h - - W i n z f e l d e n , T h a l a c k e r 41. G u e b e r s w i h r , R o s s l a u f 42. L a u c h - - R o s s w e i d

795 950 955 1000 825 800 905 920 920 1195 1220 1350 1288 1028 1020 835 970 695 715 500 820 900 1110 380 340 230 195

Sphagnum Bryales Bryaies Polytrichum Sphagnum a. S p h a g n u m b. P o l y t r i c h u m a. S p h a g n u m b. P o l y trich u m Sphagnum Bryales Bryaies Bryales Sphagnum Sphagnum Sphagnum Sphagnum B r y ales Bryales Bryales Bryales Bryales P o l y trichu m Bryales Bryales Bryales Bryales

raised bog a. h u m m o c k in b o g b. f l a r k in b o g spring area heath area heath area heath area raised bog raised bog fen n e x t t o l a k e raised bog bog subaipine meadow subalpine meadow fen bog bog spring area h a y field heath area hayfield spring area r o c k in m e a d o w subalpine meadow meadow marsh m a r s h r u d e r a l site d r a i n e d SaUx fen

is (with a possible exception for grass pollen) much shorter, usually not more than a f e w meters (cf. also Raynor et al., 1968). Thus, to ensure that the pollen deposition of trees and shrubs is regional, no samples were taken within a distance of 150 m of tree pollen sources. There are some exceptions, where an unusually high pollen percentage in the surface samples must be explained by the presence of trees nearby (extralocal deposition): viz.

221 sample 26 ca.50 m from low Fagus woods, sample 11, 31 close to local Betula trees, sample 6, 80 m from pine forest, sample 3, 42 close to Salix shrub and sample 4 close to some shrubs of Frangula alnus. Surface sample studies have also shown that the distance from the pollen source over which the local influence is noticeable is very short for most of the herb pollen types, usually within a few meters (Janssen, 1973). With this in mind, the plant species present in a 25-m 2 plot around each surface sample locality were tallied. Pollen values were considered to be "regional" when the corresponding species was absent from the 25-m 2 plot, and "local" when the corresponding species was present, even when the pollen values were similar to "regional" values at neighbouring localities. At any rate the values of a pollen t y p e in a sample from a locality where the plant species is absent must approach the "regional" level. In this regional study lists of local plant species are n o t given, b u t in Figs.3, 4, 5 values from pollen types of plants present in the plot are shown by a narrow line and those that are regional by a thick (solid or open) line. The same applies for trees and shrubs, although of course the size of the plot was larger, with a radius of at least 150 m. For two types no separation into local and regional values could be made: (1) grass pollen: grasses are almost always present in the plots; and (2) Sphagnum spores: when possible the moss polsters consisted of Sphagnum, only once (sample 38b) were these spores found when no Sphagnum was observed at the site.

Laboratory methods Moss polsters were boiled with 10% KOH, sieved (width of meshes 250/~m) and washed several times with water. The residue was centrifuged and subjected to the acetolysis m e t h o d for 5 min. The unstained material was m o u n t e d in silicone oil 2000 CS.

Pollen analysis Whole numbers of slides were counted until a total of more than 500 tree and shrub pollen grains was reached. The pollen sum included the regional pollen types (all the types of Fig.3): herb pollen was excluded because of presence at the site. Where the values of tree pollen showed local overrepresentation (Salix in sample 3, 42;Betula in 11, 31;Pinus in 6 and Fagus in sample 26), the values of the other elements were calculated u p o n a sum including an adapted (regional) figure of these overrepresented species. The results are displayed in three pollen diagrams (Figs.3, 4 and 5), on the maps showing the pollen values o f the dominant pollen types and the vegetation values for each species in each forest lot (Figs.6--14) and on the vegetation map (Fig.2). From the maps, showing the relation between pollen and vegetation values, it is readily clear that the various tree species are quite differently represented in the pollen deposition. However, the representation differs

222 from one region to another, due to differential dispersal of pollen and for that reason correction factors were n o t established. REGIONAL FLUCTUATIONS IN POLLEN VALUES OF INDIVIDUAL POLLEN TYPES In this section the pollen values of the individual pollen types will be discussed for each of the landscape regions. They are expressed in three parameters: (1) range, maximum and minimum pollen value - - w h e n the pollen t y p e occurs in only one sample the actual value found is listed; (2) frequency, expressed in decimal proportion; (3) median value, not calculated when (a) frequency is less than 50% (indicated by + ), (b) when pollen t y p e occurs in one sample only. N.B. : For the c o m p u t a t i o n of these parameters sites that (may) show a local effect were omitted. The values of samples from the subalpine meadows on t o p of the Kastelberg and H o h n e c k were compared with those in Tamboer-Van den Heuvel and Janssen (1976). The latter, however, are discussed only when they differ significantly from those in this study. 1. Tree and shrub pollen types (Fig.3) Abies alba (Fig. 6; pp. 225--232)

I-II: 0--2.9 IIIa: 1.2--10.5 IIIb: 3.5--18.6 IIIc: 2.1--12.5

(0.7) (1.0) (1.0) (1.0)

0.9 4.3 10.5 7.6

IIId: 8.8--14.0 IV: 4.5--5.1 V: 0.5--1.5

(1.0) (1.0) (1.0)

11.7 4.7 1.0

Abies pollen reputedly is poorly dispersed and its representation is consequently low. As a result the pollen values remain relatively low even at sites surrounded by Abies forest. In regions I, II and IIIa, at sites 1--10, the pollen values of Abies do not exceed 5%; and on approaching the vast belt of montane A bies forest east of site 10, the pollen values begin to rise (6--7% at site 11). However, the extensive Abies forests around Bas Beillard (sites 12, 13) are n o t apparent from the pollen deposition, although more to the east the higher representation of A bies in the forests is reflected in the higher pollen percentage of 7--10% at sites 14--17. In the montane Abies belt in region IIIb the Abies pollen values fluctuate considerably. These are generally higher here than elsewhere. In region IIIc there is a sudden drop in the pollen values of Abies to 3--4% in the montane Fagus forest, reflecting the absence of that tree there. In the open areas of the Kastelberg--Hohneck massif of region IIIc (sites 27--30) the values for Abies are 8--12%, those in the Petit Ballon massif slightly higher, 9--14% (sites 36--38), a possible reflection of the A bies belt east of the main chain of the Vosges. Between these areas at lower

223

altitudes (sites 31--34) lower pollen values for Abies are found, even at sites surrounded by mixed A bies forests. East of the Abies belt in region IIId, the role of Abies in the forest decreases rapidly and so do the pollen values: 4--5% at site 40, down to 1% in the upper Rhine Plain. A cer I--II: 0--0.6 IIIa: 0 - 0 . 4 IIIb: 0 - 0 . 6

(0.5) (0.4) (0.5)

0.3 0.1 0.1

IIIc: 0--1.6 IIId: 0 - 0 . 6 IV--V:-

(0.9) (0.7)

0.5 0.4

Acer pseudoplatanus is an important constituent of the Aceri-Fagetum and Ulmeto-Aceretum (the latter with some A. platanoides) at higher altitudes (A. platanoides reaches 1100 m in the mountains). A. campestre has its upper limit at 800 m. The regional pollen deposition reflects this, with the highest values and frequencies being found at sites 21--37. Pollen values are generally higher than those of Tilia despite lower pollen production, no d o u b t a result of a much higher abundance of Acer in the forests. A esculus This ornamental tree is often planted along roads at lower altitudes. Only three pollen grains were found, one in region IIIa and two in region IIIc, of which one was near the crest of the mountains, to which it must have been transported from the valley floor. Alnus I--II: 5.0--14.0 IIIa: 2.0--15.0 IIIb: 4.8--13.8

(1.0) (1.0) (1.0)

8.5 6.5 7.0

IIIc: 1.6--13.2 IIId: 3.3--7.9 IV--V: 1.2--8.9

(1.0) (1.0) (1.0).

6.2 5.6 3.5

Ranges and median are approx, similar in all the regions, except region IV--V. Alnus glutinosa is the native species, occurring up to 1000 m altitude. The introduced A. incana and, especially A. viridis, are rare. The maximum pollen value is found at site 7a, clearly the effect of some local alder scrub, b u t other maxima and minima in the pollen deposition are more difficult to explain: Site 12: no alder in the vicinity, sample is perhaps contaminated with moss from a period prior to the present cultivation of the area. Sites 22, 23: pollen values depressed, probably because of dominance of Picea pollen in the assemblage. Site 32: pollen values rather high. Although at present no Alnus was observed some m a y have been there in earlier times.

224

Betula I-II: 1.6-8.2 IIIa: 3.4-13.5 IIIb: 3.6--9.9

(1.0) (1.0) (1.0)

(24.8)

4.1 9.2 6.8

IIIc: 1.6-11.2 IIId: 3.5--5.8 IV--V: 2.4--7.9

(29.2) (11.0)

(1.0) (1.0) (1.0)

6.0 4.9 3.5

Pollen values in region I - I I are low, but increase in region IIIa, and are like o t h e r pollen types in t ha t Betula is negatively correlated with the decline in Pinus values. However, in region IIIb the Betula values decline as do those of Pinus, indicating t h a t the higher values in region IIIa are real and probably a reflection o f the generally less-forested aspect o f region IIIa com pared with region IIIb, giving rise to Betula pioneer scrub after cultivation. Part o f the Betula pollen m a y have c om e from Betula pubescens around bogs. In region IIIc the pollen values o f Betula are low in m o n t a n e Fagus forest and A bies forest and in subalpine meadows. The dense character of these forests and the absence o f bogs along the eastern slopes o f the main chain m a y explain this. Site 30 however shows inexplicably high Betula pollen percentages. On the massif o f the Petit Ballon the Betula pollen values are comparable with those on the H o h n e c k massif.

Buxus I-II:IIIa: 0.1-0.4 IIIb: 0.2

(0.4) + (0.2) +

IIIc: 0.2 IIId: 0.2 IV--V:0.2

(0.4) +

(0.3) +

Within the transect Buxus is absent. Pollen values are low in all the regions with frequencies 0.2--0.4. The closest pollen sources are in the calcareous region to the west (Meuse and u p p e r Marne valleys) and comes f r o m Fagus forests with a dense Buxus u n d e r s t o r e y , (Durin et al., 1964), t he Upper Rhine Plain and, in the southeast, t he Sundgau.

Carpinus I: II: IIIa: IIIb: IIIc: IIId: IV: V:

2.2-3.2 0.8--5.8 3.1-11.7 3.2--9.2 (21.5) 3.3--8.7 2.4--7.8 9.0--12.9 3.0--4.4

(1.0) 2.2-5.8 (1.0) (1.0) 7.0 (1.0) 7.3 (1.0) 6.3 (1.0) 6.2 (1.0) 10.6 (1.0) 4.2

(1.0)

3.8 (I + II)

Carpinus is absent above 800 m and even below t hat level it is rare in the area covered b y the transect. As a result, m o s t of the pollen f o u n d in the samples must have travelled over rather long distances. Two possible pollen

_pp. 225--232.

N

ABIPS RELATION ]IIe I

ALBA

POLLEN PERCENTAGES I TREE PERCENTAGES

®

-~

ebsent <5% 5-15~

25-35 ~ 35-55 55-?5%

/

I.

75-100~

ZEa2 @

~J ®

V

I 0 I

1 I

2 I

; IV

3 Km ]

Fig.6. Regional pollen percentages and distribution of Abies alba.

I

@

<_ ~/_~..>I_.

N

CASTANEA SATIVA ROBINIA PSEUDOACACIA RELATION

POLLEN PERCENTAGES

/ TREE PERCENTAGES

III c I ( ~ (CASTANEA)

absent <5% 5-15% 15-25% 25-35 % 35-55% 55-?5% 75-100 %

/

@

III e z

~q

Rob.~ Rob.

/

f V

® IV 0 I

1 I

2 I

3 Km 1

Fig.7. Regional pollen percentages and distribution of Castanea sativa and distribution of Ro binia pseudoacacia.

/

O:)

pp. 241--248.

N

FAGUS SYLVATICA RELATION

POLLEN PERCENTAGES / TREE PERCENTAGES

®

---] ebsent

O

5-15% 15-25% 25-35% 35-55% ">.-~.,..2.,.,...

R

55-?5%

mm

75-100%

I

< IIIa 2

<<
/ / (9

V

(9 /

0 I

1 I

2 I

IV

3 Km I

Fig.8. Regional pollen percentages and distribution of Fagus sylvatica.

I

®

3 N

LARIX DECIDUA RELATION

POLLEN PERCENTAGES I TREE PERCENTAGES

®

--~ absen t <5% 5-15%

15-25% 25-35 % 35-55%

55-?5%

m

75-100%

I

III a 2

/

D:.

\ W

/

®

0

I

1

I

2

I

3 Km

I

V

/

7 IV

Fig.9. Regional pollen percentages and distribution of Larix decidua.

I

Of)

1

/

pp. 257--26; i

N

PI CEA ABI ES

/

RELATION

POLLEN PERCENTAGES / TREE PERCENTAGES

03

III c I

absent

<5% 5-15% 15-25% 25-35% 0

0~

35-55 %

m m

55-?5% 75-100 %

I

©

d

/ © 0

1

2

I

I

t

3 Km

I

/

7

Fig.10. Regional pollen percentages and distribution of Picea abies.

I

v ®

N

PINUS SYLVESTRIS RELATION

POLLEN PERCENTAGES' / TREE PERCENTAGES

CJ

absent

< 5% 5 -15% 15-25%

,

lillll·::IIIIIII::ill:liiil·:1

25-35 %

\~ji::"'::::lt~

35 -55 %

:~~~~~~~

,::;:

55-75% •

,

75-100%

o

V

o

1

2

I

I

I

3 Km I

Fig.II. Regional pollen percentages and distribution of Pinus sylvestris.

§

pp.273-280.

N

PSEUDOTSUGA MENZIESII RELATION

11101

POLLEN PERCENTAGES / TREE PERCENTAGES

D D

o

absent

<5% 5-15%

C'

o I

•

15-25% 25-35% 35-55%

•

55-75%

•

75-100%

v 1

2

I

I

3 Km I

Fig.12. Regional pollen percentages and distribution of Pseudotsuga menziesii.

QUERCUS sp. pp.281-288.

N

RELATION

POLLEN PERCENTAGES / TREE PERCENTAGES

G

D

absent

(5% 5 -15% 15-25% (

25-35%

/

35-55%

•

55-75%

75-100%

V

o I

1 I

2 I

3 Km I

Fig.13. Regional pollen percentages and distribution of Quercus sp.

s

289

sources are: (1) the extensive forests of Quercus and Carpinus in the Upper Rhine Plain east o f the forests along the Ill and Thur river, and (2) the calcareous soils in the northwest. As a result of this distant pollen source the differences in the pollen values between the regions are slight. It is because of this regional character of the pollen deposition that in pollen diagrams from the crest region Carpinus is one of the best markers of synchronous horizons.

Castanea (Fig.7; pp. 233--240) I--II: -IIIa: 0.2 IIIb: 0.2--0.4

(0.3) (0.~)

+ +

IIIc: 0.2--0.6 IIId: 0.2--0.4 IV--V: 0.2--0.5

(0.2) (0.6) (0.5)

+ 0.2 +

The most probable source of Castanea pollen is from the east of the transect where the species is c o m m o n beyond the vineyards. It does n o t occur above an altitude of 800 m. There are few differences between the regions in the transect except that frequencies and m a x i m u m values are slightly higher closest to the source, in regions IIId, IV and V. Higher values of Castanea, however, were also reported from the subalpine meadows of region IIIc (Tamboer-Van den Heuvel and Janssen, 1976).

Cory lus I--II: 0.8--6.0 IIIa: 4.0--15.4 IIIb: 3.0--16.6

(1.0) (1.0) (1.0)

3.0 7.8 10.2

IIIc: 5.0--14.2 IIId: 8.0--14.0 IV--V: 1.4--13.8

(1.0) (1.0) (1.0)

9.0 12.3 8.6

The upper limit of Corylus is at 1300 m, often above the actual upper limit of Fagus along steep south-facing slopes. It is most abundant on limestone in the west and east, occasional on hercynic rock and rather rare on sandstone. Maximum pollen values and the median are very low in region II, as m a y be expected from forested areas on base-poor soils. The Corylus values increase in region IIIa, probably a result of the presence of Corylus scrub here. The gradual increase eastwards, however, does n o t correlate with an increase in the a m o u n t of Corylus present, rather the opposite. The increase in pollen values therefore appears to be related to the decrease of the values of Pinus pollen. The Corylus values in region IIIb and IIIc are, at sites 18--21, around 15%, at sites 22--32 ca.10%, then again around 15% at sites 33--35. These fluctuations reflect the fact t h a t Corylus t o d a y is almost absent at higher altitudes. The changes are, however, also relative to the importance of other forest elements. The samples from sites 18--21 are from rather open, cultivated areas, and those of sites 22--24 and 28--32 are from densely forested areas in which Corylus is almost absent and where, in the pollen deposition, the

290

forest pollen types are better represented. In the open spaces of the Hohneck-Kastelberg massif {sites 27--29} there is, however, rather inexplicably, no rise of Corylus pollen values, despite the absence of Corylus and other trees. A rise of Corylus is however noticeable in another open area, that of the Petit Ballon (sites 36--38). In contrast to the Hohneck--Kastelberg area, Corylus is present in the vicinity of sites 36 and 37, even within 100 m of site 37. The massif of the Petit Ballon is also less extensive than that of the Hohneck and closely surrounded b y valleys and lower slopes. In region I and regions IV--V the Corylus values are very low, despite abundant Corylus in these regions, especially at sites 40 and 41. These low values are difficult to explain. One reason might be the high Pinus values, depressing the values of other pollen types.

Fagus (Fig.8; pp. 241--248) I: 2.9--3.8 II: 10.2--12.0 IIIa: 4.5--16.3 IIIb: 9.4--28.5

(1.0) (1.0) (1.0) (1.0)

3.5 11.2 10.0 15.0

IIIc: 8.1--30.8 IIId: 5.2--13.0 IV: 12.7--15.4 V: 1.0--1.6

(1.0) (1.0) (1.0) (1.0)

16.0 9.0 13.0 1.6

The lowest pollen values of Fagus are shown in the most western and the most eastern parts of the transect {sites 1, 2, 3, 41, 42), both from areas with wide open spaces w i t h o u t much forest. In region II, however, 10--12% of the tree pollen is from Fagus. This m a y be partly a result of the proximity of Fagus and partly due to the relative effect of lower Pinus percentages. In the western part of region IIIa (IIIa--1) the percentages remain at this level. Although n o t always listed in the forestry notes, Fagus is indeed an important tree in the forest, especially in the northern tip of the For~t Dominiale de Jussarupt and Herpelmont. East of the plateau of Champdray the role of Fagus in the forest declines and this is clearly reflected in the pollen percentages, which decrease to 5%. The percentages of Pinus pollen decline too, thus the decrease of the Fagus values is not the relative effect of changes in the a m o u n t of Pinus pollen b u t rather a result of a real diminishing pollen output. In the eastern part of region IIIa, the values of Fagus rise to 9--16%. Again there is no negative correlation with Pinus b u t a positive correlation with an increasing a m o u n t of Fagus trees in the forest. In region IIIb the Fagus values are somewhat higher than in the two most eastern samples from region IIIa, reflecting the increased importance of Fagus in these forests. However, the Fagus values fluctuate considerably, apparently due to local conditions that are not easy to understand in detail. L o w values are sometimes from open areas {sites 18, 23), although in site 19, from the t o p of a bare mountain, the Fagus values are much higher. L o w values of Fagus are, of course, also correlated with high percentages of other pollen types, like those of Picea at sites 22, 23. At any rate the highest Fagus values (30%) are foun~l in the westernmost samples of region IIIc from m o n t a n e Fagus forests.

291

In the subalpine meadows in the crest region the Fagus percentages decrease to 15--17% or slightly lower, they decrease to 10% (TamboerVan den Heuvel and Janssen, 1976) in samples from the same region. In the eastern part of region IIIc, at lower altitudes, similar or lower values are shown, reflecting the unimportant role of Fagus in these forests. An exception is the Frankenthal cirque (site 29) where values reach up to 23%, an example of the crest effect (p. 218). In the region of the Petit Ballon (region IIId) the number of Fagus trees is small in the forest, reflected by low pollen values at sites 36--38. The considerable amount of Fagus in the area between the Petit Ballon and the Osenbach--Winzenheim area is n o t reflected in the samples. The value at site 40 (13%) is similar to that of the Petit Ballon (site 38). In samples from the Upper Rhine Plain Fagus pollen is almost absent.

Fraxinus I--II: 0.4--2.0 IIIa: 0.3--3.0 IIIb: 0--4.0

(1.0) (1.0) (0.9)

1.2 1.4 1.2

IIIc: 0--3.0 IIId: 1.0--2.7 IV--V: 1.3--5.3

(1.0) (1.0) (1.0)

1.2 1.4 2.8

Fraxinus excelsior is present along the entire transect in damp habitats, especially in valleys and along brooks, up to 1200 m altitude. Ranges: the high frequencies and median values are similar in regions I--IIId, b u t from individual sites in region IIIb and IIIc, however, they differ considerably. The pollen values at sites 29--31 are located around a cirque and may have been responsible for higher Fraxinus pollen values, since special air currents in cirques could transport Fraxinus pollen more than 150 m below the cirque where this tree is abundant.

Juglans nigra type One pollen grain was found in region V where this tree is planted in alluvial forests (Toussaint and Toussaint, 1969).

Juglans regia I--II: -IIIa: 0.2--0.4 IIIb: 0.2--0.4

(0.2) (0.3)

+ +

IIIc: 0.1--0.2 (0.8) IIId: 0.2--0.5 IV--V: 0.2--2.4

(0.3) (0.5) (0.6)

+ + 0.6

Like Castanea the most probable source is in the valleys of regions IIId, IV and V, where the walnut is cultivated and often used as a roadside tree. Pollen is present in all the regions. The upper altitudinal limit is 800 m. Ranges and frequencies in regions IIId, IV and V are distinctly higher than elsewhere.

292

Lar/x (Fig.9; pp. 249--256) I--II: 0.2 IIIa: -IIIb: 0.2

IIIc: 0.1---0.4 IIId: 0.2 IV--V: 0.2--0.5

(0.25) (0.15) (0.5)

Larch (tamarack) plantations date from late in the last century. From Fig.9 it appears that Larix plantations in the east are present in regions IIIc and IIId, although not as extensive as those of Pseudotsuga. Larix and Pseudotsuga have a similar pollen morphology but despite this, pollen of Larix is better represented than that of Pseudotsuga. Larix pollen is, however, smaller and this, together with the generally older plantations, may explain its higher range and frequency in the east. Picea (Fig.10; pp. 257--264) I--II: IIIa: IIIb: IIIc:

0.5--5.5 0.5--14.8 3.5--26.6 2.5--19.4

(1.0) (1.0) (1.0) (1.0)

1.9 9.0 10.0 8.4

IIId: IV: V:

3.6--8.8 2.4--4.3 2.2--6.3

(1.0) (1.0) (1.0)

6.1 3.4 2.3

In regions I--II, IIIa the pollen values of Picea are similar to those of Abies: viz. very low at sites 1--10 (except at site 9) and 10--15% at sites 11, 14--18. The amount of Picea in the forests is generally lower than that of Abies, but the pollen representation of Picea is better than that of Abies, probably a result of the well-known differences in dispersal capacities between pollen of A bies and Picea. In the western part of region IIIb the Picea pollen values remain the same as in region IIIa, except at sites 22 and 23 (16--20%), which reflects an extensive local Picea stand there. In the eastern part of region IIIb, where Picea is less frequent in the montane Fagus forest and in the subalpine meadows, the Picea pollen values are correspondingly lower: 4--8% in the Upper Mosellotte valley, 2--5% in the Fagus forest and in the subalpine meadows (except at site 28). The Picea stands below the Frankenthal cirque are clearly reflected by the higher percentages at sites 29, 31 and 32. At lower altitudes the Picea pollen values decrease, reflecting the decreasing amount of Picea in the forests. In region IIId, at the massif of the Petit Ballon, Picea pollen values range from 4 to 9%, generany somewhat higher than at the Kastelberg--Hohneck massif, reflecting the more important role of Picea in the Petit Ballon. Finally, in region IV and V the Picea values decrease. Site 40, closest to spruce stands, does not show elevated values. Pinus (Fig.11; pp. 265--272) I: II: IIIa: IIIb:

66.0--77.5 41.3--77.5 41.3--52.5 20.5--55.5 13.6--25.4

(1.0)

66.0

(1.0) (1.0)

30.6 22.2

IIIc: IIId: IV: V:

13.5--45.8 27.2--37.4 25.3--37.0 66.0--75.0

(1.0) (1.0)

26.0 30.4

293 The highest values of Pinus are found in regions I and V, where this tree certainly is present b u t n o t to a very wide extent. The relative importance in the pollen deposition thus must be a result of the low total pollen o u t p u t of other trees. Indeed, in these regions the dominant tree species are Quercus, Fagus, A bies and Picea with pollen being dispersed over shorter distances. Pinus trees, however, are nearby (in regions II and IV), its pollen is produced in large quantities and is easily dispersed which results in an overrepresentation of Pinus pollen in regions I and V and thus depressed pollen values for most of the other pollen types. In region II and up to site 11 in region IIIa, the Pinus pollen values are lower (except at site 6, where Pinus is within 150 m distance) as a result of an increased share of pollen o f Fagus and Picea in the regional pollen deposition. From site 11 eastwards, the values of Pinus begin to decline, marking the eastern limit of that tree in the western part of the transect. At two sites east of the Pinus sylvestris limit, Pinus montana (uncinata) grows on bogs, in the Tourbi6re de Beillard and in the Gohte Loiselbt (Les Ht. Pinasses: Guinier, 1959), b u t this is n o t reflected in the pollen deposition. In region IIIb Pinus is absent and the values of Pinus pollen decline due to the increasing distance from Pinus to the west and the increasing amounts of Fagus, Abies and Picea in the forest. On top of the Kastelberg--Hohneck massif, trees are absent and a small increased "long distance e f f e c t " is shown here. In region IIIc Pinus is present again at lower altitudes and this is reflected in increased Pinus pollen values at the sites closest to the Pinus plantations, especially sites 33--35. On t o p of the Petit Ballon (region IIId) Pinus is absent and the Pinus pollen values have declined b u t do n o t reach the low values shown on top of the Kastelberg--Hohneck massif. Unlike the Kastelberg, Pinus is present all around the base of the Petit Ballon, and this is neatly reflected in the pollen deposition. No such reflection whatsoever is seen at site 40, despite the presence of extensive Pinus stands.

Pseudotsuga (Fig.12; pp. 273--280) One pollen grain was found at locality 40b from the middle of the area in which this tree has been planted below an altitude of 900 m since about 1910 (For~t Communal de Hattstatt), often in large quantities. Pseudotsuga pollen is much under-represented because of its great weight. Quercus (Fig.13; pp. 281--288) I: 11.5--16.5 II: 3.8--14.0 IIIa: 6.9--1.2.7 IIIb: 6.3--12.8

(1.0) (1.0) (1.0) (1.0)

15.0 9.2 9.6 9.5

IIIc: (4.7) 7.1-10.8 (12.7) (1.0) 9.8 IIId: (5.2) 9.5--10.8 (12.6) (1.0) 10.2 IV: 9.4--14.8 (1.0) 14.2 V: 9.3--12.0 (1.0) 9.3

Quercus robur does n o t exceed the 500 m altitudinal limit, Q. sessiflora that of 750 m. As a result, this species is absent in the eastern part of regions IIIa, IIIb and the western part of region IIIc.

294

In regions I, II, IIIa (west), IIId, IV and V Quercus is a constituent of the forest. It is dominant especially in the eastern part of region IV. However, the Quercus pollen values are only slightly raised when compared with those in other regions. Again the reason m a y be the overrepresentation of Pinus pollen. Another explanation is possible selective preservation of Quercus pollen in moss samples. Sal/x I--If: 0.2--1.0 Ilia: 0.2--1.2 IIIb: 0.2--0.8

(I.0) (0.8) (0.7)

0.6 0.4 0.4

IIIc: 0.1--0.9 IIId: 0.2--1.2 IV--V: 0.3

(1.0) (0.8) (0.15)

0.4 0.8

In the regional pollen deposition there are no striking differences between the regions. Sorbustype I--II: 0.2 IIIa: 0.2--0.4 IIIb: 0.2--0.4

(0.3) (0.2) (0.3)

IIIc: 0.2--0.6 IIId--IV--V: --

(0.3)

Sorbus aucuparia is present everywhere on hercynic rock and on sandstone. Most of the pollen grains were found in the middle part of the transect, at sites 16--35, and are thus in the higher parts of the Vosges. The conspicuous occurrence of Sorbus in the Aceri-Fagetum of the main chain is n o t reflected in the samples of that area. Higher percentages than found here are only present in local samples (Tamboer-Van den Heuvel and Janssen, 1976). Other Sorbus species that m a y have been a source for this pollen t y p e include S. aria and S. mougeotii, both present at middle and higher altitudes in the mountains. Tilia I--II: -IIIa: 0--1.5 IIIb: 0--0.6

(0.5) (0.3)

0.1 +

IIIc: 0--0.3 IIId: 0--0.3 IV--V: 0--0.6

(0.3) (0.3) (0.5)

+ + +

Apart from ornamental trees along roadsides and in towns, Tilia is rare everywhere. The main occurrence of non-planted Tilia is in region IIIb and IIIc. Tilia platyphyllos is present occasionally in Fagus--Abies forests, and especially in the Ulmeto-Aceretum (Canyons) up to 900 m. Tilia cordata has its upper limit at 800 m. At the Hohneck and Rainkopf a few specimens of T. intermedia are found at 1200 m. This distribution is n o t reflected in the regional pollen deposition. The pollen values in region IIIa are slightly higher than in other areas, perhaps a reflection of planted T. cordata. Overall pollen values are very low as may be expected from the entomophilous nature of pollen dispersal.

295

Ulmus I--II: 0--0.6 IIIa: 0--1.0 IIIb: 0--2.2

(0.5) (0.9)

(0.9)

0.15 0.4 0.8

IIIc: 0.2--1.9 IIId: 0--1.5 IV--V: 0--1.5

(1.0) (0.8) (0.8)

0.6 0.4 0.7

Pollen of Ulmus is most abundant in regions IIIb and IIIc. Ulmus scabra occurs in the same sites as Fraxinus, up to 1000 m altitude. The highest values are f o u n d at sites 29--31, like those of Fraxinus. The same transport mechanism over longer distances as assumed for Fraxinus may thus also apply to Ulmus.

Hedera According to Issler (1942), Hedera helix occurs in Quercus--Carpinus and Quercus--Betula forests at lower altitudes. It also can be quite abundant in m o n t a n e Abies--Fagus forests (Kalis, in prep.; De Valk, in prep.). It does not exceed the 1000 m altitudinal limit. In spite of this no Hedera pollen was found in samples from the east and from the mountains. Hedera pollen however was present in samples from the subalpine meadows in the study of Tamboer-Van den Heuvel and Janssen (1976). Three grains were from sites in the westernmost part of the transect. Viscu m Only one pollen grain was f o u n d in locality 7a, despite the presence of Viscum on A bies, especially in the middle part of the transect below 800 m. Frangula alnus (4 x), Ligustrum (1 x), Comus mas (2 x), Comus sanguinea (3 x), Viburnum (1 x) and Lonicera (1 x) These taxa do n o t occur in the upper altitudinal areas (the upper limit of Cornus and Ligustrum is at 800 m, that of Lonicera at 900 m, and that of Viburnum at 1200 m). The regional pollen values of these insect-pollinated plants with low dispersal capacity are low, not over 0.2--0.3%. The high values of Frangula at site 4 are due to local occurrence. There is no explanation for the rather high values of Comus sanguinea at site 41. 2. Upland herb pollen types (Fig.4)

a. Pollen from cultivated plants Ambrosia One pollen grain was f o u n d from the top of the Petit Ballon massif. Ambrosia is n o t native in Europe and pollen may have escaped from urban areas. Centaurea cyanus One pollen grain has been f o u n d at site 11, at the margin of the Champdray

296 plateau. The plant is a companion of cereal fields. In the earlier study of Tamboer-Van den Heuvel and Janssen (1976) a grain of Centaurea cyanus was also observed in a sample from the Kastelberg. Cerealia

Secale

non-Secale

IIIa: IIIb: IIIc: IIId: IV: V:

0-2.7 0--0.7 0.2--0.4 0.2---0.4 0.4--0.5 (0.6)

(0.6) (0.3) (0.3) (0.5)

0.2

2.3--3.9

0-0.8 0--0.8 0--0.4 0--0.6

(0.5) (0.5) (0.25) (0.7)

0.2--2.1

(0.7)

0.4

Presence a n d / o r ranges are generally higher in the western part of region IIIa and in regions IV and V, where extensive cereal fields are present. Secale does n o t exceed the 900 m altitudinal limit. Outside these regions uniform pollen values are found. Fagop y ru m One pollen grain was f o u n d in a sample from the Petit Ballon massif and Tamboer-Van den Heuvel and Janssen (1976) occasionally f o u n d it in the subalpine meadows of the Kastelberg. This plant is cultivated in the Upper Rhine Plain, southeast of the transect area. Vitis Vitis vinifera is present at lower altitudes in regions IIId and V. Vitis pollen is present in regions IIIb--V and absent from the western part of the transect, indicating that the pollen source is in the east. The values in regions IIIb and IIIc are n o t over 0.2%, frequencies ca.0.25. At site 41 (region V), surrounded by vineyards, still only five pollen grains were found. The wild vine (V. sylvestris) is rare, restricted to forests in the Upper Rhine Plain. Zea mays Two pollen grains have been found at site 42 from a region where maize fields are present. However, also two pollen grains have been observed in samples from the top of the crest of the mountains and in a sample from region IIIb, both from regions where Zea is completely absent. b. Upland pollen types, present in all regions A n t h e m i s type I--II: 0.6 IIIa: 0--0.9 HIb: 0.2

(0.2) (0.55) (0.15)

0.2

IIIc: 0.2 IIId: 0--0.5 IV--V: 0--1.0

(0.1) (0.3) (0.4)

The highest pollen values are from region IIIa, reflecting the relation with cultivated fields. Pollen source is therefore probably Matricaria, A n t h e m i s and Achillea, genera associated with cereal fields.

297

The proximity of fields is also reflected in samples from regions IIId, IV and V. Pollen values throughout the middle part of the transect are not over 0.2% and frequencies not over 0.2. Artemisia /Chenopodiaceae Chenopodiaceae

Artemisia

I--II: IIIa: IIIb: IIIc: IIId: IV--V:

0.2 0--0.4 0--0.8 0--0.8 0-0.6 0.8

(0.3) (0.5) (0.8) (0.5) (0.3) (0.2)

0.2

0--1.0 0--0.6 0--0.8 0--0.5 0--1.2 0.3--0.9

(0.5) (0.55) (0.6) (0.6) (0.7) (0.8)

0.7 0.7

The ranges and frequencies of pollen of Chenopodiaceae are slightly higher in the western and eastern part of the transect, reflecting the presence of Chenopodiaceae in ruderal areas. The species of Artemisia have a similar distribution, although the highest pollen values are from the middle and more forested part of the transect (region IIIb and region IIIc), a p h e n o m e n o n difficult to explain. Asteraceae liguliflorae I--II: 0--0.6 IIIa: 0--2.2 IIIb: 0--1.6

(0.75) (0.55) (0.35)

0.3 0.2

IIIc: 0--1.6 IIId: 0.9--2.4 IV--V: 0--0.9

(0.4) (0.8) (0.7)

1.5 0.4

The high pollen values in a number of samples are due to local occurrence b u t when these sites are left o u t of consideration there is a marked maximum from the Petit BaUon massif. The source is probably Leontodon pyrenaicum, abundant in subalpine meadows. Maxima in the pollen values are also found in the eastern part of region IIIc (sites 33--35) and at sites 19, 21 (region IIIb), also near to meadows b u t at lower altitudes. Minimum pollen values are found near cereal fields and in forested areas. R u m e x acetosella I--II: 0.3 IIIa: 0--1.6 IIIb: 0--1.1

(0.15) (0.5) (0.7)

0.5

IIIc: 0--1.5 IIId: 0--0.8 IV--V: 0.3--1.0

(0.7) (0.8) (1.0)

R u m e x acetosella is a regular c o m p o n e n t of the pollen assemblage from site 10 eastwards with maximum values reaching 1.5% and with slightly increasing frequencies towards the east.

0.4 0.3 0.3

298

Urtica Pollen values are low throughout; however, identification is n o t always reliable and therefore ranges and frequencies are n o t listed.

c. Upland pollen types from regions I--IIIc Cen taurea pratensis t y p e Pollen values n o t over 0.4% and frequencies not over 0.15%.

Heracleum H. sphondylium is present in meadows and at forest edges up to 1200 m. Pollen is most frequent in region IIIa: 0--1.0, (0.2), and regions I--II: 0--0.6 (0.3). In region IIIc--d it is less c o m m o n .

d. Upland types from region III Pteridium IIIa: 0.2 IIIb: 0--0.2 IIIc: 0--0.2

(0.9) (0.9)

(0.05) (0.3) (0.25)

Pteridium aquilinum is very c o m m o n in open areas in regions with acid rock. The pollen distribution in the transect reflects the presence of extensive

Pteridium fields at middle altitudes in region IIIb--c. Pollen values and frequencies, however, remain low, even when the plant is locally present because of sterile stands. At only two sites (9, 34) were higher local values observed.

Genista/Sarothamnus Sarothamnus scoparius is c o m m o n on acid rocks below 1000 m. Genista sagittalis has its upper limit at 1350 m, Genista pilosa at 1400 m. Despite the widespread occurrence of these species, the regional pollen values are low (0.2--0.3%) and frequencies are n o t over 0.1. Increased pollen values are seen where scrub is locally present (sites 20a, 33). However, around sample 37 (2.1%) no Genista or Sarothamnus scrub has been observed.

Galeopsis type, Polygonum convolvulus, Scabiosa, Sanguisorba officinalis, one grain in region IIIa.

e. Upland types from region IIIb Scleran thus, Phyteuma, Oxalis, Sanicula, Polypodium Pollen grains o f the species were observed only once or twice. Except

Scleranthus these pollen types are from taxa typical of forest vegetation and are therefore only present in the regional deposition from the most forested part of the transect.

299

f. Upland pollen and spore types from region IIIc, IIId L ycopodium selago This plant occurs above 800 m in Calluna heath and subalpine meadows. One spore was found in a sample from the crest region (IIIc).

Helianthemum, Polygonum aviculare Pollen values do n o t exceed 0.2%. Pollen was observed in samples from the lower eastern slopes of the Kastelberg massif and from the subalpine meadows of the Petit Ballon.

g. Upland pollen types from regions IIIc, IV--V Sanguisorba minor Two pollen grains were recorded from areas where the plant is present and one pollen grain from the subalpine meadows of the Hohneck where the plant is absent.

Epilo bium One pollen grain recorded from region IIIc.

Solanum nigrum, Vicia villosa type, Geum type, Polygonum persicaria One or two pollen grains were found in regions IIId and/or V. 2. Upland pollen types from meadows

a. Pollen types from all regions Plantago lanceolata I--II: (0.4) IIIa: IIIb: IIIc"

2.8--6.5 0.5--5.0 0.7--5.7 1.0--3.0

(1.0) (1.0) (1.0) (1.0)

3.7 2.5 1.8 2.1

IIId: IV: V:

1.4--1.8 5.0--7.4 0.3--1.4

(1.0)

1.5

(1.0)

0.8

The pollen values are rather irregular, the highest values tend to be from regions I--II, IIIa and IV. These are generally the most cultivated areas where hayfields dominate (sites 19, 20, 21, 40, 2--5), the lowest values from densely forested areas (sites 14, 15, 22--24), or from the subalpine meadows on t o p o f the mountains (sites 27, 28) where Plantago lanceolata is absent. The upper altitudinal limit is at + 1200 m. Region V also shows a low Plantago lanceolata pollen deposition.

Cirsium type Pollen values rarely exceed 0.2%; frequencies 0.2 in regions IIIa, IIIb, IIIc. Site 2 has 2.2% Cirsium : no specimens of this taxon were observed here.

300

Plantago media Pollen values usually not over 0.2%; frequencies around 0.15.

Lychnis type Pollen values in regions IIIa, b, c and IV--V 0.2%; frequencies 0.1. On the Petit Ballon (region IIId) however 0.2--0.9 (0.6) 0.4.

Lathyrus Pollen only locally in the pollen deposition.

Rhinanthus type Pollen values usually not over 0.2%; frequencies 0.1. An exception is on the Petit Ballon: 0.2--0.6 (0.4).

Campanula Pollen values in region I--III 0.2--0.4%; frequencies not over 0.15.

b. Pollen types from regions I--II, eastern part of IIIc, IIId, I V - - V (absent from the central core of the Vosges)

Trifolium repens Pollen values in region I--II: 0.2--0.6% (0.6) 0.3; in the eastern part of region IIIc and region V 0.2%.

Trifolium pratense Pollen values not over 0.2%; frequencies not over 0.15.

c. Pollen and spore types from region IIId Veronica, Melandrium, Botrychium and Viola Pollen and spore types only once observed in the samples. 3. Bog and fen pollen types (Fig.5) These are pollen types that, because of the selection of sites for sampling, most often show local values. The sites are, however, not always from similar vegetation types and regional values are therefore displayed when the taxa involved are absent from the local vegetation.

a. Bog pollen types Andromeda Andromeda pollen was recorded twice, at site 23b and at site 26, both from raised bogs. Like Empetrum pollen the source may have been the bog although of course not within 10 m of the sampling site.

301

Calluna I-II: 0.3-0.6 IIIa: 0.2-0.8 IIIb: 0.2--0.4

(0.4) (0.3) (0.75)

0.3

IIIc: 0.2-0.7 IIId: 0.2 IV--V: 0.2

(0.7) (0.3) (0.2)

0.2

Local pollen values are much elevated, but regional values do not exceed 0.8%. Frequencies are highest (0.7) in regions IIIb and IIIc. In these regions most of the samples where Calluna was not locally present still come from heath areas in which the most probable source must be sought. Elsewhere pollen frequencies do n o t exceed 0.4.

Comarum type Pollen f o u n d only locally in region IIIb.

Comarum or Potentilla I--II: 0.2 IIIa: 0.2--1.8 IIIb: 0.2--1.2

(0.3) (0.5) (0.7)

IIIc: 0.2-0.4 IIId: 0.2 IV--V:-

(0.25) (0.15)

Regional pollen values are very low, except in region IIIa, IIIb and this probably is also related to the presence of heath areas and bogs in these regions, perhaps even from bogs and heath areas from which the samples were collected.

Drosera Pollen f o u n d only locally in region IIIb.

Empetrum Pollen only f o u n d once, at site 23b, from a raised bog. Empetrum was not present within 10 m of the sampling site; it is present elsewhere in the bog and the source of the pollen grain was probably nearby.

Melampyrum Pollen f o u n d once, on the top of the Kastelberg. Melampyrum may grow in bogs, but the nearest source is the subalpine meadows.

Menyan thes Pollen only recorded once at site 21a from a dry heath area. Menyanthes is a poor fen species, thus the pollen source must be outside the area of the sampling site.

Sphagnum Whenever possible Sphagnum was used as the surface sample collected. Therefore, no regional pollen value could be determined in these cases.

302

Only in t w o samples {sites 24, 38a) consisting of other mosses, were single Sphagnum spores observed. Regional value is therefore very low, almost zero.

Succisa Pollen found twice in region IIIc, in samples from areas where this plant is absent. Vaccinium Pollen was only regionally present in samples from sites at the western slope of the main chain {sites 21--27) and one pollen grain in sample 34a. Values low, 0.2--1.0%. Vaccinium was never locally present but the pollen source could be bogs (V. uliginosum) or heath areas (V. myrtillus) from which the samples were collected. Valeriana dioica Pollen was only locally present at site 37. Summary. Bog pollen types, even pollen from Calluna and spores from Sphagnum are very underrepresented in the regional pollen deposition. When they do occur, the pollen source either is local (and then sometimes huge amounts of pollen are found), or the source is within the general area of the vegetation t y p e from which the samples were collected; in the latter case just a few pollen grains are usually present in the samples. b. Pollen types from rich-fen species Bidens t y p e Pollen was only once observed in regions IIIa, IIIb and twice in region IIIc. In only one case (site 2 1 ) B i d e n s pollen must have travelled over larger distances than from within the general sampling area. Mentha type Pollen was recorded once in regions II and IIIa. Three pollen grains were found in region IIIb and region IIIc. In the vegetation of the sampling area Mentha was n o t observed. Caltha Pollen was only present in regions IIIa, b, c. Values do n o t exceed 0.6%, frequency 0.15. Filipendula I--II: 0.2 IIIa: 0.2--0.3 IIIb: 0.2--0.4

(0.6) (0.25) (0.3)

IIIc: 0.2--0.8 IIId: 0.2 twice IV--V: 0.4

(0.5)

0.3

303

Pollen grains were present in samples from all regions, mostly in region IIIc. Values were low (< 1.0%), although frequencies were higher than for any other fen pollen type. More than other bog or fen pollen types, Filipendula pollen is present from sites where this plant is absent. Pollen of Filipendula is small and produced in large quantities and therefore regularly present in the regional pollen deposition.

L ysimachia Pollen was only occasionally present in region III with values of 0.2% and frequency of 0.15.

Polygonum bistorta Pollen only recorded once in region IIIc.

Rumex--eu Rumex type Three pollen grains were found at the Petit Ballon.

Lythrum Pollen was only locally present in region IV.

Blechnum Spores only found once in regions IIIa and IIIc. 4. Ecologically indeterminate pollen types (Fig.5)

Apiaceae (includes Meum, Peucedanum, Anthriscus, Apium type, Chaerophyllum ) I--II: 0.2--1.8 IIIa: 0--1.2 IIIb: 0.2--0.3

(1.0) (0.5) (0.6)

0.5 0.2 0.3

IIIc: 0--1.0 (0.8) IIId, IV, V indeterminable

0.4

Pollen of Apiaceae is least abundant in forested regions (sites 12--17, 22--26). Asteraceae tubuliflorae (non-Anthemis, non-Centaurea cyanus, non-Artemisia,

non-Bidens) Pollen was found in samples from all regions. Values not over 0.5% and in region III frequencies not over 0.3. Brassicaceae I--II: 0.3--0.6 IIIa: 0--0.9 IIIb: 0--0.9

(0.3) (0.6) (0.8).

+ 0.3 0.3

IIIc: 0--1.2 IIId: 0--0.8 IV--V: o n l y o n c e

(0.3) (0.8)

Pollen was present everywhere in the samples from region III with rather high frequencies, usually not over 0.9%.

+ 0.6

304 Cyperaceae I--II: 0.2 IIIa: 0--1.2 IIIb: 0--1.0

(0.7) (0.6}

0.4 0.8

IIIc: 0-1.1 IIId: 0.4, 0.6 IV--V: 0.2

(0.8)

0.6

(0.3)

L o w pollen values r e c o r d e d t h r o u g h o u t , with slightly higher values f r o m regions IIIa, IIIb, IIIc. Pollen sources were p e r h a p s t h e bogs and fens c o n c e n t r a t e d in these regions.

Dryopteris t y p e ( m o n o l e t e fern spores) I-II: 0.2--0.4 (0.3) IIIa: 0.2--1.4 (0.9) IIIb: indeterminable

+ 0.6

IIIc: 0.2-5.8 IIId: 0.2--0.5 IV--V: 0.2

(0.9) (0.3) (0.8)

2.2 +

T h e spore values are l o w outside t h e m o u n t a i n s . In regions IIIa and IIIc, w h e r e ferns are a b u n d a n t in t h e u n d e r s t o r e y o f the forests, m a x i m a and m e d i a n are m a r k e d l y higher, u p t o 5.8% in region IIIc. Dryopteris spores are t h u s t r a n s p o r t e d f r o m t h e u n d e r g r o w t h o u t into the o p e n areas. A similar result was o b t a i n e d b y T a m b o e r - V a n den Heuvel and Janssen (1976). Ferns are also a b u n d a n t in the u n d e r s t o r e y o f region IIIb, b u t the regional values c o u l d n o t be d e t e r m i n e d because o f local presence o f ferns at a l m o s t all t h e sites.

Equisetum Spores were o n l y locally p r e s e n t in sample 18.

Galium t y p e I--II: 0.4-0.6 IIIa: 0-0.5 IIIb: 0-0.9

(0.4) (0.5) (0.9)

+ + 0.6

IIIc: 0-1.2 (0.6) 0.3 IIId--IV--V: indeterminable because of local occurrence

Pollen values are slightly raised in region IIIb--c, reflecting p e r h a p s the presence o f Galium hercynicum in treeless areas o n t o p o f t h e m o u n t a i n s . Ranunculaceae I--II: 0--1.7 IIIa: 0-1.4 IIIb: 0-1.2

(0.75) (0.6) (0.85)

0.5 0.2 0.3

IIIc: 0-1.0 (0.75) IIId, IV, V indeterminable

0.3

305 RELATIONSHIPS OF TREE-POLLEN ASSEMBLAGES TO LANDSCAPE REGIONS One of the purposes of this study was to detect whether the regional pollen deposition was related to the seven landscape regions as defined on the basis of parent material, topography, climate and vegetation. We have seen (p.184) that in non-mountainous regions a clear relationship exists not only when the landscape regions are very large, covering hundreds of kilometers, b u t also on a much smaller scale, such as in McAndrews' (1966) study covering a distance of 105 km. The scale of the present study in the Vosges is similar to that of McAndrews (the transect is 70 km long). However, the results of McAndrews' study cannot be applied to the Vosges without reservations. In mountains like the Vosges during clear skies, air currents move upslope and up the valleys during daytime, and downslope and d o w n the valleys at night (Geiger, 1971). A second factor is wind velocity, which generally increases with altitude. According to Jochimsen {1979), regional air currents dominate in the crest region of the (~tztaler Alps in Austria from wind speeds of 6 m/sec, upwards, resulting in a considerable increase of the pollen concentration during times of low air pressure. Carbiener (1966, 1972) recognized a crest effect (l'effet de cr~te): in west--east oriented valleys on the western side of the main chain, wind velocities increase, resulting in a low-pressure area behind the main chain. This gives rise to a compensating air current in cirques. These mechanisms may be responsible for the mixing of pollen from one region to another, blurring the possible delimitation of the landscape regions on the basis of pollen. From the preceding section, dealing with the individual pollen types it is readily clear that the regional pollen values do differ from region to region. In this section an a t t e m p t will be made to define the landscape regions by differences in the regional pollen deposition.

Differences of the regional pollen values Figs.3, 4 and 5 reveal that the most striking differences in the pollen values are between the dominant wind-pollinated species such as Fraxinus, Ulmus, Fagus, A bies, Picea, Corylus, Pinus, Quercus, Betula and Alnus. N o t surprisingly, Pinus pollen is the most important t y p e present in the samples, with widely fluctuating percentages of 33--77%. The pollen values are relative and the values of the other pollen types therefore bear a direct relationship to that of Pinus. It appears that the areas of the seven landscape regions can be defined on the basis of differences in the regional pollen values of the most dominant tree-pollen t y p e s viz. Pinus, Quercus, Fagus, Picea, Alnus, Corylus and Betula. However, in some regions, a subdivision can be made on the basis of combinations of these pollen types, viz. in regions IIIa, IIIb, IIIc. The pollen assemblages of the various landscape regions will be discussed below. Table III shows a summary of the main results.

306 TABLE III Characterization of landscape regions by main tree pollen types Pollen assemblage Pinus--Quercus Pinus--Fagus Picea--A bies---Fagus Fagus--A bies--Picea Fagus Fagus--Abies--Picea Fagus--Quercus--Pinus Pinus-Quercus

Region I ~ II IIIa-1 IIIa-2 { IIIb-1 IIIb-2 IIIc-1 IIIc -2 ~IIId IV V

Difference

Corylus--Betula with region II Alnus--Corylus---Betula with IIIb-2

A bies

In addition to differences in the values of the main tree pollen types there are also differences between those of herb and shrub pollen types and in some minor tree pollen types. These data are given in a differential figure (Fig.14), depicting in which regions a particular ponen t y p e shows values markedly different from adjacent regions. The lines in Fig.14 are not based on calculations b u t on estimates b y visual inspection of Fig.3, Fig.4 and Fig.5. Regional pollen assemblages o f the landscape regions Region 1: Pinus--Quercus assemblage This region is clearly delimited in the regional pollen deposition. Although Quercus is dominant in the forest vegetation, the principle pollen type is Pinus, followed b y Quercus. On account of the strong overrepresentation of Pinus most of the other tree pollen types have low percentages. Region H: Pinus--Fagus assemblage This region is characterized b y lower values of Pinus and b y increased values of Fagus. Region IIIa: IIIa-1: Pinus--Fagus assemblage IIIa-2: Picea--A bies--Fagus assemblage The pollen assemblages of this region are n o t homogeneous. In the western part, up to sample 10 (region IIIa-1), the Pinus and Abies values are similar to those in region II; in the eastern part of region IIIa (region IIIa-2) the Pinus values decline, whereas those of Abies and Picea rise. The differences are clearly related to the eastern boundary of Pinus west of the main chain of the Vosges. East of sample 10, in region IIIa-2, Pinus is absent and the higher degree of aforestation is reflected in the raised A bies and Picea values. Quercus also has its eastern b o u n d a r y here, b u t this is n o t reflected

307

<

H

I F

landscape regions HERACLEUM ALNUS GENISTA- PTERIDIUM RUMEX ACETOSELLA DRYOPTERIS CYPERACEAE OXALIS- SCL ERANTHUS- PHY r-SANICUL AMENYAN THE5 -POLYPODIUM GALIUM CALLUNA VACCINIUM ANDROMEDA FIL/PENDULA SUCC/.SA ACER SANG MINOR-EPILOBIUM-SOLA DULC.-VIC/A V/T/5 VILLOSA TRIFOLIUM ASTERACEAE L/GULIFLORAE ANTHEMIS TYPE CHENOPODIACEAE PLANTAGO LANCEOLATA CEREALIA CORYLU5 BETULA

Fig.14. Differential distribution of some minor pollen types over the landscape regions.

in the pollen assemblages. The distinction from region II may be seen in the Corylus and Betula values: low in regions I--II, increased in region IIIa. This probably reflects the degree of aforestation in the two regions. It is small in region II, and larger in region IIIa. Essentially this is the same feature often seen in pollen diagrams from the Postglacial Period during phases of increased land occupation promoting expansion of pioneer vegetation of Corylus and

Betula. Region IIIb: Fagus--A bies--Picea assemblage This region is, compared with region IIIa, characterized by increased Fagus values and decreasing Pinus values. The A bies values, though fluctuating, are at a generally high level. In these features the general character of the forest vegetation, viz. a zone with Fagus forest above a Fagus--A bies forest, is neatly reflected. There is a strong difference between cultivated areas (sites 18--21) and forested areas (sites 22, 23}. In cultivated areas Alnus, Betula and Corylus have increased pollen values, whereas in the forested region of the upper

308 Chajoux river, a large Picea stand makes its influence on the pollen deposition felt beyond the 150 m limit.

Region IIIc: IIIc-l : Fagus assemblage IIIc-2: Fagus--A bies--Picea assemblage In the pollen deposition there is a clear separation between montane Fagus forest (Fagus maximum, Picea, A bies and Pinus minimum in the pollen values) and subalpine meadows and the eastern slopes of the central chain. It is, however, doubtful whether the two assemblages from the montane Fagus belt are truly regional in character, since large open areas, necessary for the determination of regional tree pollen assemblages, are rare in the area covered by montane Fagus forest. The pollen differences along the eastern slopes are small, except that the Pinus and Corylus pollen values are somewhat raised at lower altitudes. Region IIId : Fagus--A bies--Picea assemblage The pollen values of the massif of the Petit Ballon are .similar to those from the top of the Kastelberg--Hohneck massif, except that the Abies values are slightly higher, reflecting the dominance of A bies at lower altitudes here. Region IV: Fagus--Quercus--Pinus assemblage Region IV is clearly delimited by low Abies and Picea pollen values and by increased Quercus values. Region V: Pinus--Quercus assemblage This region is, like region I, characterized by greatly increased Pinus values. Consequently most of the values of the other pollen types are low, except those of Quercus. The landscape regions also show, in addition to the main tree pollen types, differences between the values of some herb and shrub pollen types. These differences plus those of Alnus, Betula and Corylus are shown in Fig.14. Pollen types related to cultivation, e.g. Anthemis type, Chenopodiaceae, Plantago lanceolata and Cerealia are clearly most abundant at the two ends of the transect where fields and meadow areas prevail. Cerealia pollen is most abundant in region IIIa--1 (the cultivated plateau of Champdray), in region IV (the cultivated area near Osenbach), and in region V (the Upper Rhine Plain). Pollen of Asteraceae liguliflorae are, in pollen diagrams, usually typical of phases of cultivation. However, in the transect, the main distribution of this pollen type is restricted to regions IIIc--2 and IIId, probably reflecting the presence of subalpine meadows in these regions. Trifolium type is also restricted to the part of the transect east of the main chain. Pollen of Vitis, Sanguisorba minor, Epilobium, Vicia villosa type and Solanum dulcamara also have a mainly eastern distribution along the transect,

309

although these types are also found west of the crest region, in region IIIc--1. Quite a few pollen types are most abundant in region III, the large middle portion of the transect. Pollen of Genista and spores of Pteridium are, although in low frequencies, almost exclusively found in all the subregions of region III. Pteridium aquilinum and Sarothamnus scoparius are indeed typical for region III, occurring on abandoned fields and meadows on hercynic rock. A more restricted distribution shows Dryopteris type, coinciding with the middle, forested part of the transect in region IIIa--2 and region IIIc--2, where ferns are often quite abundant in the understorey of the forest. Other " f o r e s t " pollen types, e.g. Oxalis, Phyteuma, Sanicula, Polypodium, have an even more restricted occurrence viz. in region IIIb. Also restricted in various degrees to the middle part of the transect is the pollen of Menyanthes, Galium, Calluna, Vaccinium, Filipendula and Succisa. Most of the corresponding taxa are constituents of peat bogs, from which most of the samples were taken. Although these pollen grains certainly do n o t have a strictly local origin, they may originate from these bogs and therefore cannot be considered really regional. The same may apply to pollen of Cyperaceae. CONCLUSIONS

The major vegetational sequence from west to east across the mountains, from Quercus--Pinus, Fagus--A bies--Picea, Fagus, and back to Fagus-Abies--Picea and again Quercus--Pinus is reflected in the pollen values of the major forest dominants and in some herb pollen and spore types, such as Genista, Pteridium and Dryopteris. There is also a good reflection of the degree of aforestation of the area, viz. b y the values of pollen of Anthemis, Chenopodiaceae, Plantago lanceolata, Cerealia and by Corylus, Betula and Alnus. The differences reflect, of course, changes in the vegetation pattern. One of the objects of this study was to detect to what extent vegetation and regional pollen deposition is based on differences of abiotic parameters like parent material and topography. We have seen (p.000) that these characters are stable features in the landscape. Certainly the vegetation was different in the past. However, the patterns m a y be determined partly by these stable features and the past vegetation might be partly patterned along lines that separate present-day landscape types. This study shows that the separation of regions based on these abiotic features is also reflected in the regional pollen deposition. In Figs.3, 4, 5 this is indicated b y solid vertical lines. However, three regions, IIIa, IIIb and IIIc, could be subdivided, based on vegetational criteria solely, viz. region IIIa--1/2: degree of aforestation and distributional limit of Pinus and Quercus; region IIIb--1/2: degree of forestation; region IIIc--1/2: altitudinal limit of Fagus.

310

These boundaries do not have roots in the non-biotic features of the landscape. It therefore looks likely that these boundaries will move with time. However, the original boundaries of the regions are more fixed features in the landscape and possibly the regional pollen assemblages from the past can be delimitated along similar lines. Once this regional groundwork has been established in the past the ever-changing pollen assemblages from local stands within the individual regions can be interpreted in terms of species composition. ACKNOWLEDGEMENTS

This study could n o t have been carried o u t without the helpful assistance of the "Ing6nieurs des Travaux des Eaux et For~ts" of the forestry districts within the transect, viz. Mr. G. Deschaseaux (Bruy~res), Mr. J. Desforges (G6rardmer), Mr. J. Toussaint et Mr. J.M. Bosshardt (Colmar) et Mr. A. Labarri6re (Guebwiller). They all kindly allowed me to use the data of the forest inventory of their districts. The drawings were capably done by J. Linhart, H. R y p k e m a and T. Schipper.

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