Biologiske Skrifter udgivet af Det Kongelige Danske Videnskabernes Selskab Bind 11, nr. 5

Biol. Skr. Dan. Vid. Selsk. 11, no. 5 (1962)

EXPERIMENTAL AND CYTOLOGICAL STUDIES ON PLANT SPECIES

VII. GERANIUM SANGUINEUM

BY

TYGE W. BOCHER and MARTIN G. LEWIS D et Kongelige Danske Videnskabernes Selskab udgiver følgende pub­ likationsrækker :

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E jn a r M u n k sg a a r d , Publishers, 6 Nbrregade, Kobenhavn K, Denmark. Biologiske Skrifter udgivet af Det Kongelige Danske Videnskabernes Selskab Bind 11, nr. 5

Biol. Skr. Dan. Vid. Selsk. 11, no. 5 (1962)

EXPERIMENTAL AND CYTOLOGICAL STUDIES ON PLANT SPECIES

VII. GERANIUM SANGUINEUM

BY

TYGE W. BÜCHER and MARTIN C. LEWIS

København 1962 i kommission hos Ejnar Munksgaard Synopsis The experimental material comprises 36 strains of Geranium sanguineum from various localities in Europe, grown under uniform conditions. All strains have the same chromosome number i.e. 2n = 84. Variation patterns investigated: (a) Leaf morphology. A cline in leaflobe width is found from E. to \V. Europe. A striking environmental modification of leaf shape is shown, (b) Leaf size. A cline from open to shaded habitats is found in the Danish material, (c) Petal moiphology. Variation in petal shape shows a degree of correlation with leaf shape, (d) Height and habit. Low coastal races and taller continental races are discernible. Three habit-types are found i. e. prostate, cushion and erect, (e) Flowering and vegetative phenology shows considerable variation which can be correlated with the climates of the native areas, (f) Hairiness. In Danish material there is a relationship between degree of hairiness and openness of the original habitat, (h) Flower colour. Although there is considerable variation, this is randomly distributed. Within this experimental material it is possible to recognize 2 race groups (I) Coastal race group (II) Woodsteppe race group. However these 2 groups only re­ present the ends of a more or less continuous clinal system. The taxonomic status of var. prostratum and var. lancastriense are considered. The significance of the variation in leaf morphology is discussed at some length. For several reasons it is believed that the leaf shape is of definite selective value: (I) The same distribution pattern of leaf types is found in several other species. (II) Variation in leaf shape is concurrent with ecotypic differentiation. (Ill) Narrow leaf lobing, such as is found in continental strains, is a well-known xeromorphic character, considered to be of selective value in dry climates. (IV) Environmental modification of the leaf parallels genotypically controlled adaption.

PRINTED IN DENMARK BIANCO LUNOS BOGTRYKKERI A/S Introduction his paper is designed as another in the series of experimental studies undertaken Tby the senior author in cooperation with others. The purpose of the series is to supplement our taxonomic knowledge with information on the genetic-biological structure of the plant species. This involves studying genetic variability of the species, using information obtained from a study of living plants growing in cultivation. Inter-race patterns investigated include those involving the morphology, cytology and phenology of the species. Apart from the desirability of recording this variation, studies of this kind can be of value in revealing recent evolutionary trends within a species. In Geranium sanguineum, a striking variation in leaf shape was noted between strains from various parts of Europe. A more detailed study of this aspect was thought to be important, considering the taxonomic significance of leaf shape in the genus. Here, as in other genera (e.g. Ranunculus) leaf shape often provides the most dist­ inctive morphological character separating species. Variation in leaf morphology has proved useful in understanding evolutionary mechanisms in certain other genera. In this connection several investigations can be mentioned. W oodson (1947) was able to make a detailed analysis of introgression in the Asclepias tuberosa complex using leaf shape characters. Zimmermann’s work (1959) on the Pulsatilla aggregate illustrates how leaf characters can help in eluci­ dating recent history and modes of spéciation within a genus. In Taraxacum and Alchemilla it is possible to distinguish apomictic micro-species using small differences in leaf outline. Petersen’s examination of ecological leaf differentiation within the species Anthriscus sylvestris (1915, 1922), Potentilla erect a (1926) and Pimpinella saxifraga (1921) is particularly pertinent to the present paper and will be mentioned later in more detail. Leaf size has also proved useful in understanding the differentiation in tetra- ploid Veronica officinalis (B ôcher 1944). Genetically-determined differences in leaf area within this species form both a topocline from oceanic to continental regions and an ecocline from open to wooded localities. Other work on leaf morphology has centred mainly around morphogenetic problems. The striking changes in leaf morphology incident on submersion in water have received much attention. There is also a large volume of work on leaf xeromor- l* 4 Nr. 5 phism. Some plants, when grown under dry conditions will develop xeromorphic characters. In many plants, the upper leaves are more xeromorphic than those below. There has been much discussion about the adaptiveness of these structural changes to a water deficit (Maximov 1929; Shields 1950). However, only a few workers (e.g. Tukesson 1922, 1925; Fisher 1960) have investigated genetic variation in leaf shape within a species in terms of the selective response of the species to differences in environment. The junior author intends to study this aspect more fully using Geranium sanguineum and other species. In Veronica officinalis, leaf size forms “the backbone of the variational structure of the species” (B ocher 1944). In Geranium sanguineum, leaf shape fulfils the same role; it is central to an understanding of the organization within this species. In this paper other interracial patterns have been considered against this background. The characters investigated include petal morphology, height and habit, flower colour, flowering and vegetative periodicity, and hairiness. The senior author acknowledges with gratitude financial aid given by the Carls­ berg Foundation for the collection of material in various parts of Europe. The junior author expresses his thanks to N.A.T.O. Studentship Schemes for financial support during the period of this work.

Material and Methods The experimental material comprises 36 strains of Geranium sanguineum from various localities in Europe (see Table I), grown under uniform conditions in the Experimental Garden of Copenhagen University.

Field nos. 647-3195 have been in cultivation since 1953 - 3400-3533 - - - - - 1954 - 3796-4001 - - - - - 1955 - 4145-5111 - - - - - 1959 B 1-B 6 - - - - - 1959

The plants were grown from seed with the exception of strains 10, 14, 23, & 24 which were transplants from nature. Except in 4 numbers, 5-15 plants per strain were available. The leaf samples were pressed in 1960; 5 leaves were taken from each plant at different levels between the stem apex and base. The flower samples were harvested at the same time. Pressings of entire shoots were made in 1961; these were later used in studies on hairiness and leaf size. Root-tips from pot-grown plants were fixed in Miintzing’s modification of Navashin-Karpechenko and embedded in paraffin wax; these were later sectioned and stained in Feulgen’s Fuchsin. Root tips from strains 23, 24, and 32 were fixed in 3:1 Alcohol-Acetic after pretreatment in saturated aqueous solution of Monobromo- naphthalene; these were later stained in Feulgen’s Fuchsin and squashed. Nr. 5 5 T a b le 1.

Country Field No. Locality Ecology Collector of origin No.

Denmark 1 647 Dybdal, nr. Aalborg, N. Jutland ...... T. W. B. Steep grassy 2 648 Fosdal, nr. Fjerresslev, N. J u tla n d ...... — 3 649 Vilsund, N. Jutland ...... — 4 650 Boserup, nr. Roskilde, Zeeland...... Wood edge — — 5 657 Skredbjærg, N. Zeeland...... 1 Steep grassy 6 2073 ...... slope — 7 2558 Dybesø, nr. Rørvig, N. Zeeland...... Stony rais. beach — Dry grassland K. Larsen 8 2563 Hirtsholmene, Island off N.E. Jutland. . . i 1 at the sea 9 2940 Bulbjærg, N. Ju tlan d ...... Fixed dunes T. W. B. 10 3400 Hammershus, Bornholm ...... Grassy slope — 11 3514 Tværsted, nr. Hirtshals, N. Jutland ...... Fixed dunes — 12 3533 Gudhjem, N.E. Bornholm ...... Coastal wood K. Larsen 13 5032 Kyndby, nr. Frederikssund, Zeeland...... Steep slope T. W. B. 14 5111 Bulbjærg, N. Ju tlan d...... Fixed dunes — 15 B 1 Kaas, nr. Lime, N. Jutland...... Oak wood — 16 B 2 Løgstør, N. Jutland ...... Calcareous slope —

Poland 17 m i Mikolajki, M azuria...... Steppe H. Jonassen

British 18 1189 Avon Gorge, nr. Bristol, England...... Limestone cliif Hope-Simps. Isles 19 1190 Yorkshire, England...... — grassland — Crevice in 20 2598 Poulsallagh, Co. Clare, E ir e ...... ' limestone D. A. Webb 1 pavement 21 3180 Loch Bunny, nr. Gort, Co. Clare, Eire . . . — T. W. B. 22 3193 Loch Derg, nr. Terryglass, Tipperary, Eire Lake edge — 23 B 5 Walney Island, Lancashire, E ngland...... Fixed dunes T. G. Tutin 24 B 6 — — — — ...... — — —

Sweden 25 2559 Vickleby, Öland...... Alvar vegetation T. W. B. 26 2560 — — ...... — — — 27 2562 G otland...... — — — 28 2561 — ...... Dry grassland —

France 29 2381 ♦Besançon...... Seeds from 30 2382 ♦Rouen ...... nature obt. 31 2383 ♦D ijon...... through var. 32 B 3 ♦Rouen ...... Bot. Gardens

Germany 33 2384 ♦Frankfurt am Main...... —

Italy 34 3796 Pogerola, nr. Salerno...... Gravel beach K. Larsen

Erica carnea Yugoslavia 35 4001 Samobor, nr. Zagreb...... < T. W. B. heath

Austria 36 4145 Gumpoldskirchen, nr. V ien n a...... Steppe T. W. B. Biol. Skr. Dan. Vid. Selsk.ll,no.5. 2 6 Nr. 5 T a b le IL

Flowering Height of Plants Mean & range, cms. Flower Period Leaf index colour Strain No. type (See 17/6 1961 acc. to (See Fig. 2-6) 17/9 1953 20/9 1960 New cultiva­ Wanscher tions Fig. 12)

1 40 (30-49) 34 (32-35) 40 (35-45) 1 Strong violet 3 2 19 (10-28) 25 (20-30) 13 (10-18) 2 Str. purplish 4 violet 3 47 (40-56) 40 (40) 48 (45-50) 1 — 3 4 27 (19-32) 27 (27) 17 (15-20) 1 Strong violet 4 5 18 (10-24) 15 (10-25) 22 (20-25) 3 — 4 6 18 Dead Dead 4 7 15 (13-18) 7(7) 10 (10) 4 Strong purplish 5 violet 8 26 (15-36) 32 (30-34) 33 (30-45) 2 — 3 9 17 (15-20) 15 (15) 2 — 4 10 20 (19-20) 18 (15-20) 5 Light violet 4 11 7(5-9) 7 (5-10) 4 Strong violet 5 12 ' 25 (15-30) 33 (30-40) 5 Strong purplish 2 violet 13 7(7) 17 (15-20) 4 — 3 14 15 (11-17) 4 Strong violet 5 15 10 (10) 4 — 5 16 7(7) 4 Light violet 5 17 34 (34-35) 39 (37-40) 35 (30-40) 7 Strong violet 2 18 27 Dead Dead 4 19 24 (20-28) 13 (10-15) 10 (10) 4 Strong violet 3 20 29 (20-34) 37 (35-40) 5 Light violet 2 21 17 (10-21) 14 ( 7-25) 5 Strong purplish 4 violet 22 23 (16-31) 21 (10-25) 5 Str. viol. - str. 3 purpl. viol. 23 Still in pots Very pale red 4 24 —— 4 25 27 (22-37) 28 (23-35) 22 (20-25) 6 Strong violet 1 26 36 (25-44) 35 (32-40) 27 (20-35) 6 Strong purplish 1 violet 27 33 (30-38) 38 (35-40) 37 (30-40) 6 — 1 28 28 (26-30) 23 (20-25) 1 Strong violet 1 29 15 (15-16) Dead Dead 2 30 30 (24-38) 21 (15-30) 22 (20-25) 2 Strong violet 5 31 45 (43-47) Dead Dead 2 32 Still in pots 4 33 37 (30-45) 19 (18-20) 27 (25-30) 7 Strong violet 1 34 10 20 8 Light purp. viol. 5 35 31 (22-40) 37 (30-40) 7 Str. purp. viol. 2 36 22 (22-25) 23 (20-30) 7 — 2 Nr. 5 7

Observations Chromosome number Due to the high number and small size of the chromosomes in this species, an exact count using root-tip sections was found to be extremely difficult. No more than a good estimate of the number could be obtained in this way. However, with the squash technique, after pretreatment, an exact count was possible. Thus, in strains 23, 24, and 32 a somatic num ber of 84 was found (see Fig. 1). This is in agreement with previous counts; W a r b u r g 1938, G a u g e r 1937, S a n so m e 1936, S akai 1935. An estimate of the number in the rest of the strains was obtained from sectioned root tips. In all cases 2n approximated to 84 ± 4. Thus, it can be assumed that all strains have the same num ber i.e. 2 n= 84.

Fig. 1. Mitoses in root tips — squash preparations; strain 23 to the left and strain 32 to the right.

Leaf shape Photography was considered to give the most comprehensive description of variation in leaf shape. Although in certain respects a more quantitative method would have been desirable, this would have involved a rather complex series of meas­ urements. Photography was found quite sufficient for present needs and has the advantage of ease of interpretation. Figures 2, 3, 4, 5, and 6 are silhouettes showing 4 representative leaves (vertical series) of each cultivated strain, harvested from the stem at different levels from the base. There was very little intrastrain variation. These samples have been arranged in a horizontal direction with respect to the width of the leaf lobes. Although some­ times it is difficult to differentiate between samples lying close together in the series, the general trend is clear. There is no marked discontinuity in the series. Thus, in Geranium sanguineum, variation in leaf-lobe width follows a more or less continuous cline. 2* 8 Nr. 5

1__ 5cm. i I__ 5cm. i

^ s k 5# * * # % * Hk * * h $ s M # ^ 4% ^

27 25 33 26 28 17 35 12 20 36 Fig. 2. Silhouettes of 4 representative leaves (vertical Fig. 3. Silhouettes of 4 representative leaves (vertical series) from each strain. Leaf index 1. (see text). series) from each strain. Leaf index 2. (see text).

m # * % # # * * * * * * i ^ -¿i ^ q k ^ ¡ k . -Jk- JL- vjkfe iiftt * # ’ v ' V - r w

8 22 1 19 13 3 10 21 5 2 9 4 Fig. 4. Silhouettes of 4 representative leaves (vertical series) Fig. 5. Silhouettes of 4 representative leaves (vertical series) from each strain. Leaf index 3. (see text). from each strain. Leaf index 4. (see text).

* # * - * ■ * * * * # * * ^ ^ ^ j k 4 k J k * ^

16 30 7 11 15 34 Fig. 6. Silhouettes of 4 representative leaves (vertical series) from each strain. Leaf index 5. (see text). Nr. 5 9 Furthermore, it became apparent from the cultivated strains, that this cline has a geographical correlation. Plants from continental Europe show a strong tendency towards narrow leaf-lobes. On the other hand, plants from Western and Northern Europe have leaves with broader lobes (see Map, Fig. 8). This trend was also apparent in herbarium material. It was decided to use available herbarium collections to supplement the cultures; material from the British Museum, the Stockholm Riksmuseum, and the Copenhagen Botanical Museum was used. The series of cultured leaves was arbitrarily divided into 5 sections i.e. Figs. 2, 3, 4, 5 and 6, corresponding to lobe-width indices 1, 2, 3, 4 and 5 respectively. The

jjMi. ^|fc jfe jjtk . M fe ” VJH*

20 36 8 22 I 19 15

atfe JlfeWlP inn W 3 10 21 5 2 9

A jXfcte ¿Ufe ” C v “ ' 5cm>' 1

16 30 7 11 15 34 Fig. 7. Rosette leaves (see text). Numbers refer to strain nos., which are arranged in the same order as in Figs. 2-6. herbarium specimens were then scored according to this index system. The resulting indices were plotted on an outline map of Europe (see Map, Fig. 8). The general geogra­ phical arrangement of the cline becomes apparent from this map. Broad-lobed plants have a mainly north-western distribution, and narrow-lobed plants a mainly eastern one. This pattern may possibly represent a topocline but the more detailed aspects of the distribution can perhaps better be explained in terms of climatic and ecological responses. If it is assumed that broad-lobed types arc associated with coastal climatic conditions, this would explain the occurrence of broad-lobed types along the east coast of Sweden and the Mediterranean coast. On the other hand, that narrow lobes are correlated with continental conditions, would account for the exist­ ence of narrow-lobed types on the drier limestone areas of Central England, the Alvar of 01and, and Central Sweden. Local microclimatic conditions undoubtedly would also effect the distribution pattern. Nr. 5 Nr.

Fig. 8. Map showing distribution and leaf indices of cultivated strains. (Symbols used are the same as in Fig. 9). Nr. 5 Nr.

Fig. 9. Map showing distribution and leaf indices of herbarium material. 12 Nr. 5 In the Mediterranean region, particularly the Balkan peninsula, apparent anomalies occur in this pattern, these could be due to very variable local conditions, but until more material is available nothing definite can be stated. It has been noticed in the S. Alps that Geranium sanguineum is found in moister situations e. g. on north-facing slopes. The possibility of the distribution pattern being interpreted as a response to environmental conditions will be discussed more fully later.

Rosette leaves Fig. 7 shows silhouettes of one representative rosette leaf from each strain. These were harvested in Sept. 1961. The rosette leaves appear first in the spring but usually die down before June. New rosette leaves appear in the autumn and persist until killed by the first severe frosts. The rosette leaves in Fig. 7 are arranged in the same order as the stem leaf samples in Figs. 2-6. There is the same trend in leaf lobe width, although this is not as marked as in the stem leaves.

Environmental Modification of Leaf Morphology In March 1961, clonal rhizome divisions of one plant from each of 10 cultivated strains were made. 2 pieces from each clone were potted separately. One series of clonal divisions was then placed in the Cactus house and the other in the Fern house of the University Botanical Garden. These have the following characteristics: Cactus House : — Low Humidity, higher light intensity and with a large tempe­ rature amplitude between night and day. Fern House: — High Humidity, lower light intensity and with a small tempera­ ture amplitude between night and day. The plants continued to grow in these houses until September 1961, when the leaves of each strain from the 2 houses were compared. In this way it was possible to investigate the “reaction pattern” (Clausen, Keck & Hiesey 1940) in leaf mor­ phology of each individual i.e. the environmental influence when one individual is compared in different environments. Fig. 10 shows the result of these cultivations in 3 individuals which are repre­ sentative of the 10 clones. In all cases the leaves from the fern house had distinctly broader lobes than those from the Cactus House. Also leaves from the Cactus house were more hairy and smaller than their counterparts in the Fern house. Broadly the environmental conditions in the Fern and Cactus house are compar­ able with an oceanic and continental climate respectively. Thus, as regards leaf-lobe width, reaction pattern appears to parallel the inter-race pattern in Geranium sangui­ neum i.e. environmental modification simulates genetically - determinded variation. This is a situation occurring fairly commonly in plant species (e.g. T u r e s s o n 1922; B o c h e r 1944). Its importance in this particular case will be discussed later. Nr. 5 13

lunk. H k 4 ! * * * * * * * # # #

C F C r c F 26 22 Fig. 10. 3 representative clones from the greenhouse experiment (see text). Numbers refer to strain nos. C-leaves from plants grown in the cactus house. F-leaves from plants grown in the fern house.

M ü l l e r -S t o l l (1935) found the same environmental modification of leaf shape apparent in naturally occurring populations of Geranium sanguineum from the Kraichgau district of Germany.

Leaf size. T a b le III1).

Leaf Diameter cms. Leaf Diameter cms. Strain No. Strain No. Mean & Range Mean & Range

1 5.3 (5.0-5.5) 19 3.8 (3.5-4.0) 2 3.8 (3.5-4.0) 20 4.0 (4.0) 3 5.0 (4.8-5.2) 21 4.8 (4.5-5.0) 4 5.3 (5.0-5.5) 22 5.2 (5.0-5.5) 5 3.3 (3.0-3.7) 25 4.4 (4.3-4.5) 7 4.0 (4.0) 26 4.7 (4.5-4.8) 8 3.9 (3.8-4.0) 27 4.6 (4.5-4.7) 9 3.9 (3.8-4.0) 28 4.1 (4.0-4.2) 10 3.4 (3.3-3.5) 30 4.1 (4.0-4.2) 11 3.4 (3.3-3.5) 33 5.4 (5.0-5.5) 12 5.7 (5.2-6.0) 34 5.2 (4.5-6.0) 13 4.2 (4.0-4.5) 35 4.4 (4.2-4.4) 14 3.9 (3.8-4.0) 36 5.9 (5.8-6.0) 15 4.9 (4.8-5.0) 16 3.3 (3.2-3.5) 17 4.8 (4.5-5.0)

x) Measurements are taken from the 3 lowest stem leaves on each shoot. 14 Nr. 5 Size of the mature stem leaves varies considerably within each plant; there is a gradual decrease in size with increasing height of insertion on the stem. For this reason exact comparisons of leaf size between strains is difficult. However, by taking the diameter of the 3 lowest on each shoot a reasonable interstrain comparison can be made (see Tabel III). From table III it is seen that the variation in leaf size is clinal, but with some ecological correlation. The Danish strains from open coastal habitats tend to have small leaves. This trend was also apparent in the herbarium collections, where var. postratum was more or less uniformly small-leaved. The Danish strains from shaded wood margin localities (Nos. 4, 12, 15) are large-leaved. Thus, at least in Danish material, Geranium sanguineum shows a leaf size cline from open coastal habitats to shaded woodland ones. This is essentially the same situation as found in Veronica officinalis. In the greenhouse experiment, the leaves from the Cactus house were in all cases smaller than their counterparts from the Fern House. The difference sin condi­ tions between the 2 greenhouses are in some ways comparable with those between open and shaded habitats. Thus, here again, environmental modification apparently parallels genetic differences. Leaf size in the extra-Danish strains is very variable, but because of insufficient material and ecological information it is thought inadvisable to make any comparisons.

Petal Morphology Flowers of the cultivation strains were harvested at the same time as the leaf samples. A sample of 5 flowers per plant was taken. Petals of these flowers were then measured for length and breath, and the mean length/breadth ratio calculated for each strain. Within each strain the ratio is very uniform, but between strains it is very variable. This variation in petal shape shows a degree of correlation with the variation in leaf-lobe width. Plants with broad-lobed leaves tend to have broad petals, and narrow-lobed leaves to have narrow petals (See Fig. 11). This state of affairs is not unexpected from what is known of pleitropic effects. That is, the same gene system affects simultaneously leaf and petal morphology. This condition is already known from Anderson & de W inton’s (1935) work on Primula sinensis. However, in general this correlation is not strong. Its weakness may be ex­ plained by a mechanism of adaptive compensation (Clausen, Keck & Hiesey 1948). The reduction in leaf-lobe breadth may be of some selective advantage to plants in more continental regions. However, the gene system favoured by this selection, also results in narrower petals (pleiotropic effect) and thus less distinct flowers. This secondary effect may be disadvantageous to this predominantly insect pollinated species. Other genes which compensate for the harmful effect will now have a high Nr. 5 15 selective value. These compensatory genes will result in an only partial correlation between leaf and petal morphology, such as is observed. In this connection it is worth noting in Fig. 11, that the curve is depressed in intermediate strains towards a low petal ratio. This is as would be expected from the postulated mechanism.

Height and Habit (See Table II) The height of cultivated plants was measured in 1953, 1960 and 1961; the 1961 measurements are of root divisions of the original plants. Although there is a large variation in height within some strains, an inter-strain pattern can be discerned.

Fig. 11. Diagram of mean petal length/breadth ratio plotted against leaf index. The petal length/breadth ratio used here is the average value of the individual strain means in each leaf index group.

Strains from coastal habitats of Western Europe are distinctly shorter than those from inland ones. These low races, however, represent only the lower end of a contin- ous height cline; they are not separated by any abrupt discontinuity from the inland types in this respect. Strains 5, 6, 7, 9, 10, 11, 13, 14, 16, 23 & 24 are typical of this coastal type. Most of these are prostrate in cultivation, but nos. 5, 9 & 10 were of low “cushion” habit. This cushion habit is probably another response to the coastal ecosystem involving shortening of internodes, as an alternative to the prostrate habit. This situation is known from other species e.g. in Solidago virga-aurea the low dense habit is found in maritime as well as montane races (personal observations, T u r e s - so n 1925; B o c h e r 1955). The inland races of Geranium sanguineum are mainly erect in habit. Thus, here, as in many other species, can be seen the beginnings of “life form” differentia­ tion (i.e. prostrate, cushion & erect habits), in response to the differing selective pressures of the various habitats in which the species occur. 16 Nr. 5 Time and length of Flowering period Results for this are based on observations made on cultivated plants in 1953, 1955, 1960, and 1961. At weekly intervals during the summer the strains were scored as to degree of flowering using an arbitrary index (see Fig. 12). Using this method, it was possible to classify the strains according to 8 main flowering period types (see Fig. 12 and Table II).

Fig. 12. Diagram showing time and degree of flowering in the cultivated strains. As shown the strains fell into 8 main flowering period types. 1-10 flowers/plant. m m 1^-20 flowers/plant. I > 20 flowers/plant. i.e. in full flower.

All West European strains are included in types 1-5. Most of the coastal strains behave as type 4 and the West Irish material is included in type 5. Flowering type 6 is characteristic of the Øland Alvar race, type 7 of the continental steppe races and type 8 of the Italian strain (No. 34). Although it is difficult to predict how the phenology of a plant will be altered when moved from its native area to that of the experimental field in Denmark, it is clear from the results that the strains from the various climatic zones do differ genetic­ ally with respect to their flowering period. Furthermore this genetically determined Nr. 5 17 variation in flowering periodicity is correlated with the prevailing climate of the regions of origin. Thus the strains from coastal and extreme western areas (Type 4 & 5) have an extended flowering period; often flowering continues into October. This extension of the flowering period is presumably facilitated by the more equable climate of these extreme oceanic regions. The W. Irish material (Type 5) resumes flowering in September after a break in August. This discontinuity can be explained by assuming that the Danish summer (higher temperature and lower humidity) is too rigorous for plants adapted to the climate of W. Ireland with its high rainfall and low summer temperature. This results in a temporary break in llowering during high summer, followed by a resumption in September when conditions are more favourable. The short llowering period of the continental, Øland and Italian strains is prob­ ably an adaption to the hot dry summer of these regions, when conditions are unfavour­ able for the plant; here, spring and early summer are the optimal periods for plant growth. The early spring of Øland is rellected in the early commencement of flowering in this race.

Longevity of the Vegetative State Based on observations made in 1960, and 1961, an inter-race pattern was noticed with regard to the length of time over which the foliage remains healthy. Two groups were recognizable: (I) N.W. European and coastal types — Strains 2, 5, 7, 9, 11, 13, 14, 15, 16, 20, 21, 22. — Foliage persistent at least until late October. (II) Continental and Southern European types — Strains 17, 25, 26, 34, 35, 36.— Foliage withers and shoots die down before early October. This difference can also be interpreted as an adaption to the climate in the same way as the flowering period.

Hairiness The degree of hairiness varied greatly between the strains under cultivation, although none were completely glabrous. Using pressings of entire shoots (made in 1961) the strains were scored for hairiness of upper and lower surfaces of the leaves, A purely arbitrary index system was used (See Table IV). The hairs on the stem are long and erect; those of the upper leaf surface are short and adpressed whilst the lower surface has long, more or less adpressed hairs restricted to the veins and leaf margin. Hairiness in leaf and stem appears to be inde­ pendently determined e.g. plants with a low leaf hair index can have a high stem hair index as in strain 15 and vice-versa as in strain 5. Biol. Skr. Dan.Vid. Selsk.ll,no.5. 3 18 Nr. 5 T a b le IV.

Leaf Index Strain No. Stem Index Leaf Hair Index Surface upper Surface lower

1 1 1 2 3 2 3 1 2 3 3 1 1 1 2 4 2 1 1 2 5 1 2 3 5 7 3 2 2 4 8 3 2 2 4 9 3 1 3 4 10 3 1 1 2 11 • 3 1 3 4 12 3 1 1 2 13 3 2 3 5 14 3 2 3 5 15 3 1 1 2 16 3 1 3 4 17 3 1 2 3 19 3 2 2 4 20 3 1 1 2 21 1 1 1 2 22 2 1 1 2 25 3 3 3 6 26 2 2 2 4 27 2 2 2 4 28 3 1 1 2 30 3 1 2 3 33 2 1 2 3 34 3 2 3 5 35 2 1 1 2 36 2 1 2 3

Index used: Increasing degree of hairness 1—v2->3. Leaf Hair Index is the sum of the indices for upper and lower leaf surface.

From Table IV it is possible to recognize some relationship between degree of leaf hairiness and the environmental conditions of the original habitat. All Danish material from open coastal habitats has a high leaf hair index. On the other hand, plants from Danish wood margins have a low leaf hair index. Thus, at least, in this material there is some correspondence between hairiness and openness of the habitat. The rest of the material is also very variable, but not comprehensive enough for any conclusions to be drawn. However, it is perhaps worth mentioning that the Irish material has a low leaf hairiness index. Stem hair index appears randomly distributed in the material investigated. Nr. 5 19

As mentioned earlier, in the greenhouse experiments, leaves from the Cactus House were consistently more hairy than those from the Fern House. The two green­ houses can be compared to an open and a shaded habitat. Thus, here again, environ­ mental modification parallels genetically-controlled variation.

Flower colour

Wanscher’s simplified colour chart (W anscher 1953) was used as the standard for determining the flower colour. The flower colour of the strains is given in Table II, column 6. Although there is considerable variation between the strains this appears to be randomly distributed. The Horticultural Colour Chart (1938-40) code-number for each colour de­ scription used in Table II is given below. Strong purplish violet H. C. C. 32 Light - - H. C. C. 33 Strong violet H. C. C. 35 Light violet H. G. C. 36 Very pale red H. C. C. 20

Discussion Over most of its range Geranium sanguineum is a plant of rather open, dry, warm habitats with base-rich soils. Broadly, it is characteristic of 2 types of natural community. (a) Open coastal areas, where it occurs on fixed dunes, stony beaches, grassy sea slopes and cliff ledges. (b) Dry, open woodland. In central and eastern Europe it is characteristic of the open Karst scrub (woodsteppe) over limestone, base-rich sand and even rock. In some regions it occurs in hanging woodland where the tree canopy is kept open by the slope; in Denmark it is frequently found on the more or less wooded sea slopes. In addition, a few other habitats can be mentioned. In highland areas of Southern Europe it is found in open, exposed habitats such as stony slopes and rock ledges, and in montane meadows. In the bare limestone area of the Burren, West Ireland, Geranium sanguineum is found growing in the crevices of the limestone pavement. It is sometimes found along wood margins and in woodland kept open by man’s activities, but it does not normally occur in man-made habitats (see Meusel 1955). Within the cultivated material of Geranium sanguineum investigated in this paper it is possible to recognize 2 race groups, corresponding to the two habitats described above. Biol. Skr, Dan. Vid. Selsk.ll.no.5. 4 20 Nr. 5 (a) Coastal race group — low with prostrate to cushion habit, 5-25 cms. high; broad leaf lobes corresponding to leaf indices 4 and 5; small (3.0-4.5 cms. diam.) and pubescent leaves; long llowering (mid-May-early Oct.) and vegetative (until late Oct.) periods (See Plate I). (b) Woodsteppe race group — erect habit, 20-45 cms. high; narrow leaf-lobes corres­ ponding to leaf indices 1 and 2; variable leaf size and pubescence; shorter flower­ ing (mid-May-mid-July) and vegetative (until early Oct.) periods. (See Plate II).

Although these two race groups are recognizable they are not distinctly separated from each other. They represent the two ends of a more or less continuous clinal system. They are connected through strains from intermediate habitats e.g. cliff ledges, wooded sea slopes, wood margins etc. In the Burren area of West Ireland, where limestone Karst occurs at the coast, the distinction between the coastal and steppe habitats disappears. Here, all transitions from the coastal race group to the wood-steppe race group are found in a small area. As yet a suitable taxonomic treatment of this type of continuous clinal varia­ tion has not been generally accepted. For this reason, it is considered inadvisable to confer taxonomic rank on these recognisable race groups, although this has been the policy with some workers (e.g. Valentine 1941, Clausen, Keck & H iesey 1940, P ritchard 1959, 1960). Subspecific and varietal ranks do not adequately express the type of variation found in Geranium sanguineum. They presuppose distinct limits which are not found here. There is great need of a generally accepted system which gives a satisfactory taxonomic treatment of clinal systems. That end of the clinal system associated with coastal habitats has been called Geranium sanguineum var. prostratum (Cav.) Pers. (see Plate I). Its main diagnostic features are its prostrate growth and broad leaf lobes. It is impossible to define the exact limits of this taxon using these two characters. However, because (in the absence of a better system) such a division may be of limited value when referring to a clinal system we have attempted to establish the distribution of this variety using the cultiva­ tion material and herbarium specimens (see Map, Fig. 13). A pink-flowered race within the coastal race group (strains 23, 24) has also received taxonomic treatment as var. lancastriense (W ith.) D ruce. This probably represents a genotypic difference of one or a few genes as apart from flower colour, var. lancastriense resembles the normal coastal races. The petals of our material are pale pink with red venation. The two recognizable race groups are almost certainly polytopic in origin i.e. they have differentiated from the biotype pool independently in more than one locality. They are the result of genotypic convergence of populations in response to similar environment conditions. In this context, the race groups correspond with T uresson’s ecotype concept. In formulating his ecotype concept T uresson (1925) used the phenomenon that the same habit-type has become differentiated within a wide variety of species in response to the same ecosystem. In fact, morphologically distinct ecotypes can Nr. 5 Nr.

Fig. 13. Map showing the distribution of Geranium sanguineum var. prostratum (Cav.) Pers. as determined from herbarium and cultivated material. 22 Nr. 5 be said to correspond to incipient life forms. As already seen, in Geranium sangui- neum habit-type differentiation has occurred along with morphological and phenolo- gical differentiation: 3 habit-types are recognizable (prostrate, cushion and erect types) but these are connected by a more or less continuous series of transitional types. This situation probably represents a relative young stage in life-form differentiation. In Sarothamnus scoparius this process has continued further to give a marked discontinuity between the prostrate ssp. maritimus, and the erect ssp. scoparius; inter­ mediate forms do exist, but these are few in number, and do not form a continuous transitional series. The closely related Silene marítima and S. cucubalus illustrate a third stage in this process. Two distinct life forms are present between which intermediates are rare, and which are distinct enough to be given specific rank. What does the distribution of the various leaf types represent in Geranium sanguineum? We can envisage several possible explanations, which will be discussed separately below. Variation in leaf shape is concurrent with ecotypic differentiation. It would there­ fore be most reasonable to assume that the different leaf types are direct genotypic adaptions to environmental conditions. That is, they are of selective advantage in the different habitats. This explanation gains more weight from the fact that the same distribution of leaf types occurs in several other species. In Danish Pimpinella saxí­ fraga P etersen (1921) found a higher percentage of broad-lobed types in Jutland whilst in more continental Zealand the narrow-lobed types predominated. Another example of this pattern is found in Californian populations of Layia gaillardioid.es (Clausen 1951). Leaves of populations from the dry inner Coast Range are more dissected than those from the more oceanic outer Coast Range. Within our own cultivations of Centaurea scabiosa, we have noticed a parallel race pattern (unpublish­ ed). Strains from Ireland, Jutland and Norway have leaves with broad segments, whereas strains from Poland, Austria and E. France have narrowly-segmented leaves. Petersen (1915, 1922) also demonstrated a parallel ecological differentiation of leaf shape in Danish populations of Anthriscus siluestris; broad-lobed leaf types are commoner on the fertile, heavy soils and narrow-lobed leaf types are commoner on the dry sandy soils. This recurring distribution pattern is possibly correlated with the water economy of plants from the different regions. Those individuals which have the mesophyll collected closely around the veins may be better able to withstand larger transpira­ tion stresses that would be met with in more continental areas (Gregor 1956). This localization of mesophyll is a well known xeromorphic trend. In Geranium sanguineum in addition to the genotypically-determined variation in leaf shape, there is also considerable phenotypic plasticity (reaction pattern) in this respect. The reaction pattern is a genotypically-conditioned phenomenon and as such will be selected for in the normal way (W addington 1961). That reaction Nr. 5 23 pattern which allows the plant to extend its tolerance range is at a selective advantage (S chmalhausen 1949; F is h e r 1960). In Geranium sanguineum, leaf shape reaction pattern simulates the inter-race pattern; there are many other examples of this peno- menon (e.g. T u r e s s o n 1922, 1925; S chmalhausen 1949). That the two patterns coin­ cide in this way can be taken as evidence that the leaf types are directly adaptive (T u r e s s o n 1922, 1925). However, more physiological work is needed before anything more definite can be stated. Although this first explanation appears the most reasonable, other possibilities cannot be excluded. The differences in leaf morphology may only be “symptoms” of direct meta­ bolic adaptions to the environment, having no survival value in themselves (M axim ov 1929; S h ie l d s 1950). This is M a x im o v ’s (1929) interpretation of xeromorphism and may give an explanation of the association in several species of narrow leaf lobes with dry habitats; all the species have undergone similar metabolic changes in response to similar environmental factors. The coincidence of inter-race and reaction patterns could be due to physiological adaption at the genotypic and phenotypic level involving the same morphological changes. These first two explanations assume that the differences in leaf shape are either adaptive in themselves, or incident upon physiological adaption. The cline of varia­ tion may be the result of adaption to a more or less continuous habitat gradient within existing populations. It could also represent introgression of the two previously iso­ lated ecotypes, each with its own distinct leaf type i.e. a narrow-lobed and a broad- lobed ecotype. This assumes that at some time in the past, factors causing spatial disjunction were operative within the species, factors which have since disappeared. Two possibilities are apparent; this disjunction could have occured during the last ice age or in post glacial times. There is good evidence (Z im m e r m a n n 1959) that the present day variation within the Pulsatilla vulgaris — grandis complex (see later) is due in part to spatial disjunction during the last ice age caused by the Mid-European Ice sheets. With the retreat of the ice there was an expansion of areas, followed by introgression, resulting in the present day situation. A parallel process could have occurred within Geranium sanguineum. An alternative situation is that the disjunction was the result of forest expansion during post-glacial times. The development of forest would have restricted Geranium sanguineum to warm, more or less open habitats. Two such habitats would be provided by the sea coasts in Western Europe (B o c h e r 1955, 1961) and the woodsteppe areas of Central and Eastern Europe (i.e. the 2 most important habitats today). During this disjunction, adaptive differentiation in leaf shape could have occurred between the two populations. With forest clearance both populations were able to extend their areas and introgression possibly occurred to obliterate the morphological dis­ continuity between the two populations. 24 Nr. 5 A further possibility is that during spatial disjunction within the species, non­ adaptive leaf differentiation occurred between the two populations, to give a narrow- lobed eastern population and a broad-lobed western one. This was followed by area expansion and introgression between the populations. In the Pulsatilla vulgaris-g ran­ dis complex the distribution of leaf-types is the reverse of that in Geranium sanguineum, Pimpinella saxífraga etc. Pulsatilla grandis with broad-lobed leaves has a more eastern distribution than P. vulgaris. In Southern Bavaria, Tirol and Upper Austria these two species are sympatric and have introgressed to give transitional types (Z im m e r m a n n 1959). That the reverse distribution pattern is found in this complex is some evidence for believing that the leaf differentiation in Geranium sanguineum etc. is non-adaptive. However, the situation in the Pulsatilla complex is rather more complex than in Geranium sanguineum. Both P. grandis and P. vulgaris are tetra- ploid but, according to Zimmermann, have arisen independently from two distinct allopatric diploid stocks. Thus, the situation in Geranium sanguineum is not directly comparable with that in the Pulsatilla complex. However, there still remains the possibility that the leaf variation in Geranium sanguineum is non-adaptive. It may represent a topocline similar to that found by G r e g o r (1939) in Plantago maritima-, here there is a clinal increase in the ratio scape-length to spike-length from W. America to N.W. Europe. This appears not to be correlated with climatic or ecological factors. However, as mentioned previously, in Geranium sanguineum the details of the distribution pattern are best explained by assuming some adaptive significance in the leaf variation. The junior author will continue studying the problem of the significance of leaf variation. Investigations of transpiration, drought resistance etc. will be under­ taken together with the effect on leaf shape of separate components of the environment, e.g. temperature, humidity, light.

Literature

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F ish e r , F. J. F. 1960. A discussion of leaf morphogenesis in Ranunculus hirtus, — N. Z. J. Sei. 3: 685-93. Ga u g e r , W. 1937. Ergebnisse einer zytologischen Untersuchung der Familie der Geraniaceae. — Planta 26: 529-531. Gregor, J. W. 1939. Experimental Taxonomy. IV. Population differentiation in North America and European sea plantains allied to Plantago maritima L. — New Phytol., 38: 293-322. Gregor, J. W. 1956. Genotypic-environmental interaction and its bearing on a practical problem of international interest. — Proc. 7th Int. Grass. Gong., 202-210. H orticultural Colour Ch a r t. 1938/40. — R. F. Wilson, London. Maxim ov, N. A. 1929. The plant in relation to water. A study of the physiological basis of drought resistance. — London, Allen & Unwin. 451 pp. (English trans. by Yapp). Me u s e l , FI. 1955. Verbreitungskarten mitteldeutscher Leitpflanzen, 8 Reihe. — Wiss. Z. Univ. Halle. Math.-Nat., 5: 297-334. Müller-S to ll, W. R. 1935. Ökologische Untersuchungen an Xerothermpflanzen des Kraich- gaus. — Z. f. Bot., 29: 161-253. P e t e r s e n , H. E. 1915. Indledende Studier over Polymorphien hos Anthriscus silveslris (L.) Hoffm. — Dansk Bot. Ark., 1: 1-150. (French summary). — 1921. Nogle Studier over Pimpinella saxifraga L. — Bot. Tidsskr., 37: 222-240. — 1922. Études ultérieures sur la polymorphie de Y Anthriscus Silvester (L.) Hoffm., — Dansk Bot. Ark., 4: 1-28. •— 1926. Über die Variation der Potentilla erecta (L.) Dalla Torre. — Bot. Tidsskr., 39: 368-374. P r it c h a r d , N. M. 1959. Gentianella in Britain.I. G. amarella, G. anglica and G. uliginosa. — Watsonia, 4: 169-193. — 1960. Gentianella in Britain. IL Gentianella septentrionalis (Druce) E. F. W arb.— Wat­ sonia, 4: 218-237. Sa k a i, B. 1935. Studies on the chromosome number in alpine plants. II. — Jap. Jour. Genet., 11: 68-73. Sa n so m e, F. W. 1936. Some experiments with Geranium species. — Journ. Genet., 33: 359-363. S chmalhausen , I. L. 1949. Factors of Evolution. — New York, Blakiston (Me Graw-Hill). 327 pp. Sh ie l d s, L. M. 1950. Leaf xeromorphy as related to physiological and structural influences. — Bot. Rev., 16: 399-447. T u r e ss o n , G. 1922. The genotypical response of the plant species to the habitat. — Ilereditas, 3: 211-350. — 1925. The plant species in relation to habitat and climate.— Hereditas, 6: 147-236. V a l e n t in e , D. H. 1941. Variation in Viola riviniana Rchb.—-New Phytol., 40: 189-209. W a d d in g t o n , C. IF. 1961. Genetic Assimilation.-— Advances in Genet., 10: 257-293. W a n sc h e r , J. H. 1953. A simple way of describing flower colours, and a flower colour chart. — Roy. Vet. Agr. Coll., Copenhagen Yearb.: 91-104. W a r bu r g , E. F. 1938. Taxonomy and relationship in the Geraniales in the light of their cytology. — New Phytol., 37: 130-159. W ood son, R. E. 1947. Some dynamics of leaf variation in Asclepias tuberosa. — Ann. Mo. Bot. Gard., 34: 353-432. Zim m erm a nn, W. 1959. Die Phylogenie der Pflanzen. II. — Stuttgart, G. Fischer. 777 pp.

Indleveret til Selskabet den 28. juni 1962. Færdig fra trykkeriet den 31. december 1962.

PLATES Plate I. Typical specimen of the coastal race group (referable to var. prostratum (Cav.) Pers.). From Hirtshals, N. Jutland, Denmark; leg. E. Warming. Plate I Plate II. Typical specimen of the woodsteppe race group. From Kbszeg (Guns), Hungary; leg. Waisbecker. Plate II

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Printed in Denmark Bianco Lunos Bogtrykkeri A/S Id: 85 Forfatter: Bocher, Tyge W., and Lewis, Martin C. Titel: Experimental and Cytological Studies on Plant Species. VII. Geranium anguineum. År: 1962 ISBN: Serietitel: Biologiske Skrifter Serienr: SB11:5 SerienrFork: SB Sprogkode: Eng