Blackwell Science, LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2003 141 Original Article L. SÁEZ and J. J. ALDASOROREVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX Botanical Journal of the Linnean Society, 2003, 141, 215–241. With 19 figures A taxonomic revision of Campanula L. subgenus Sicyocodon (Feer) Damboldt and subgenus Megalocalyx Damboldt (Campanulaceae) LLORENÇ SÁEZ1* and JUAN JOSÉ ALDASORO2 de Botànica, Facultat de Ciències, Universitat Autònoma de Barcelona. E-08193, Bellaterra, Barcelona, Spain 2Real Jardín Botánico de Madrid-CSIC. Plaza de Murillo, 2, E-28014, Madrid. Spain Received May 2002; accepted for publication October 2002 A taxonomic study of Campanula L. subgenera Megalocalyx Damboldt and Sicyocodon (Feer) Damboldt is presented here. Taxonomical, nomenclatural, morphological, chromosomal, geographic and ecological data are recorded for each taxon. Based on these data, one species is included in C. subgenus Sicyocodon and 12 in C. subgenus Megalocalyx. In addition, 11 lectotypifications of specific and infraspecific names are made, and a taxonomic key and descriptions of subgeneric and specific taxa are given. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241. ADDITIONAL KEYWORDS: endemism – Mediterranean basin – phylogeography – pollination – taxonomy. INTRODUCTION The genus Campanula comprises c. 300 species, largely distributed in temperate areas of the northern hemisphere. Campanula includes annual and perennial plants. Annuals were included in sect. Annuae by Boissier (1875), but later were separated by Damboldt (1976) into four subgenera: Megalocalyx, Sicyocodon, Roucela (Dumort.) Damboldt and Brachycodonia (Fedorov) Damboldt. The subgenus Roucela includes plants lacking appendages between lobes of the calyx (Damboldt, 1978), and the monotypic subgenus Brachycodonia holds an intermediate position between Campanula and Legousia (Federov, 1957). Subgenus Sicyocodon and Megalocalyx are closely related and form a natural group. They have two synapomorphies in common: (i) the dichasial or subdichasial inflorescence, and (ii) the reflexed appendages present between the calyx lobes (Damboldt, 1976). These subgenera have been recognized by different authors as several other taxonomic ranks: as a subsection *Corresponding author. E-mail: llorens.saez@uab.es (Fedorov, 1957; Rechinger & Schimann-Czeika, 1965), or as a section (Post & Kuntze, 1904; Charadze, 1949). The monotypic subgenus Sicyocodon can be easily differentiated from taxa belonging to subgenus Megalocalyx on the basis of the conspicuous, broadly campanulate corolla and very long exserted style (35– 50 mm long). Unfortunately, the reported taxonomic affinities of subgen. Sicyocodon are speculative. On the other hand, plants included in subgenus Megalocalyx show tubular-campanulate or nearly rotate corollas, the style is smaller (4–15 mm long) and always included in the corolla. Some authors (Contandriopoulos, 1972, 1976, 1984; Gadella, 1966; Damboldt, 1978) suggested that Sicyocodon with its long exserted style probably holds a very isolated position in the genus, which is attributed to a different pollination syndrome related to flies, unique in the Campanulaceae (Damboldt, 1978). Plants included in the genus Campanula, and particularly those in subgen. Megalocalyx, are mainly pollinated by bees that become coated with pollen when they take the nectar. When the bee visits a flower in female-phase, the pollen is deposited on the stigma while the bee searches for nectar. Other nonspecialized pollen-feeding insects were observed, such as small flies and beetles. Nevertheless, they are less © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 215 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 1Unitat 216 L. SÁEZ and J. J. ALDASORO pared. All taxa were then differentiated using qualitative characters. This separation was tested by means of Discriminant Analysis (DA) for the most difficult species. This method (Sneath & Sokal, 1973), which requires the a priori assignment of OTUs to groups, indicates whether the recognized groups are statistically definable entities or whether there is a too much variation within groups to allow classification. For DA, a raw matrix was obtained, the results sorted into discrete groups, and calculations carried out using the STATISTICA package. The parameters used as descriptors and the results are summarized in Tables 1 and 2. Both quantitative and qualitative Table 1. Correct classifications and values of P obtained in DA C. dichotoma C. hierosolymitana C. strigosa C. saxonorum C. propinqua C. reuteriana C. camptoclada Number of OTUs Correct classifications (%) P 25 9 17 7 16 8 10 100 100 100 100 100 100 100 0.271 0.097 0.184 0.076 0.173 0.086 0.108 Table 2. Standardized coefficients obtained in DA for canonical variables MATERIAL AND METHODS This revision is based on 423 herbarium specimens, including type specimens, from the following herbaria (abbreviations according to Holmgrem, Holmgrem & Barnett, 1990): BC, BCC, BCF, COI, G, LD, LE, MA, MAF, MPU, P, RAB, UPS, VAB, W and the herbarium of the Universitat de les Illes Balears (Balearic Islands, Spain). Features of gross morphology were studied under a binocular stereoscopic microscope. Samples for scanning electron microscopy (SEM), were glued to aluminium stubs, coated with 40–50 nm gold and examined with a JEOL-TSM T330A at 20 kV. The density of spinules in the pollen surface was recorded by counting the number of spinules in a known area of surface of a SEM photograph. Twelve quantitative characters were recorded and measured using a Brown and Acutee 599-571-3 digital caliper. Each character was analysed for its mean and median values, range, standard deviation and significance, using the STATISTICA package. To represent the variability of each descriptor within species, boxplots containing medians and percentiles were pre- Ratio leaf width/leaf length Calyx length during fructification Calyx appendage length during fructification Corolla length Ratio lobe length/total corolla length Length of the basal part of the stamen filament Length of the narrow part of the stamen filament Anther length Style length Stigma width ratio seed width/seed length Eigen values Total Cumulative Proportion Root 1 Root 2 -0.2191581 -0.10640835 0.14559311 -0.26435199 0.16281492 0.28133583 0.23898034 0.11932179 0.02054952 0.45656919 0.43252316 -0.20315267 -1.14482856 0.54066497 0.23132282 -0.10097335 0.68195915 0.50054479 0.42487493 -0.04147021 -0.18845785 0.5991323 41.5903473 0.79949468 5.16856527 0.89885038 © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 important as pollination agents. In the absence of insects, autopollination is common, being probably more frequent in annual plants with small flowers (Nyman, 1993; Proctor, Yeo & Lack, 1996). On morphological grounds, subgen. Sicyocodon is related to subgen. Megalocalyx with which it shares most of the vegetative characters and the basic flower features. However, this could be simply due to a convergence. In the absence of more conclusive evidence (molecular or anatomical) for subgeneric delimitation in the genus, we prefer to accept subgenus Sicyocodon as independent of Megalocalyx. Subgenera Sicyocodon and Megalocalyx are distributeed throughout the Mediterranean basin, but the number of species is higher in the east Mediterranean. The limits are the Canary Islands in the west and the Caucasus in the east. The annual habit and the irregular reproductive patterns could be responsible for the observed morphological variation of some species. These species can have autogamous and allogamous flowers during different phases of their life cycle. Taxonomic difficulties sometimes led to different and somewhat confusing treatments in Floras and other local studies (Battandier & Trabut, 1904; Fomin, 1907; Maire, 1932; Rechinger & Schimann-Czeika, 1965; Damboldt, 1978; Greuter, Burdet & Long, 1984). Thus a comprehensive study throughout the distribution area is necessary to understand the group. The object of this work is to obtain comprehensive information about these species, to discover useful characters to define them taxonomically, and to define their geographical distribution. REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX characters were used in the key, the most discriminant quantitative characters being inferred from box-plots. RESULTS TAXONOMIC CHARACTERS Inflorescences, flowers and pedicels: Inflorescences can vary between dichasia and monochasia, with many intermediate forms, but the commonest form is a dichasium. In some cases, the monochasial or dichasial development seems to depend on actual growth conditions. The exceptions are C. camptoclada, which always has monochasia, and C. rimarum, with solitary flowers. Flowers are usually erect but in the case of C. rimarum, they are pendulous. However, all the species have reflexed pedicels during the fruiting period. The indumentum of pedicels is useful to discriminate species because, as previously noted, they can show one or two types of hairs (Figs 1,2). Calyx: The calyx is five-lobed, deeply divided, with reflexed appendages located between the adjacent lobes. The lobes are ovate, ovate-lanceolate or narrowly lanceolate, but in C. strigosa the lobe apex is subulate. The length of lobes is a useful discriminating character: they are longer in C. cecilii, C. reuteriana and C. macrostyla and shorter in the remaining species. In many cases, the calyx is accrescent and the appendages are connivent during fruiting, especially in C. strigosa and C. reuteriana. One species shows a different type of calyx development: C. stellaris has fruit with a stellate calyx. Corolla: The corolla is considerably variable in shape, varying from tubular-campanulate, campanulate, broadly campanulate, to nearly rotate. The length is also variable, from 4 to 5.5 mm long in C. rimarum to c. 30(40) mm long in C. dichotoma and C. macrostyla. Most species have lobes dissected from only a third to a half of the corolla length, but C. semisecta and C. stellaris have lobes longer than one half of the corolla length and in C. rimarum they are shorter than a quarter of the total corolla length. The corolla colour is violet or violet-blue in subgen. Megalocalyx, with two exceptions: yellow in C. sulphurea, and reddish to violet in C. strigosa. The colour in subgen. Sicyocodon (C. macrostyla) is purple-whitish outside, and inside it is purple with darker veins. Androecium: The filaments show two parts: a triangular-ovate (rarely obovate, elliptical or subpentagonal) base and a filiform apex. The base is ciliate in most species, showing 0.1–0.2 mm long hairs, but in C. rimarum the base margin is papillate. Campanula macrostyla has long hairs (c. 1 mm) in the margin. The abaxial surface can also bear hairs, as occurs in C. camptoclada, C. cecilii and C. macrostyla. The filiform part of the filament varies in length, from 0– 0.5 mm in most species, to 0.5–2 mm in C. dichotoma, C. macrostyla, C. saxonorum and C. cecilii. The anthers are also considerably variable in size, from 3–4.5 mm long in C. stellaris, to 7–7.5 mm in C. macrostyla. Pollen is spherical, triporate (rarely tetraporate), 18– 30 mm in diameter, and has a surface ornament of more or less crowded spinules, which are more or less prominent (Figs 3,4). The density of spinules in the pollen surface was recorded, and varies from 0.24 spinules mm-2 to 2.68 spinules mm-2. As occurs in other Campanula species, anthers dehisce while flowers are still in bud, depositing the pollen on pollen-collecting hairs of the upper style (Fig. 5). Gynoecium: The style is included in all species of subgen. Megalocalyx (rarely subexserted in C. propinqua) and prominently exserted in subgen. Sicyocodon. The stigma is trilobate in all species of both subgenera. Campanula macrostyla is characterized by its larger stigma (12–16 ¥ 3.5–5 mm) with patent lobes at maturity. The pollen-collecting hairs of the style accumulate pollen (Nyman, 1992a,b, 1993). Later, pollen is retrieved by pollinators and the hairs usually retract into their expanded bases (Fig. 6). However, there are two species (C. macrostyla and C. cecilii) which show a weak or very late invagination of these hairs (Fig. 5). Capsules and seeds: The fruit of both subgenera is a 3-locular, pendulous capsule, dehiscing by three basal valves. The capsules are usually hispid, with indumentum distributed on the keels or on the valves. The indumentum is often deciduous, the mature fruit being glabrous or glabrescent. The seeds are ±broadly elliptical, with the surface smooth and shining, the © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Habit and leaves: Annuals, showing generally erect stems (the exception is C. rimarum which is flexuous and filiform). Stem simple, often dichotomously branched. Leaves sessile or subsessile, obtuse. The shape varies between lanceolate, elliptical, ovate and oblong-lanceolate, with a reticulate venation. The leaf margin is entire, crenate or finely denticulate. The indumentum is hispid in leaves and stems of both subgenera, showing rigid unicellular eglandular hairs (0.5)1–2.2(2.8) mm long. Hair density varies considerably among species included in subgen. Melgalocalyx: it is dense in C. hierosolymitana, and usually sparse in C. dichotoma. A few species possess two types of hairs in pedicels, calyx, capsule and higher parts of stems: longer patent hairs and shorter more retrorse ones. This feature is useful in distinguishing some species. Lower parts of stems and leaves show only one type of hair. The hair surface is generally granulate (Fig. 1). 217 218 L. SÁEZ and J. J. ALDASORO colour yellowish to brown. The seeds are rather similar in length (0.7–1.1 mm long), only C. cecilii having longer seeds (1.2–1.5 mm long). Chromosome number: Subgenus Megalocalyx shows several basic numbers: x = 8, 10, 11, 12 (Contandriopoulos, 1972), while subgen. Sicyocodon has x = 10 (Marchal, 1920; Contandriopoulos, 1972; Damboldt, 1978). Processes of dysploidy could explain these numbers (Contandriopoulos, 1972), considering that x = 17 occurs frequently in Campanula (Gadella, 1966). B chromosomes have been reported in two species, C. reuteriana and C. rimarum (Contandriopoulos, 1972). MORPHOMETRIC ANALYSIS As previously mentioned (see Material and methods), DA was used to test the differentiation made using qualitative characters. The group of more similar species (C. dichotoma, C. hierosolymitana, C. strigosa, C. saxonorum, C. propinqua, C. reuteriana and C. camptoclada) was analysed and the results are presented in Figure 7 and Tables 1 and 2. The descriptors used are: width/length ratio of leaves, calyx length during fructification (Fig. 8), calyx appendage length during fructification, corolla length (Fig. 9), ratio lobe length/total corolla length (Fig. 10), length of the basal part of the stamen filament (Fig. 11), length of the © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Figures 1–6. Scanning electron micrographs of different parts of Campanula subgen. Megalocalyx. Fig. 1. Indumentum of pedicels of C. saxonorum showing two hair types (Haussknecht 2600, MA 628444). Fig. 2. Indumentum of pedicels of C. strigosa showing only one hair type ( Dinsmore 6014, MA 120925). Fig. 3. Pollen of C. macrostyla (unknown origin, MA628444). Fig. 4. Pollen of C. propinqua (Magmanesian s.n., MA 560705). Fig. 5. Pollen-collecting hairs of the upper style in C. macrostyla (unknown origin, MA628444). Fig. 6. Retracted pollen-collecting hairs in the upper style in C. saxonorum (Haussknecht 2600, MA 628444). REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX 219 fructification, appendage length during fructification, corolla length, ratio lobe length/total corolla length, length of the basal part of the stamen filament, length of the narrow part of the stamen filament, anther length, style length, stigma width and ratio seed width/seed length. narrow part of the stamen filament (Fig. 12), anther length (Fig. 13), style length, stigma width and ratio seed width/seed length (Fig. 14). The least discriminating among them were: width/length ratio of leaves, calyx length during fructification, appendage length during fructification, and style length. The rest of characters were sufficiently discriminant (Tables 1,2). The calyx is longer in several species. The longest are C. cecilii, C. strigosa, C. saxonorum, C. sulphurea, C. reuteriana and C. macrostyla (Fig. 8), but they are more acrescent in C. strigosa and C. reuteriana. In C. cecilii and C. macrostyla they suffer a proportionally weaker upgrowth during fructification. The longest corollas occur in C. cecilii, C. strigosa, C. reuteriana and C. macrostyla (Fig. 9). However, only two species have the lobes deeply dissected, C. stellata and C. semisecta (Fig. 10). As shown in the box-plot (Fig. 12), the filiform part of the filament is very useful in differentiating C. dichotoma, C. cecilii, C. saxonorum and C. propinqua from all other species of subgen. Megalocalyx. The length of the basal part of the stamen filament differentiates C. propinqua, with lower values from all others (Fig. 11). The anthers are longer in C. semisecta, C. strigosa, C. sulphurea and C. saxonorum, and shorter in the others (Fig. 13). All these values are considerably higher in C. macrostyla due to the fact that this species possesses broad flyattracting flowers (Damboldt, 1978). The seeds are wider in C. strigosa and C. reuteriana, producing a nearly rounded shape and higher width/length ratios (Fig. 14). GEOGRAPHIC DISTRIBUTION The species of Campanula subgen. Megalocalyx are native to western Asia, north Africa, south and west Europe and Macaronesia (Canary Islands). Many species are endemic to relatively small areas, with the exception of C. dichotoma and C. propinqua (Figs 15– 19). The centre of diversity lies in western Asia, mainly the area formed by Anatolia, Caucasus, Syria and Palestine (Fig. 15). The coast of Syria, Lebanon and Palestine and many zones of the interior of Anatolia are extraordinarily rich in species of C. subgen. Megalocalyx (Figs 15–19). The monotypic subgenus Sicyocodon is endemic to south-western Anatolia (Fig. 15). Conversely, only two species grow in the western Mediterranean, C. semisecta and C. dichotoma. Campanula semisecta is endemic to the south-east of Iberian Peninsula, while C. dichotoma is widespread in Macaronesia, north Africa, Italy, Sicily and several other Mediterranean islands (Figs 16,17). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Figure 7. Plot of Discriminant Analysis of the group of C. dichotoma, C. hierosolymitana, C. strigosa, C. saxonorum, C. propinqua, C. reuteriana and C. camptoclada. The descriptors used are: width/length ratio of leaves, calyx length during 220 L. SÁEZ and J. J. ALDASORO Fig. 9 Fig. 10 Fig. 11 Fig. 12 Fig. 13 Figures 8–14. Box-plots representing the variability of descriptors in all Campanula species studied. Fig. 8. Calyx length in fruit. Fig. 9. Corolla length. Fig. 10. Ratio of lobe length/total corolla length. Fig. 11. Length of the basal part of the stamen filament. Fig. 12. Length of the narrow part of the stamen filament. Fig. 13. Anther length. Fig. 14. Ratio of seed width/seed length. All measurements are in mm. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Fig. 8 REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX 221 ¥ 10 Fig. 14 Figure 15. Number of species of Campanula L. subgenera Sicyocodon (*) and Megalocalyx in the east and west of the Mediterranean Basin. TAXONOMIC TREATMENT Campanula subgen. Sicyocodon (Feer) Damboldt, in Notes Roy. Bot. Gard. Edinburgh 35:43 (1976). ∫ Sicyocodon Feer in Bot. Jahrb. Syst. 12: 614 (1890). ∫ Campanula sect. Sicyocodon (Feer) Kuntze in T. Post & Kuntze, Lex. General Phan. 95 (1904). Description: Annual, subdichotomously branched, rarely simple. Leaves ovate-lanceolate, entire or nearly so, alternate. Flowers usually terminal. Calyx © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Figures 8–14. Continued 222 L. SÁEZ and J. J. ALDASORO Figure 17. Area of distribution of C. dichotoma, C. reuteriana and C. hierosolymitana in the Mediterranean Basin. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Figure 16. Area of distribution of C. cecilii, C. semisecta, C. macrostyla and C. stellaris in the Mediterranean Basin. REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX 223 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Figures 18, 19. Area of distribution of Campanula species in the east of the Mediterranean Basin. Fig. 18. C. camptoclada, C. saxonorum and C. strigosa. Fig. 19. C. sulphurea, C. propinqua and C. rimarum. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 224 L. SÁEZ and J. J. ALDASORO KEY TO THE SPECIES 8¢¢. 9. 9¢¢. 10. 10¢¢. 11. 11¢¢. 12. 12¢¢. Style 35–50 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. C. macrostyla (subgen. Sicyocodon) Style 4–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2. (subgen. Megalocalyx) Corolla divided more than half of its length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Corolla divided less than half of its length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Calyx appendages spread during fructification, with a stellate aspect. . . . . . . . . . . . . . . . . . . . . . . . . . 2. C. stellaris Calyx appendages not spread during fructification, not stellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. C. semisecta Anthers 1–2 mm long. Corolla usually 4–5.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. C. rimarum Anthers longer than 3 mm. Corolla usually longer than 6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Corolla yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. C. sulphurea Corolla blue, violet or pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Lobes of calyx ending in a ±subulate apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Lobes of calyx acute, not ending in a subulate apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Subulate apex on calyx lobes up to 2 mm long. Seed width/length 0.5–0.7 . . . . . . . . . . . . . . . . . . . .6. C. saxonorum Subulate apex on calyx lobes up to (3)5–10 mm long. Seed width/length 0.8–0.9 . . . . . . . . . . . . . . . . . 7. C. strigosa Pedicels of flowers and base of calyx with two types of hairs: some shorter, retrorse and others longer, patent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Pedicels of flowers and base of calyx with only a kind of hairs, all similar and patent . . . . . . . . . . . . . . . . . . . . . 11 Appendages of calyx 5.5–7 mm long during fructification. Corolla 20–30 mm long. . . . . . . . . . . . . . 8. C. reuteriana Appendages of calyx 2–4 mm long during fructification. Corolla 9–20(23) mm long . . . . . . . . . . . . . . . . . . . . . . . . 10 Stamens with a very short (0.1 mm long) filiform part or without it. Stigma 2–3 mm long . . . . 9. C. camptoclada Stamens with a filiform part 0.2–1 mm long. Stigma 1–1.5 mm long. . . . . . . . . . . . . . . . . . . . . . . . 10. C. propinqua Stamens with a filiform part 0–0.2 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. C. hierosolymitana Stamens with a filiform part longer than 0.4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Lobes of fruiting calyx 5.5–15 mm long. Seeds 0.7–0.9 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. C. dichotoma Lobes of fruiting calyx longer than 15 mm. Seeds 1.2–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. C. cecilii divided nearly to base, accrescent after anthesis, with reflexed appendage at each sinus between adjacent lobes. Corolla large, very broadly campanulate. Style very long-exserted. Stigma spindle-like, spreading in 3 long linear-oblong segments. Capsule 3-locular, dehiscing by three basal valves. Type: C. macrostyla Boiss. & Heldr. in Boiss. 1. Campanula macrostyla Boiss. & Heldr. in Boiss., Diagn. Pl. Orient. Ser. 1, 11: 65 (1849) ∫ Sicyocodon macrostylus (Boiss. & Heldr.) Feer in Bot. Jahrb. 12: 614 (1890) Ind. loc.: ‘Hab. in glareosis ad littora lacûs Egirdir Pisidiae et in saxosis ad Ermenek Isauriae. (Heldr.). Fl. Julio ineunte.’ Lectotype: (designated by Damboldt, 1978: 52): [Turkey, Isparta], in glareosis ad littora lacus Egirdir, 3 lieues d’Egirdir sur la route de Kovich, 4.vi.1845, Heldreich s.n. (G). Description: Annual 15–35(70) cm, hispid. Stem erect to subflexuous, simple or branched from base. Leaves 30–65 ¥ 10–20 mm, scattered, sessile, entire or nearly so, ovate-lanceolate, acute or subacute; upper cordate and auricled at base, reflexed. Flowers solitary, terminal or axillary on stout pedicels. Pedicels hispid, hairs 2–3 mm long. Calyx 22–29 ¥ 5–6 mm (25–28.2 ¥ 7.2– 8.7 mm in fruit), hispid, hairs 0.7–2 mm long; lobes 17–23 mm long (19–32 mm long in fruit), ovate-lanceolate, acute, hairy at margin and on the nerves; appendages 4.7–6 mm long (5–9 mm long in fruit), rounded to ovate, obtuse, concealing ovary in fruit. Corolla 22–32(40) mm, divided to c. 1/5, broadly opencampanulate, glabrous or hairy towards base, outside whitish to purple, purple within with violet veins; lobes 6.5–11 ¥ 8–15 mm, broadly triangular, acute; tube 15–17 mm long. Stamen 13–14.5 mm long; filiform part of filament 1.5–2 mm long; base 4.2–5 ¥ 4– 5 mm, triangular-ovate to broadly obovate, densely hairy on the abaxial surface and on the margin, hairs 0.7–0.9(1) mm long; anthers 7–7.5 mm long; pollen 19–25 mm diameter, with a low density of spinules (0.22–0.31 spinules mm-2). Style 35–50 mm long, long exerted, straight. Stigma 12–16(20) ¥ 3.5–5 mm. Capsule (7.5)11–14 ¥ (6)12–15 mm, concealed by the acrescent connivent calyx lobes and appendages, sparsely hairy on the keels (hairs 0.7–2 mm long). Seeds 1–1.2 ¥ 0.7–0.9 mm, light brown. Chromosome number: n = 10 Contandriopoulos, 1972). (Marchal, 1920; Area: South Anatolia: Konya, Ermenek. Habitat: Scrubs, stony places, 600–1400 m. Phenology: Flowering June Illustration: Hooker (1878; table 6394), Feer (1890; table VIII), Damboldt (1978: 51, fig. 3). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 1. 1¢¢. 2. 2¢¢. 3. 3¢¢. 4. 4¢¢. 5. 5¢¢. 6. 6¢¢. 7. 7¢¢. 8. REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX Representative specimens examined: TURKEY: In saxosis Ermenek, 2.vii.1845, Heldreich (G); Ad lacum Egirdir, 1854, Tchiatchef s.n. (G); Ermenek, vii.1872, A. Peronin s.n. (P); 50 km W Mirh, 1000 m, 7.vi.1966, Damboldt 66-29-7 (W); oberhalp d. Apa Baraji, 1000 m, 11.vii.1978, Sorger, 78-25-6 (W); Konya: Konya-Seydisehir, 2 km SW Karacaören, 1300 m, 27.vii.1981, Nyedgger 16997 (G); Ermenek, Cilicia, 1290 m (G); Cilicie, 1874 (MPU). Unknown origin: ex Hort. Paris. Herbier Delacour s.d. (MA 628481). 225 obovate to subpentagonal, sparsely hairy in the upper margin, hairs 0.1–0.2 mm long; anthers 3–4.5 mm long; pollen 19–23 mm diameter, with a low density of spinules (40–47 spinules mm-2). Style 6–12 mm long, ±included. Stigma 1.9–2.7 ¥ 0.9–1.5 mm. Capsule 5– 6.5 ¥ 4–6 mm, surrounded by the spreading calyx, hairy on the keels (hairs 0.8–1.5 mm long) and sometimes on the valves (hairs 0.5–0.8 mm long), irregularly dehiscing. Seeds 0.8–1 ¥ 0.3–0.5 mm, yellowish. Chromosome number: Unknown. Description: Annual herbs, subdichotomously branched, rarely simple. Leaves ovate, elliptical or oblong-lanceolate, entire or nearly so, alternate. Flowers usually in shortly pedunculate dichasia. Calyx divided nearly to base, accrescent after anthesis, with a reflexed appendage at each sinus between adjacent lobes. Corolla tubular-campanulate, broadly campanulate or nearly rotate, usually blue or violet, rarely yellow or purple. Style included. Stigma-lobes 3. Ovary growing upward after anthesis. Capsule 3-locular, pendulous, dehiscing by three valves, rarely dehiscing irregularly. Seeds c. 1 mm, pale yellow, shining. Type: C. strigosa Banks & Sol. 2. Campanula stellaris Boiss., Diagn. Pl. Orient. Ser. 1, 11: 63 (1849) Ind. loc.: ‘Hab. in lapidosis dumosis montis Carmeli paulò supra conventum. Legi fructif. Maio 1846.’ Lectotype: (designated by Damboldt, 1978: 50): Palestine, Mt. Carmel, vs. 1846, Boissier s.n. (G). Isolectotypes K, G, P, W. Description: Annual 3.5–26 cm, hispid, greyish. Stem erect or flexuous, simple or dichotomously branched form base or in the upper part. Leaves 5–30 ¥ 3– 11 mm, usually sessile, lower sometimes petiolate, entire or nearly so, ovate to elliptical, obtuse. Flowers in shortly pedunculate dichasia. Pedicels hispid, hairs 0.4–0.9 mm long. Calyx 5–13.1 ¥ 2–2.4 mm (9.6– 14.3 ¥ 2.5–4.2 mm in fruit), hispid, hairs 0.5–1 mm long; lobes 3.5–9.1 mm long (7.1–10.2 mm long in fruit), oblong-lanceolate, acute; appendages short, 1.1– 3.5 mm long (2.5–4.1 mm long in fruit), narrowly triangular, acute, usually concealing ovary in fruit; fruiting calyx stellately spread, accrescent, reaching up to 25 mm diameter. Corolla 7–16.5 mm long, deeply lobed (to its middle or beyond), broadly campanulate, nearly rotate, glabrous, violet to violet-blue, tube white; lobes 5.5–11.5 mm long, elliptic-lanceolate, obtuse or subacute; tube 2.4–4.7 mm long. Stamen 5–7.1 mm long; filiform part of filament absent; base 2–3 ¥ 1–2 mm, Area: South-eastern Turkey, Iraq, Syria, Palestine and Lebanon. Habitat: Scrub, rocky and sandy places. 200–1675 m. Phenology: Flowering February – May. Illustration: Feinbrun-Dothan (1977: 476). Representative specimens examined: IRAQ: Gadir, Siria, 23.iv.1880, Peyron s.n. (G). LEBANON: S Liban, Ras el Bayada, S de Tyr, 8.iv.1957, Pabot s.n. (G); S, Baabda, près Beyrouth, 200 m, 2.v.1957, H. Pabot s.n. (G); Devi el Kamar, SE Beyrouth, 28.iv.1957, Pabot s.n. (G); Ain Hesban, Palestine, 660 m, 27.iv.1911, Meyers & Dinsmore 848 (G); Beyrouth, 8.iv.1889, Vicent s.n. (MPU). PALESTINE: Mt. Carmel, iv.1846, Boissier s.n. (G, W); Zerka to Jbel Atturus Transjordan, 26.iv.1945, Davis 9381 (G); Tiberias-Migdal, dry slopes, 3.iv.1942, Davis 4253 (W); Wadi Jhannam, Nahr Michmich, 14.vi.1946, Mouterde 8572 (G); Jerusalem, 1894, Makowski s.n. (W); Nahr el Kelb, 18.iv.1948, Mouterde 9237 (G); Kinnrot Valley, upper Jordan Valley, 2 km NE of Kibbutz Haen, 2.iv.1989, Damin 39045 (G). SYRIA: cultures sablonneuses à Abarouch, près Saïda, 6.iv.1853, Blanche s.n. (MPU, P, W); champs au dessous de Bacamie a l’E de Saïda, 20.iv.1858, Gaillardot s.n. (MPU); Saïda, 21.iv.1880, Barbey s.n. (G); Beryti, 8.v.1890, Vincent s.n. (MPU); Collines du Liban à l’Est de Saïda, 18.iv.1818, Gaillardot s.n. (G); M. Libani prope Sidonem, s.d., Gaillardot, s.n. (W). TURKEY: Sehch Meram, monte Nur, iter Cilicico-Kurdico, 29.iv.1859, Kotschy 21 (W); Syriae borealis, mont Carsuis, 4–5500¢, VI-1909, Haradjan 3128 (G); prov. Hatay, dist. Iskenderum, 7 miles N of Iskenderum, 21.iv.1957, Davis & Hedge 26945 (G); Camaine, Toprakkale, bassalt gulley below the castle, 20.iv.1957, Davis & Hedge s.n. (W); Antalya, 49 km W Anamur, 14.iv.1985, Sorger 85-59-3 (W). 3. Campanula semisecta Murb. in Acta University Lund. 33(12): 115 (1897). ∫ C. dichotoma var. semisecta (Murb.) Pau in Mem. Soc. Esp. Hist. Nat. 12: 357 (1924). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Campanula subgen. Megalocalyx Damboldt in Notes Roy. Bot. Gard. Edinburgh 35: 43 (1976). 226 L. SÁEZ and J. J. ALDASORO Ind. loc.: ‘Connu jusqu’ici seulement en Espagne. Albacete: Broussailles à Riopar (Bourg. 1850) Jaen: Cerro de San Vicente; Puerta (Blanco, 1849).’ Lectotype: (here designated) [Spain] Broussailles à Riopar, 4.vii.1850, Bourgeau 779 (LD). Isotypes MA, P, G. Description: Annual up to 45 cm, hispid. Stem erect or flexuous, dichotomously branched from base or in the upper part. Leaves 6–21 ¥ 2–14 mm, usually sessile, lower sometimes petiolate, entire or nearly so, ovate to elliptical-lanceolate, acute or obtuse. Flowers in shortly pedunculate dichasia. Pedicels hispid, with adpressed hairs 0.05–0.25 mm long and patent hairs 0.7–2 mm mm long. Calyx 5–16 ¥ 1.3–1.5 mm (7– 13.4 ¥ 2–2.3 mm in fruit), hispid, hairs 0.8–1.9 mm long; lobes 4–11 mm long (9.3–15 mm long in fruit), oblong-lanceolate to lanceolate, acute; appendages 2.1–2.7 mm long (2.2–3 mm long in fruit), narrowly triangular, acute, usually concealing ovary in fruit. Corolla 5–25 mm long, deeply lobed (to its middle or beyond), broadly campanulate, glabrous or hairy in the abaxial side, violet to violet-blue; lobes 4–12 mm long, elliptical-lanceolate, obtuse or subacute; tube 7– 8 mm long. Stamen 5–9 mm long; filiform part of filament 0–0.3 mm; base 1.5–2.5 ¥ 2–3.1 mm, broadly ovate, sparsely hairy in the upper margin, hairs 0.2– 0.3 mm long; anthers 3.5–7 mm long; pollen 20–25 mm diameter, with a high density of spinules (2.5–3.1 spinules mm-2). Style 4.5–15 mm long, included. Stigma 1.1–2.5 ¥ 1.2–1.4 mm. Capsule 5.5–8 ¥ 5.5– 7 mm, with patent hairs on the keels (0.7–2 mm long) and adpressed hairs on the valves (hairs 0.05– 0.25 mm long). Seeds 0.9–1.1 ¥ 0.3–0.6 mm, yellowish. Chromosome number: Unknown. Area: South-eastern Iberian Peninsula. Habitat: Scrub, rocky and sandy places. 80–1700 m. Phenology: Flowering May – July Illustration: Sáez & Aldasoro (2001: 128) Notes: There is some confusion about the distribution of this species. There are several herbarium specimens collected in north Africa and determined as Representative specimens examined: SPAIN: Albacete, Sierra de Taibilla, near Nerpio, 1000 m, 26.vi.1988, Valdés et al. s.n. (G); Albacete, Santa Elena de Ruidera, 25.v.1934, González Albo s.n. (MAF 2780); Albacete, Broussailles à Riopar, 4.vii.1850, Bourgeau 779 (G, P); Alicante, in monte Mongó vs. Denia, 1.vi.1885, Pau s.n. (MA 120999); Alicante, Montcabrer, 30SYH19, 1250 m, 19.vi.1988, J.R. Nebot s.n. (VAB 92/0809); Alicante, Vall de Gallinera, pic del Miserat, 30SYJ4402, 550 m, 8.vi.1997, M. Signes & J.X. Soler JS7098 (MA 590245); Castellón, Mas de Moro, Amara, Segorbe, 1885, Pau s.n. (MA 120958); Ciudad Real, Lagunas la Tomilla y San Pedro, 29.v.1934, González Albo s.n. (MA 120950), 25.vi.1935, González Albo s.n. (BC 84827); Cuenca, Contretras, vi.1899, Pau s.n. (MA 120956); Jaén, cerro de San Vicente, 1849, P. Blanco s.n. (P); Jaén, El Serrate, vert. Oriental, 17.vii.1925, Cuatrecasas s.n. (BC 39099, MAF 2781); Jaén, Almadén, vert. SE, 19.vii.1925, Cuatrecasas s.n. (BC 39100); Jaén, Santiago de la Espada, La Fresnedilla, 17.vii.1975, González Rebollar et al. s.n. (MA 479937); Murcia, Sierra de Espuña in regno Murcico, 1850, Guirdo s.n. (G); Murcia, Sierra Cantón, Abanilla, 19.v.1985 Robledo s.n. (ARAN). Teruel, Sierra de Javalambre, 11.vii.1895, Pau s.n. (MA 120957); Valencia, Bicorp, vi.1915, Vicioso s.n. (BC 39109); Valencia, Enguera, 10.vii.1919, Font Quer s.n. (BC 39105); Valencia, In saxosis Valldigna, 17.vi.1791, Cavanilles s.n. (MA 121516); Ricorp, 26.vi.1915, C. Vicioso s.n. (MA 120955, MA 120962); Valencia, Les Foies, Simat, vi.1976, Mansanet & Mateo s.n. (VAB); Valencia, 1815, Léon Dufour s.n. (MPU); Valencia, de Chiva à la sierra de Santa Maria, 14.vi.1881, Barbey s.n. (G); Valencia, Villagordo del Cabriel, 11.v.1984, Mateo & Figuerola s.n. (VAB); Valencia, Monte del Tejo, Requena, 20.vi.1984, Sanchis & Alcover s.n. (VAB); Valencia, Pico del Águila, Requena, 10.vi.1986, García s.n. (VAB); Valencia, Corbera d’Alcira, 1944, Borja s.n. (BC 100370); Valencia, Utiel, La Rampina, 30.iv.1988, Navarro s.n. (VAB); Valencia, Cerro Calderón, pr. Puebla de San Miguel, 3.vii.1988, Mateo 940 (VAB); Valencia, Llaurí, 3.vi.1989, Pascual s.n. (VAB); Valencia, Ayora, la Muela de Carcelén, 12.vi.1990, Mateo et al. s.n. (VAB); Valencia, Sinarcas, umbría del Picarcho, 25.vi.1991, Navarro s.n. (MA 599090); Valencia, Sierra de la Murta, Borja s.n. (MAF 2777). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 = C. semisecta var. basiclada Murb. in Acta University Lund. 33(12): 116 (1897). Ind. loc.: ‘Valentia: Moxente (Bourg. 1852); Sierra de la Cueva-Santa (Reverch. 1891). Albacete: Hellin, au pied de la sierra de las Caldas (Rouy, 1881). Murcia: Serra de Espuña (Guirao, 1850).’ Lectotype: (here designated) [Spain] Sierra de la Cueva-Santa, dans les maquis rocheaux, sur le calcaire jurassique, 700 m, vi.1891, Reverchon 588 (G). C. semisecta. However, we have studied most of them and they are not C. semisecta (some of them are misidentifications of C. filicaulis Durieu or C. dichotoma). Consequently, in our opinion, C. semisecta has not yet been collected in north Africa. In our view, the lectotype of C. semisecta var. basiclada should be included within the variation of the species. There is no evidence of distinction from typical C. semisecta. REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX 4. Campanula rimarum Boiss., Fl. Orient. 3: 931 (1875). Ind. loc.: ‘Hab. in fissuris rupium Lyciae ad Duden prope Elmalu (Bourg !). Fl. Mai.’ Lectotype: (designated by Damboldt, 1978: 50) [Turkey], in fissuris rupium ad Duden prope Elmalu, 1.vi.1860, Bourgeau s.n. (G). Isotypes (E, G, MPU, P, W). Chromosome number: 2n = 20, 2n = 20 + 1B, 2n = 20 + 2B (Contandriopoulos, 1970, 1976). Area: Antalya, south-eastern Turkey Habitat: Limestone rocks. 1000–2800 m. Phenology: Flowering June Illustration: Fig. 20. Representative specimens examined: We were able to see only the type specimens, although several others are known (Contandriopoulos, 1970; Damboldt, 1978). 5. Campanula sulphurea Boiss., Diagn. Pl. Orient. Ser. 1, 11: 64 (1849) Ind. loc.: ‘Hab. in arenà mobili purà collium littoralium Syriae circà Gaza (Boiss. Aucher n. 1856), circà Beyrout (Boiss. Pestalozza).’ Lectotype: (here designated) (Palestine] Gaza & Aucher-Eloy 1856 (G, isolectotype P). Description: Annual 5–30 cm, hispid. Stem erect, simple or spreadingly branched from base. Leaves 7– 20 ¥ 4–6 mm, sessile, entire or nearly so, oblong to oblong-lanceolate, obtuse to acute. Flowers in shortly pedunculate dichasia. Pedicels hispid, hairs 1.1– 1.4 mm long. Calyx 9–13.5 ¥ 1.7–2.5 mm (11.3– 13.8 ¥ 2.6–3.7 mm in fruit), hispid, hairs 0.5–1.7 mm long; lobes 7–10 mm long (8.9–14 mm long in fruit), lanceolate, subacute; appendages 2–3.5 mm long (2.4– 4.2 mm long in fruit), ovate, obtuse, concealing ovary in fruit. Corolla 18–21 mm long, divided to 1/3–1/2, campanulate, glabrous or hairy on nerves outside, sulphur-yellow; lobes 6.5–10.6 mm long, triangularovate, obtuse; tube 15–17 mm long. Stamen 9.4– 10.9 mm long; filiform part of filament 0.1–0.4 mm long; base 3.5–4 ¥ 2.3–2.5 mm, obovate, hairy in the upper margin, hairs c. 0.2 mm long; anthers 5.8– 6.7 mm long; pollen 20–25 mm in diam., with a low density of spinules (0.60–0.68 spinules mm-2). Style 12.5–15 mm long, included. Stigma 2.5–5 ¥ 1–1.3 mm. Capsule 6.1–7.2 ¥ 5.7–6 mm, concealed by the acrescent connivent calyx lobes and appendages, sparsely hairy on the keels (hairs 0.5–1.5 mm long). Seeds 0.8– 0.9 ¥ 0.4 mm, yellowish. Chromosome number: unknown. Area: Egypt, Sinai, Palestine, Lebanon and Syria. 200–1200 m. Habitat: Sandy soils. Phenology: Flowering February – May Illustration: Feinbrun-Dothan (1977: 478) Representative specimens examined: EGYPT: Versant W de Gebel Ammone, entre le Caire et Suez, 2.v.1880, Sickenberger s.n. (G); 15 km in WSW von Cairo, iv.1885, Schweifurth s.n. (MPU); Cairo, 29.iv.1885, Volbens s.n. (G); Egypte, environs du Cairo, Deflers s.n. (MPU). LEBANON: route de Beyrouth a Saïda, 7.v.1853, Blanche 691 (W); forets de pins au Sud de Beyrouth, 7.v.1858, Gaillardot s.n. (MPU); Bir Hassen, 12.v.1955, Mouterde 11522 (G); Nebi l’Aousaye, 11.v.1933, Mouterde 2218 (G); Beryti, 8.v.1890, Vincent s.n. (MPU); Beirut, 29.v.1901, Post s.n. (G). PALESTINE: Jaffa, prope Sarona, 12.v.1897, Bornmüller s.n. (P); Gaffa, 25.iv.1912, 1–20 m, Dinsmore 5059 et al. (G); Herzlia, near Tel.Aviv, sandy soil, 26.iv.1928, Eig et al. s.n. (MA 120984, W); Gaza, 13.v.1896, Fonck s.n. (W). SINAI: Isthmus Aegyptiaco-Palaestinus, Jebel Ekhfeynm 16.iv.1891, Deflers s.n. (MPU); Egypte, El Ary- © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Description: Small fragile spreading whitehispidulous annual 5–15 cm. Stem flexuous, filiform, branched from base. Leaves 2–15(20) ¥ (2)4–6(8) mm, lower attenuate into petiole to 15 mm, upper sessile, entire, elliptical to oblong-spatulate, obtuse. Flowers solitary, terminal. Pedicels filiform, hispid, with long hairs 1.2–1.7 mm long. Calyx 5–6 ¥ 2.5–3 mm (6.1– 6.7 ¥ 3.6–4 mm in fruit), densely hispid, hairs 0.7– 1.7 mm long; lobes 3.6–4 mm long (4.1–4.5 mm long in fruit), ovate-triangular, acute; appendages 1.4– 1.9 mm long (2–2.3 mm long in fruit), truncate to rounded, obtuse, concealing ovary in fruit. Corolla 4– 5.5 mm long, shortly exceeding calyx lobes, divided to c. 1/3, narrowly cylindrical to narrowly tubularcampanulate, hairy outside, pale violet; lobes 1.3– 1.5 mm long, broadly triangular to semielliptical, obtuse; tube 3.5–4 mm long. Stamen 3–4 mm long; filiform part of filament 0.1–0.2(0.3) mm long; base 1.7– 2 ¥ 0.8–1 mm, subelliptical to oblong-elliptical, papillose in the margin; anthers 1.5–1.7 mm long; pollen 20–23 mm diameter, with a high density of spinules (1.8–2.0 spinules mm-2). Style 4.3–4.5 mm long, ±included. Stigma 0.4–0.6 ¥ 0.4 mm. Capsule 4.3– 5.5 ¥ 4–5.2 mm, concealed by the acrescent connivent calyx lobes and appendages, sparsely hairy on the keel and on the valves, hairs 0.7–1.7 mm long. Seeds 0.75– 0.9 ¥ 0.3–0.4 mm, yellowish to brownish. 227 228 L. SÁEZ and J. J. ALDASORO A 2mm E 1cm 0.2mm B 2mm 1mm D C Figure 20. Campanula rimarum. Turkey: in fissuris rupium ad Duden prope Elmalu, 1.vi.1860, Bourgeau s.n. (P, isotype). A: habit; B: stamen and detail of the base of stamen (left); C: flower; D: seeds; E: dissection of a flower; F: capsule. sch au S du Jebel Ekhfeyn, 14.iv.1891, Deflers s.n. (MPU); prope El Arysch, iv.1855, Kotschy 455 (P). SYRIA: Jebel Amqabya N, 25v. 1945, Davis 8539 (G 154513, W). 6. Campanula saxonorum Gand. in Bull. Soc. Bot. France 65: 54 (1918). Ind. loc.: ‘Anatolia, ad Amasia (Bormüller n. 581!); Armenia, in monte Dolidayh (Bormüller n. 3429!) et ad Gumuschkhane (Sintenis n. 6014!).’ Lectotype: (designated by Damboldt, 1978: 48) [Turkey, Gümüsane], Aghakoei, 20.vi. 1894, Sintenis 6014 (B, isolectotypes B, E, G, GOET, LD, P, W, WU). Description: Annual 3–15(27) cm, hispid, green. Stem usually erect, simple or dichotomously branched from base. Leaves 10–60 ¥ 4–18 mm, entire or incised-dentate, lower subssesile, spathulae; cauline sessile, elliptical to oblong-lanceolate, obtuse. Flowers in dichasia. Pedicels hispid, with long hairs 1–2 mm long and small hairs 0.2–0.5 mm long. Calyx 9.6–14 ¥ 1.5– 3 mm (12.5–15.5 ¥ 3.6–4.6 mm in fruit), hispid, hairs 0.8–1.4 mm long; lobes 5–10 mm long (7–11.5 mm long in fruit), linear-lanceolate, acute ending in a subacuminate or subulate awn up to 2 mm; appendages 5–10 mm long (7–11.5 mm long in fruit), rounded, acute or obtuse, concealing ovary in fruit. Corolla © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 2mm 1mm F REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX Chromosome number: Unknown. Area: Turkey Habitat: Stony places. 1300–1700 m. Phenology: Flowering June – July Illustration: Fig. 21. Notes: The distinction of C. saxonorum and C. strigosa in most cases presents no difficulty. The lobes of calyx with a subulate apex (3)5–10 mm long in C. strigosa makes this species very easy to recognize. Representative specimens examined: TURKEY: Armenia turcica, Kharput, S. Kuschnas, 8.vi.1889, Sintenis 686 (G); Armenia Turcica, Egin, in campis prope Szanduk, 15.vi.1890, Haussknecht 2600 (MA 628444); Armenia Turcica, Szanduk, 20.vi.1890, Haussknecht 6014 (W); Aghakoci, Tunceli-Pülümür, 6 miles form Pülümür, 7.vi.1957, Davis & Hedge 29205 (G); Prov. Tunceli, Nozat, 14.vii.1957, Davis & Hedge 31124 (G, W); Bulandi, 16.vii.1965 (W); 2 km NE Mursal (N Yama Daglari), Sivas, 1.vii.1976, Sorger 76.12.5 (W); 25 km NE Feke, Adana, 7.vii.1977, Sorger 77.26.6 (W); Tunceli, Pülümür, 17.vi.1980, Yíldirimi & Eren 3317 (G); Armenia Minor, in monte Deli-dagh, vi.1983, Bornmüller 3429 (W); Agri: Agri-Hamur, 2 km N Hamur, 9.vi.1988, Nyedgger 43655 (G); Sivas, DivrigiSincan, 28 km W Divrigi, Nyedgger 43843 (G). 7. Campanula strigosa Banks & Sol. in Russell, Nat. Hist. Aleppo ed. 2, 2: 246 (1794). Ind. loc.: ‘Aleppo.’ Lectotype: (designated by Rechinger & SchimannCzeika, 1965: 12) [Syria], Aleppo, Russell, s.n. (BM). ∫ C. russelliana Schult. in Roem. & Schult, Syst. Veg. 5: 142 (1819) (as ‘russeliana’). Description: Annual 5–35 cm, hispid. Stem erect to subflexuous, simple or dichotomously branched in the upper part, rarely from base. Leaves 4–30(60) ¥ 6– 10(30) mm, sessile, elliptical to oblong-lanceolate, obtuse or subacute. Flowers in shortly pedunculate dichasia. Pedicels hispid, hairs 1.4–2 mm long. Calyx 10.2–18.7 ¥ 2.7–7 mm (14.2–21.8 ¥ 4.5–7.1 mm in fruit), hispid, hairs 0.9–2 mm long; lobes 9–14.7 mm long (9–14.4 mm long in fruit), ovate-lanceolate, abruptly ending in a subulate (3)5–10 mm long awnlike tip; appendages (1.2)3–6 mm long (3–7.4 mm long in fruit), ovate, obtuse, concealing ovary in fruit. Corolla 16–23.5 mm, divided to c. 1/2, tubular-campanulate, glabrous or hairy on nerves outside, reddish purple to violet-blue; lobes 7.7–12 mm long, ovate to semi-elliptical, obtuse; tube 7.5–12 mm long. Stamen 6–11.5 mm long; filiform part of filament 0–0.1 mm long; base 2–3.7 ¥ 1.5–2.5 mm, oblong-elliptical, emarginate, sparsely hairy, hairs 0.1–0.2 mm long; anthers 4–7.7 mm long; pollen 20–23 mm diameter, with a intermediate density of spinules (1.0–1.1 spinules mm-2). Style 10–15 mm long, included. Stigma 0.9–3.1 ¥ 0.2–1.5 mm. Capsule 7.5–10 ¥ 6– 7.7 mm, concealed by the acrescent connivent calyx lobes and appendages, sparsely hairy on the keels (hairs 0.8–2 mm long) and occasionally on the valves. Seeds 0.9–1.1 ¥ 0.8–1 mm, yellowish. Chromosome 1976) number: n = 10 (Contandriopoulos, Area: Palestine, Lebanon, Syria, southern Anatolia. Habitat: Fields, stony places, 100–2000 m. Phenology: Flowering March – May Illustration: Fig. 22. Representative specimens examined: LEBANON: Nahr ad Damur, 8.v.1949, Mouterde 9598 (G); Beyrouth, 18.v.1816, Gaillardot s.n. (G-Boiss.); In planitie Coelesyriaca (Bekâa) prope Rajak, 16.v.1910, Bornmüller 12118 (G); Oued Zénati, 24.vi.1911, Clavé s.n. (P); Rayak, Bekaa, cultures, 1000 m, 11.v.1957, Pabot s.n. (G); Tourzaya, audessus de Byblos, 16.iv.1957, Pabot s.n. (G); Antiliban: entre Aïn Bigé et kaf Zent, 18.v.1817, Gaillardot 2040 (G); Zahlé-Bekaa, s.d., Frere Luis s.n. (P); In felsen bei Ghazir, 28.iv.1934, Busujan 103 (W). PALESTINE : Wadi Shomrya, Carmel, 9.iv.1942, Davis 4379 (G, W); Carmel, v.1846, Boissier s.n. (G); Thabor, 19.iv.1896, Forrek s.n. (W); Jerusalem, iv.1846, Boissier s.n. (G, P); Ouadi Qandil, 5.v.1954, Pabot s.n. (G); Mt. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 10.5–16.5 mm long, divided to 1/4–1/3, narrowly tubular-campanulate, hairy outside at base and along the nerves, violet or blue-violet; lobes 4.5–8 mm long, triangular-ovate, obtuse; tube 6–8.5 mm long, hairy outside at base and on the nerves. Stamen 5.8–9.3 mm long; filiform part of filament 0.7–1 mm long; base 1.7–2.1 ¥ 1.7–2 mm, oblong to ovate, sparsely hairy in the margin, hairs c. 0.1 mm long, and on the abaxial surface hairs 0.5–2 mm long; anthers 3.3–6.5 mm long; pollen 18–21 mm diameter, with a high density of spinules (1.5–1.56 spinules mm-2). Style 8–10 mm long, included or occasionally subexerted. Stigma 0.9– 2.7 ¥ 0.6–1 mm. Capsule 6.5–7.5 ¥ 5.5–6 mm, concealed by the acrescent connivent calyx lobes and appendages, hairy on the keels (hairs 0.6–1.2 mm long) and occasionally on the valves (hairs 0.2–0.5 mm long). Seeds 1–1.1 ¥ (0.4)0.5–0.6 mm, yellowish. 229 L. SÁEZ and J. J. ALDASORO 0.2mm 230 B 5mm G 2mm C D 1cm 5mm A 5mm F E 1mm Figure 21. Campanula saxonorum. A: Turkey, Aghakoei, 20.vi.1894, Sintenis, 6014 (P); B–G: Turkey, Tunceli, Nozat, 1700 m, 14.vii.1957, Davis & Hedge D 31124 (G, W). A: habit; B: dissection of a flower; C: capsule; D: fruiting calyx; E: seeds; F: flower; G: stamens and detail of the base of stamen (right). Ebal, 19.iv.1942, Davis 4482 (G, W); Jerusalem, Messalebe, 3.v.1933, Eig et al. 292 (MA 120924); Samarie, 13.iv.1880, Barbey s.n. (G); Haifa, 15.iv.1897, Bornmüller 1093 (G); El Wadi, 5.vi.1938, Mouterde 6416 (G); Kfar Tibnite vers Beaufort, 6.v.1946, Mouterde 8526 (G); Jerusalem, Sheikh garrah, 2.iv.1963, Maitland 78 (G) Inter segetes vs. Eden, 20.vii.1855, Kotschy s.n. (G, MPU); Jerusalem, 15.iv.1912, Dinsmore 6014 (MA 120925). SYRIA: Syriae borealis, env. de Homs, v.1910, Haradjan s.n. (G); Cote de Son Qalat Makal, Banias, 19.iv.1952, Pabot s.n. (G); Champs de blé sur la rive gauche et près de l’embouchoure du Saïnih, c. Saïda, 21.iv.1853, Blanche s.n. (G,MPU); Alep, 29.iv.1841, Kotschy 182 (G); champs audessous de Aïn Hallalie, premiéres collines du Liban, a l’est de Saïda, Syrie, 11.v.1858, Gaillardot s.n. (MPU, G); Aïn Hallalie, 11.v.1858, Gaillardot 1378 (P); Port naturel du Litani, 31.v.1955, Mouterde 11536 (G); Rarhayat el Foukhar, 15.v.1938, Gombault 5190 (P); Hasmiyeh, 19.iv.1934, Mouterde 3002 (G); Syrie S, N de Deraa, Hauran, © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 2mm REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX 231 F B 1 mm 4 mm 5 mm 5 mm 2 cm 5 mm A D C Figure 22. Campanula strigosa. Lebanon: Rayak, Bekaa, 1000 m, 11.v.1957, H. Pabot (G). A: habit; B: stamen; C: flower; D: fruiting calyx; E: capsule; F: seed. 13.vi.1956, Pabot s.n. (G); Jebel Garbi supra Zebdani, 30.v.1880, Peyron s.n. (G). TURKEY: 30 km S of Antioch (Antakia), 6.vi.1938, Disnmore 14014 (G); Plaine de Mersina, Cilicie, 2.v.1855, Balansa s.n. (MPU); Plaine de Mersina, Cilicie, 2.v.1855, Balansa 625 (G, P, W); Prov. Hatay, Antakya, near St. Peter’s Church, Davis & Hedge 27256 (G); 3–6 km N Halfeti, 23.v.1983, Sorger 83-3-2 (W); 25 km E Kahta, nebefluss des Euphrat, 4.v.1980, Sorger 80-27-45 (W); Maras, Ahir-Dagin, 6.v.1990, Nydegger 45362 (G). 8. Campanula reuteriana Boiss. & Balansa in Boiss., Diagn. Pl. Orient. Ser. 2, 3: 108 (1856). Ind. loc.: ‘Hab. in Ciliciâ Kotschy pl. exs. 1836 n∞. 349, in Ciliciâ propè Gülek Boghas Tauri cl. Balansa Julio fructiferam legit.’ © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 E 232 L. SÁEZ and J. J. ALDASORO Lectotype: (designated by Rechinger & SchimannCzeika, 1965: 11) [Turkey], Cilicia, Kotschy 349 (W, isolectotypes G, P). Chromosome number: 2n = 22 1972, 1980). (Contandriopoulos, Area: South-eastern Turkey, N Iran and N Iraq 9. Campanula camptoclada Boiss., Diagn. Pl. Orient. Ser. 1, 11: 63 (1849). Ind. loc.: ‘Hab. in fissuris ruppium calidarum in vallibus Antilibani circà Banias saxis arctè incumbens et secùs ea ramos suos curvans. Legi Apr fine.’ Lectotype: (designated here) [Syria], Rochers de Banias, iv-v. 1846, Boissier s.n. (G). Description: Annual 10–32.5 cm, hispid. Stem usually erect, dichotomously branched from base. Leaves 7– 25(50) ¥ 2–11(22) mm, sessile (lower leaves sometimes petiolate), entire or nearly so, ovate to ovate-oblong, obtuse or acute. Flowers lateral and terminal. Pedicels hispid, hairs (0.4)1–1.8 mm long. Calyx 7–10.2 ¥ 2– 3.1 mm (9–12.4 ¥ 3–4 mm in fruit), hispid hairs 0.6– 1.8 mm long; lobes 5–8 mm long (6.3–9 mm long in fruit), ovate to lanceolate, acute or subacute; appendages 2–3.2 mm long (2.5–4 mm long in fruit), rounded, obtuse, concealing ovary in fruit. Corolla 9–23 mm long, divided to 1/4–1/3, tubular-campanulate, glabrous or hairy on nerves outside, limb blue, tube white; lobes 3–6 mm long, ovate to semi-elliptical, obtuse; tube 6–14 mm long. Stamen 6–7.7 mm long; filiform part of filament absent; base 2.3–2.6 ¥ 1.4– 1.9 mm, oblong-elliptical, sparsely hairy, hairs 0.1– 0.2 mm long; anthers 3.5–5.2 mm long; pollen 20– 23 mm diameter, with a low density of spinules (0.40– 0.46 spinules mm-2). Style 8–12 mm long, included. Stigma 1.8–2.7 ¥ 1–1.4 mm. Capsule 5.5–7.7 ¥ 4.9– 5.3 mm, concealed by the acrescent connivent calyx lobes and appendages, hairy on the keels (hairs 0.8– 1.5 mm long) and on the valves (hairs 0.5–0.8 mm long). Seeds 1.1–1.3 ¥ 0.9–1.1 mm, yellowish. Habitat: Rocky slopes, fields. 400–1830 m. Chromosome number: Unknown Phenology: Flowering March – May Area: Jordan, Lebanon, Palestine and Syria. Illustration: Fig. 23. Habitat: rock crevices. Representative specimens examined: IRAN: Iran W, Kuh-i-Marab v.1910, Strauss s.n. (W); SYRIA: Ain Syria (Ayn Sajriyah) prope Svedia, 1836, Kotschy 349I (W); TURKEY: Amanus, Mount Dümanly, 700 m, 11.vii, Haradjian s.n. (W); in monte Tauro, 1836, Kotschy 349 (P, W); Cilicia: village de Gulek-Boghar, prés du défilé des Portes Ciliciennes, 6.vii.1855, Balansa s.n. (P); Taurus, VII. 1856, Balansa s.n. (G); Cilicia, vi.1860, Reuter s.n. (P, W); c. 40 km NW Mardin, 23.vi.1965, Sorger s.n. (W); Deirik, sur basalte, 11.v.1955, Mouterde 493 (G); Elazig, 15 km S von Maden, 14.v.1966, Eiselt s.n. (W); Village de Gulek-Boghas, N de Tarrous, 6.vii (G); Mardin, Kasrik Bogazi, 11 km Phenology: Flowering March – April Illustration: Feinbrun-Dothan (1977: 480) Wadi Representative specimens examined: JORDAN: Tawadin (near Safad), 3.v.1942, Davis 4594 (W). LEBANON: El Wadi, près Zahle, 5.vi.1938, Gombault 5194 (P); Ruines du Château de Beaufort, S Liban, 5.v.1955, Mouterde s.n. (G). PALESTINE: Mt. Carmel, Wadi Shomrya, 9.iv.1942, Davis 4397 (G); Between Binyamina and Kabara, rock crevices at the west foot of Carmel, 9.iv.1942, Davis 4476 (W). SYRIA: Rochers de Baniyas, © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Description: Annual 8–30 cm, hispid. Stem erect, simple or dichotomously branched in the upper part, rarely from base. Leaves 10–32 ¥ 5–10 mm, sessile, entire or nearly so, elliptical to oblong-lanceolate, obtuse or acute. Flowers in shortly pedunculate dichasia. Pedicels hispid, with long hairs 0.7–2(2.8) mm long and small hairs (0.1)0.2–0.4(0.5) mm long. Calyx 13–19.5 ¥ 4–6 mm (13–20.6 ¥ 5–7.8 mm in fruit), hispid or strigose, hairs (0.5)0.8–2 mm long; lobes 10– 18 mm long (8–15 mm long in fruit), lanceolate, acute; appendages 3.7–4.8 mm long (5.6–6.8 mm long in fruit), rounded, obtuse, concealing ovary in fruit. Corolla 20–29 mm long, divided to 1/3–1/2, campanulate, glabrous or hairy on nerves outside, light violet; lobes 9.6–11 mm long, triangular-ovate, obtuse; tube 12.2.-18 mm long. Stamen 8–9.1 mm long; filiform part of filament 0.1 mm long; base 2.7–3 ¥ 2.3– 2.5 mm, triangular-ovate, hairy in the margin, hairs c. 0.2 mm long; anthers 5.5–6 mm long; pollen 20–25 mm diameter, with a intermediate density of spinules (1.00–1.16 spinules mm-2). Style 9–13.2 mm long, included. Stigma 2–2.1 ¥ 1.4–1.5 mm. Capsule 8.5– 9.9 ¥ 6–8.8 mm, concealed by the acrescent connivent calyx lobes and appendages, hairy on the keels (hairs 0.7–1.7 mm long) and on the valves (hairs 0.2–0.5 mm long). Seeds 1–1.1 ¥ 0.6–0.8 mm, yellowish. from Gizre to Sirnak, 400 m, limestone slopes, s.d., Davis 42650 (G). REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX 233 A B 3 mm 2 cm 5 mm 5 mm 5 mm 1 mm D C E Figure 23. Campanula reuteriana. Turkey [Cilicia] in monte Tauro, 1836, Kotschy 349 (W). A: habit; B: stamen; C: flower; D: seed; E: fruiting calyx; F: capsule. 4.v.1846, Boissier s.n. (G); Baniyas, 2.vi.1884, Peyron s.n. (G); Baniyas, iv.1937, Mouterde s.n. (G). 10. Campanula propinqua Fisch. & C.A. Mey., Index Sem. Hort. Petrop. 2: 32 (1836). ∫ Roucela propinquaxe ‘Roucela:propinqua’ (Fisch. & C.A. Mey.) Karadze in Zametki Sist. Geogr. Rast. 32: 55 (1976). Ind. loc.: ‘Hab. in provincia Aderbeidschan Persiae borealis, locis sterilibus siccis.’ Lectotype: (designated by Damboldt, 1978: 48) [Iran] described from material cultivated in Hort. Petrop. with provenance: in provincia Adzerbeidschan Persiae borealis, Szovits s.n. (LE, isolectotype P). = C. pestalozzae Boiss., Diagn. Pl. Orient. Ser. 1, 11: 62 (1849). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 F 234 L. SÁEZ and J. J. ALDASORO Ind. loc.: ‘Hab. in Caramaniâ ubi legit amic. Dr Pestalozza.’ Lectotype: (here designated) (Turkey], Caramania, 1846, Pestalozza s.n. (G). = C. propinqua var. parviflora Turrill in Bull. Misc. Inform. 1929: 229 (1929). Ind. loc.: ‘N. Persia: Yam, Dik Dash, north-west of Tubriz, August 1928, Gilliat-Smith 2645.’ Lectotype: (designated by Rechinger & SchimannCzeika, 1965: 13) [Iran], Yam, Dik Dosh, NW Tabriz, Gill.-Sm. 2465 (K). Description: Annual 3–15(27) cm, hispid. Stem usually erect, simple or dichotomously branched from base. Leaves 5–20(30) ¥ 3–10 mm, entire or inciseddentate, lower subssesile, spathulae; cauline sessile, elliptical to oblong-lanceolate, obtuse. Flowers terminal. Pedicels hispid, with long hairs 0.7–0.9 mm long and small hairs 0.2–0.4 mm long. Calyx 3.4– 10.4 ¥ 1.4–2 mm (9–12.5 ¥ 2.6–4 mm in fruit), hispid, hairs c. 0.8 mm long; lobes 2–7 mm long (5–9 mm long in fruit), linear-lanceolate, acute; appendages 0.9– 3.4 mm long (1.5–3.5 mm long in fruit), ovate to triangular, acute or obtuse, usually concealing ovary in fruit. Corolla 5.8–17 mm long, divided to 1/3–1/2, narrowly tubular-campanulate, hairy outside at base and along the nerves, violet or blue-violet; lobes 2–6 mm long, triangular-ovate, obtuse; tube 5–11 mm long, hairy outside at base and on the nerves. Stamen 6.4– 7.8 mm long; filiform part of filament 0.25–1 mm long; base 0.9–2 ¥ 0.7–1.2 mm, oblong to ovate, sparsely hairy in the margin, hairs c. 0.1 mm long; anthers 3.8– 5 mm long; pollen 20–24 mm diameter, with a intermediate density of spinules (2.60–2.71 spinules mm-2). Style 5.5.-7.5 mm long, included or occasionally subexerted. Stigma 0.5–3.1 ¥ 0.6–1.5 mm. Capsule 5– 6.2 ¥ 4–5.2 mm, concealed by the acrescent connivent calyx lobes and appendages, hairy on the keels (hairs 0.7–0.9 mm long) and on the valves (hairs 0.2–0.5 mm long). Seeds 1–1.2 ¥ 0.4–0.5 mm, yellowish. Chromosome number: 2n = 16 1972, 1976). (Contandriopoulos, Area: north-western Iran, eastern Turkey, and southern Caucasus (Nakhichevan and Georgia). Phenology: Flowering March – July Illustration: Fig. 24, Fedorov (1976); pag. 135: 4. Notes: Campanula propinqua is considerably variable, mainly in plant size and general shape. This high variability resulted in the description of many taxa. Victorov & Elenvsky, (1998) proposed including C. propinqua in C. dichotoma, in order to combine two rather similar and variable species. However, two characters discriminate them: the presence of two hair types in pedicels and fruits of C. propinqua, with only one type in C. dichotoma; and the different shape of staminal appendages in both species. In addition, the divergent chromosome number supports the view that C. propinqua and C. dichotoma deserve taxonomic recognition at the specific level. None of the morphological features used by Boissier (1849) to distinguish C. pestalozzae appear to be constant. Damboldt (1978) included C. pestalozzae within the the variation of C. propinqua. However, C. propinqua has a large variabiliy in the size of its floral parts, and shows no relation with any other characters or with its ecological and geographical distribution. Campanula propinqua var. parviflora Turrill is probably a mere variant of C. propinqua without any taxonomic value. Plants from northern Iran do not differ from other accessions of the species. Representative specimens examined: IRAN: Azerbaijan W, Sharhestan, Marand, 23.vi.1965, Scharef 6485 (W); Persia boreal s.d., Szovits s.n. (P, LE); Azerbaijan E, 9 km SE of Marand, 16.vii.1964, Grant s.n. (W); Azerbaijan: prope station viae ferr. Nagram, 24.v.1933, Prilipro s.n. (P); Azerbaijan W; Serow, 7.vii.1968, Petrovitz 165 (W); Aderbidjan, s.d. Aucher-Eloy 4901 (G); Azerbaijan orientate., in jugo inter Marand et Sufian, 1600– 1750 m, 6.vi.1971, Rechinger 41204 (G); Azerbaijan occidentalis, in jugo Quschi inter Shapur et Rezaiyeh, 13.vi.1971, Rechinger 41899 (G); in valle fluvii Qotur W Khvoy versus fines Turcicas, 11.vi.1971, Rechinger 41740 (W); Azerbaijan occidentalis: in valle fluvii Quotur, W Khvoy, 17.vii.1974, Rechinger 49548 (G); Azerbaijan orientate. in arenoso glareosis a Bunab boreo.occidentem vs., 15.vi.1977, Rechinger 56798 (G). TRANSCAUCASUS , NAKHICHEVAN : pr. Pag. Arincz, 3.vii.1952, Fedorov & Smoljianinova s.n. (W, G); Aznabiurt; 2.vi.1960, Magmanesian s.n. (MA 560705); Shakhbuz, 2.vi.1934, Grossheim & Gurvitsh s.n. (W); prope pag. Arincz, 3.vii.1952, Prilipro s.n. (P); (G); Nachiczevan: prope pag. Aznajurt, 8.vi.1947, A. Grossheim et al. s.n. (G). TRANSCAUCASUS , GRUCIYA: © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 = C. balansae Boiss. & Hausskn. in Boiss., Fl. Orient. 3: 931 (1875). Ind. loc.: ‘Hab. in arenosis et glareosis torrentium regionis alpinae inferioris montis Masmeneudagh Cappadociae (Bal !) et Berytdagh Cataoniae (Haussk !). Fl. Jul Aug.’ Lectotype: (designated by Damboldt, 1978: 48) [Turkey], in glaer. torrentium Berit-dagh, Taurus Cataonicus, 15.viii. 1865, Haussknecht s.n. (G). Habitat: Fields, steppes, dry slopes, sandy and stony places. 60– 2000 m. REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX 235 4 mm F 1 mm B 5 mm 2 mm 2 cm A C D Figure 24. Campanula propinqua. A-F: Iran, in valle fluvii Qetur, W Khroy, 17.vi.1971, Rechinger 41740 (W); A: habit; B: capsule; C: stamens; D: fruiting calyx; E: flower; F: seeds. Vallis Bardus-Zai, prov. Karo, 21.vi.1904, Schicloiowsky (COI). TURKEY: Armenia turcica, Egin, in campis prope Szanduk, 15.vi.1890, Haussknecht s.n. (G); Kurdistania W, Taurus Cataonicus, inter urbem Malatia et vicum Kjatcha, in declivibus glaerosis inter Kory et Furendscha, 19.vii.1910, Handel-Mazzetti 2493 (W); Berithdagh, Cataoniae, 15.viii.1865, Haussknecht s.n. (W); Adalia, 16.iv.1860, Bourgeau s.n. (P, W); Isparta, Anamas-Schlucht (Zindan Magarasi) am Ayvali Cay (Köpruirmagi), 25.vi.1969, Fitz & Spitzenberger 977 (W); Malatya, Malatya-Pörtuge, 26 km anch Abzweingung Kube Dâg Südseite, 17.vii.1982, Nyedgger 17211 (G); Bingöl, Elazig-Bingöl, 21 km W Bingöl, 19.vii.1982, Nyedgger 17243 (G); Gümüsane, TorulSiran, 40 km N Siran, 1280 m, Nyedgger 19119 (G); Dogancal, Sivas Department, 2.vii.2001, Aldasoro 2731 (MA); Anamurium, 22.iv.1985, Nyedgger 40394 (G); Antalaya: Gazipasa-Alanya, 15 km E Alanya, 2.v.1986, Nyedgger 41581 (G); Toakt, Zile-Amasya, 5 km N Zile, 27.vi.1987, Nyedgger 42959 (G); Adalia, sur les pelouses, 16.iv.1860, s.r. (G); Asia minor, oest Pontus, 1858, Tchihatchef s.n. (G); près Ispir, 6.vi.1862, Bourgeau 453 (G); Armenia prope Ispir, vi.1853, Huet du Pavillon s.n. (G). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 E 236 L. SÁEZ and J. J. ALDASORO 11. Campanula hierosolymitana Boiss., Diagn. Pl. Orient. Ser. 1, 11: 62 (1849). Ind. loc.: ‘Hab. ad pedem rupium propè Hierosolymam juxtà speluncas Regnum et Judicum dictas et in Samarià propè Naplouse. Legi Maio ineunte.’ Lectotype: (designated here) [Palestine], aux pieds des rochers à Jerusalem, Naplouse, iii-iv. 1846 Boissier s.n. (G). Chromosome number: unknown. Area: Lebanon, Jordan, Palestine, Syria and southeastern Turkey. Habitat: Rocky places and arid slopes. 100–1200 m. Phenology: Flowering March – May Illustration: Feinbrun-Dothan (1977: 479). Representative specimens examined: JORDAN: Wadi es Sik, 1880, Barbey s.n. (G); Zerka Main to Jebel Atturus, 25.iv.1945, Davis 9368 (G, W). PALESTINE: ad pagum Malcha, prope Hyerosolym., 19.iv.1855, Kotschy s.n. (W); Kinrot valley, Upper Jordan Valley, 2 km NE of Kibbutz Haon, 2.iv.1989, Danin et al. 39048 (G); Jerusalem, près de Bassis de Salomon, 26.iv.1883, Cramer s.n. (G); Jericho, in declivitatibus aridis as Ain-Sultan et Wadi-Kilt, 30.iii.1897, Bornmüller 1095 (W); Env. de Jersualem, Joffé (MPU); De 12. Campanula dichotoma L., Cent. Pl. II: 10 (1756). Ind. loc.: ‘Habitat in Sicilia, Syria.’ Lectotype: (designated by Victorov & Elenvsky, 1998: 56) Boccone P. 1674. Icones et descriptiones. p. 83, Table 45, fig. 1. = C. afra Cav. in Anales Ci. Nat. 3: 21 (1801). Ind. loc.: ‘El Sr. Broussonet la encontró en Salé.’ Lectotype: (here designated) [Morocco], Herbario Cavanilles, Mogador, Salé, Broussonet s.n. (MA 120963). ∫ C. dichotoma ssp. afra (Cav.) Maire in Cavanillesia 2: 174 (1930). = C. kremeri Boiss. & Reut., Pugill. Pl. Afr. Bor. Hispan. 75 (1852). ∫ C. dichotoma var. parviflora Ball in J. Linn. Soc., Bot. 16: 553 (1878). ∫ C. dichotoma ssp. kremeri (Boiss. & Reut.) Nyman, Consp. Fl. Eur. 477 (1879). ∫ C. afra var. parviflora (Ball) Font Quer in Mem. Real Acad. Ci. Barcelona 25: 342 (1936). Ind. loc.: ‘Hab. in rupestribus ad mare et septentrionem spectantibus promontorii Mers el Kebir propè Oran (Aprili 1849, Boiss. & Reut).’ Lectotype: (designated here). [Algeria], Oran, montes supra Mers el Kbir, iii- 1849, Boissier & Reuter s.n. (G). = C. decipiens Schult. in Roem. & Schult., Syst. Veg. 5: 142 (1819). Ind. loc.: ‘In Calabria’ (no authentic material found). = C. dichotoma var. brachiata A. DC., Monogr. Campan. 237 (1830). Ind. loc.: ‘Varietas b in Mauritaniâ (Thibaud! Salzm!)’. Lectotype: (designated here) Mauritania, 1815, Thibaud s.n. (G) = C. dichotoma var. pallida Maire in Bull. Soc. Hist. Nat. Afrique N. 23: 199 (1932). ∫ C. afra var. pallida (Maire) Dobignard in Candollea 47: 455 (1992). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Description: Annual 5–30 cm, hispid. Stem erect, simple or dichotomously usually branched in the upper part, rarely from base. Leaves 5–40 ¥ 4–15 mm, sessile, entire or nearly so, ovate to oblong, obtuse or subacute. Flowers in groups of 2–3. Pedicels hispid, hairs 1.5–2.5 mm long. Calyx 6–16 ¥ 2–3.8 mm (8– 17 ¥ 3–4.5 mm in fruit), rather densely hirsute, hairs 1.6–2.5(3) mm long; lobes 4–10 mm long (12 mm long in fruit), ovate to lanceolate, acute; appendages 2– 3 mm long (2,5–3 mm long in fruit), ovate to rounded, obtuse, concealing ovary in fruit. Corolla 10–22 mm long, divided to 1/3–1/2, tubular-campanulate, glabrous or hairy on nerves outside, light or deep violet; lobes 3–7 mm long, ovate to triangular-ovate, obtuse; tube 7–15 mm long. Stamen 6.5–8 mm long; filiform part of filament 0–0.2 mm long; base 1.5–1.9 ¥ 1.3– 1.7 mm, oblong-elliptical, sparsely hairy in the margin, hairs 0.2 mm long; anthers 5–5.8 mm long; pollen 20–22 mm diameter, with a intermediate density of spinules (0.8–1.1 spinules mm-2). Style 8.4–11 mm long, included. Stigma 1.5–2 ¥ 0.5–0.7 mm. Capsule 5–7 ¥ 5–6.5 mm, concealed by the acrescent connivent calyx lobes and appendages, hairy on the keels (hairs 1.5–2 mm long) and on the valves (hairs 0.2–0.4 mm long). Seeds 1 ¥ 0.4–0.45 mm, yellowish. Mezleh à Mà Màs, 15.iv.1906, Aaronschii s.n. (MPU); Shomron, outlet of W. Fari’a, 6.iv.1927, Eig et al. s.n. (MPU); Jerusalem, Messallebe, 3.v.1933, Amdursky s.n. (W); Wadi Beidan near Nablus, 23.iv.1942, Davis 4531 (G, W); Nebi Youn, 29.v.1950, Mouterde s.n. (G); Jerusalem, Romema, v.1954, Barkay s.n. (MA 203052); Naplouse, Jinsafoud, 6.iv.1955, Kasapligil 2460 (G); SYRIA: Nebi Youn, 29.v.1950, Mouterde s.n. (G); TURKEY: Syria, prope Alexandrette (Iskenderum), 1834, Montbret 18 (W). REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX Ind. loc.: ‘Hab. in lapidosis calcareis Imperii maroccani austro-occidentalis Agadir-n-Ighir; prope Herculis Promontorium.’ Lectotype: (designated here) Morocco, Agadir-n-Ighir, 13.iv. 1931, Maire s.n. (P) = C. occidentalis Y. Nyman in Willdenowia 20: 113 (1991). Ind. loc.: ‘Canary Islands, Tenerife: Masca, below the village, about 500 m, 12.3. 1982, Lindblad s.n.’ Holotype: Canary Islands, Tenerife: Masca, below the village, 500 m, 12.iii. 1982, Linblad s.n. (UPS). Description: Annual 5–30(57) cm, hispid. Stem erect, dichotomously branched in the upper part, rarely from base. Leaves 6–30(60) ¥ 3–15(31) mm, sessile, entire or nearly so, ovate to oblong, obtuse or acute. Flowers generally in shortly pedunculate dichasia. Pedicels hispid, hairs 1.3–2.2 mm long. Calyx 6.6–15(21) ¥ 1.5– 4.2 mm (7.8–21.5 ¥ 3–5 mm in fruit), rather densely hirsute, hairs 1.1–2 mm long; lobes 4–10(15) mm long (5.5–15 mm long in fruit), ovate-lanceolate to lanceolate, acute; appendages 1.8–4(6.5) mm long [2.3– 4.5(6.5) mm long in fruit], rounded or ovate, obtuse, usually concealing ovary in fruit. Corolla (5)10–20(30) mm long, divided to 1/4–1/3, tubular-campanulate, glabrous or hairy on nerves outside, light violet; lobes 3–10 mm long, broadly elliptical to obovate, obtuse; tube 5–15 mm long. Stamen 5–9.2 mm long; filiform part of filament (0.5)0.7–1(1.5) mm long; base 1.4– 2.8 ¥ 1–2.3 mm, triangular, triangular-ovate to broadly obovate, hairy in the margin, hairs 0.1– 0.2 mm long; anthers (2.4)4–5(6) mm long; pollen 22–29 mm diameter, with a low density of spinules (0.24–0.27 spinules mm-2). Style (4.9)7–11(15) mm long, included. Stigma 0.9–2.4 ¥ 0.5–1.4 mm. Capsule 5–9.7 ¥ (3)4.9–7 mm, concealed by the acrescent connivent calyx lobes and appendages, hairy on the keels and on the valves (hairs 0.5–1.7 mm long). Seeds 0.7– 0.9 ¥ 0.3–0.5 mm, yellowish. Chromosome number: 2n = 24 (Gadella, 1962, 1964; Thulin, 1976; Contandriopoulos, 1980, 1981). Area: Western Mediterranean region (Morrocco, Algeria, Tunisia, southern Iberian Peninsula, Ibiza in Balearic Islands, Sicily and southern Italy) and Macaronesia (Canary Islands). We have seen only one specimen from the Dalmatian coast, with an imprecise label, which generates some doubt about its presence in Yugoslavia. Habitat: Fields, steppes and stony places; 0–1400 m Phenology: Flowering March – July Illustration: Sáez & Aldasoro (2001: 128) Notes: Campanula dichotoma is considerably variable in size, leaf shape, and flower size. We have cultivated this species from seeds, and in our opinion the floral variability is associated with pollination syndrome. Well developed plants in nutrient-rich sites are usually allogamous and show larger flowers, but at the end of the flowering period these plants usually possess only small, presumably autogamous, flowers. In poor or dry sites sparse plants are found, with only small flowers. Plants with smaller corollas were described as C. kremeri while those with larger corollas were included in C. afra or C. dichotoma. Later, a plant of the Canary Islands was described as C. occidentalis (Nyman, 1991). It has a small corolla, seeming closely related to C. kremeri. We have not been able to find any constant morphological feature suggesting that plants of the Canary Islands merit recognition as a separate taxon. We have seen the types of all these species and varieties and after studying them all features suggest that they belong to one variable species, specialized to colonize new habitats and exhibiting a great variability in reproductive behaviour. Representative specimens examined: ALGERIA: Oran, iv.1839, Bové s.n. (W); Oran, coteaux argillo-calcaries, 24.v.1851, Durando s.n. (W); Rochers du Djebel Santo à Oran, 20.iv.1852, Balansa s.n. (W); rochers à LallaMaghrina, ouest de la province d’Oran, 19.v.1856, Bourgeau s.n. (G, MPU); Montagne de la Bouzardah près Alger, 25.v.1859, Romain s.n. (MPU); in sterilibus cap Falcon, Oran, v.1856, Munby s.n. (MPU); environs d’Alger, iv.1861, Bourlier s.n. (MPU); Bou-Saada, sud de la prov. de Constantine, Algérie, 1865, S. Rebaud s.n. (W); Mégrin, env. de Tlemcen, 20.iii.1872, Courcière (MPU); In umbrosis ad Mustapha, 5.vi.1894, Chevallier s.n. (COI); Alger, v.1899, Gandoger s.n. (MA 120944); Le Gouraya de Bougie, Kabylie, vi.1896, Reverchon s.n. (COI, MPU); Santa Cruz, Oran, 17.iv.1908, Faure s.n. (MA 120938); Djebel Santo, Oran, v.1915, Alleizette s.n. (G); El Biar, près Alger, coteaux, 12.vi.1916, Bianor s.n. (MA 120942, MA 120947); Oran, iv.1921, Alleizette s.n. (MA 120969); Beni Saf, Oran, 1.v.1934, Maire & Wilczek s.n. (MA 120934); Oran, montagne des Lions, 25.iv.1934, Faure s.n. (MA 120978); Wilaya, Tizi Ouzou, Djurjura, 10 km E Yakouren, 930 m, 14.vi.1984, Podlech 39292 (G); © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 = C. leptosiphon Pau & Sennen in Sennen, Diagn. Nouv. 195 (1936). Ind. loc.: ‘Hab.-Maroc: Atlas Rifain, Targuist, à BabIzugar, 1230 m. Leg. Sennen et Mauricio.’ Lectotype: (designated here) Morocco, Atlas Rifain, Targuist à Bab-Izugan, 1230 m, 20.vi. 1933, Sennen & Mauricio s.n. (MA 121002). Isolectotypes (G, MAF 2911). 237 238 L. SÁEZ and J. J. ALDASORO 120965); Beni-Bufrah, 24.v.1934, Sennen & Mauricio s.n. (MAF 61348); Anti Atlas, El Arba des Aït Baha, 10.iii.1935, Gattefossé s.n. (MPU); Beni-Snassen, collines de Tahil, 28.v.1935, Sennen & Mauricio s.n. (MA 161405); El Kansera, iii.1937, Gattefossé s.n. (MPU); Boulhaut, Quercetum 10.v.1937 Gattefossé s.n. (MPU); Anti Atlas occid., 30.iii.1954, Rungs s.n. (RAB 4886, COI); Sefrou, 1.vii.1958, Giscard 40 (RAB 5167); Plateau Central, El Khakarake, Takesbir, 8.vi.1963, Mathez s.n. (RAB 44707); Tanger in monte Gibel Kibir, Durand s.n. (MPU); Beni Ersine, Jbel Tasiat, 35 km von Ketama, 1330 m, 8.vii.1971, Dittrich 1221a (G); Beni Mellal, entre le col du Tizi Mlil et Beni Mellal, 1250–1300 m, 5.vi.1980, Charpin et al. (G); Entre Oulad M’Barek y Ouaouizarthe, cerca de Beni-Mellal, 800 m, 12.vi.1982, Fernández Casas 6752 (MA 430481); 4 km de Boujad, cerca de Kasba-Tadla, 710 m, tomillar ralo sobre calizas, 12.vi.1982, Fernández Casas 6741 (MA 430480); Agadir, Col de Kerdouss, 930 m, 24.v.1985, Blanché et al. 9237 (MA 340938); province d’Agadir, rive gauche de l’Oed Emdel 3 km en amont S de Sidi Ifni, 16.iv.1989, Jacquemoud 4006 (G); prov. Taza, 4 km S Sidi Abdallades-Rhiata, 400 m, 10.v.1989, Podlech 46391 (G, MA 472213). SICILY: Cefalu, Siciliae septentrionalis, 14.vi.1840, Heldreich s.n. (G); ad colles propè Sta. Agostina Piana dei greci, 25.v.1855, Huet du Pavillon s.n. (MPU); Palermo a San Martino, v.1879, Todaro s.n. (P); Prov. Messina, Galatis, 1.v.1898, Rigo 209 (P, MA 120943); Messina, v.1901, Ross s.n. (BC 39111); Taormina, 100 m, 26.iv.1926, Spencer s.n. (BC 39113); Taormina, 300 m, 23.v.1933, Bornmüller 430 (P); Palermo a Gilbrossa, Todaro s.n. (MPU). SPAIN: Almería, in rupestribus montis Mugron, v.1890, Porta & Rigo s.n. (MPU); Ronda, broussailles, 21.v.1902, Dominguez s.n. (BC 39093); Almería: Barranco del Caballar, 2.v.1921, Gros s.n. (MA 120937); Murcia, s.d. Bernardé s.n. (G); Cádiz: Provincia de Cádiz, 1925, Gros s.n. (MA 190501); Málaga: In quercetis dumetisque Casares; Monte del Duque, 14.v.1932, Vicioso s.n. (MA 120961); Sierra Bermeja, alcornocales sobre silúrico, 19.v.1969, Rivas Goday & Izco s.n. (MA 219731); Benahavis, Sierra Palmitera, in rupestribus peridotiticis, 4.vi.1978, Fernández Casas 2289 (MA s.n); Murcia, 12.iv, Trèmols s.n. (MAF 2778); Coto de Alquerías, 19.v.1902, Ibáñez Jiménez & Pau s.n. (MA 120936); cerca de Cehegín, sierra de Quipar, 19.vi.1975, Fernández Casas 2289 sn. (MA 411999); in cistetis supra Estepona, v.1937, Boissier s.n. (P).TUNISIA: c. Temram, 2.v.1884, Letorneux s.n. (P); Tunis, 1922, s.d. (MPU); Jendouba, Tabarka, dunas litorales, 28.v.1992, Benedí et al. s.n. (MA 557453, MAF 148325). YUGOSLAVIA: Dalmatia, Petter s.n. (G). 13. Campanula cecilii Chitt., Gardeners’ Chronicle Ser. 3, 89: 397 Fig. 210(1931) © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Constantine, s.d. Joly s.n. (MPU). BALEARIC ISLANDS: Ibiza, Sommet du Puig d’en Serra, 28.iv.1852, Vigneix s.n. (MPU); Ibiza, Puig des Molins, 9.v.1918, Gros s.n. (BC 39096); Ibiza, Cala Jondal, 16.v.1918, Gros s.n. (MA 120959, BC 39095); Ibiza, Sant Miquel, 19.v.1919, Font Quer s.n. (BC 39094); Ibiza, Punta Corb Marí, 23.iv.1988, Aloina et al. s.n. (Herb. University Illes Balears); Ibiza, Es Vedrà, 19.iv.1991, Bibiloni s.n. (Herb. University Illes Balears). CANARY ISLANDS: Barranco Hondo près Santa Cruz de Tenerife, ii.1852, Bolle s.n. (MPU); Tenerife, 1852, Bolle s.n. (W); Teneriffa ad barranco Santo près Santa Cruz, 16.iv.1855, Bourgeau s.n. (P, W); Teneriffa, Las Mercedes, 4 km NE La Laguna, c. 600–700 m, 13.iv.1968, Ehrendorfer et al. s.n. (W); Alrededores de Sta. Cruz de Tenerife, 28.iii.1879, MasFerrer s.n. (BC 39110); Jandia, 80 m, 6.iii.1969, Kunkel 12580 (G); Lanzarote, road to Orzola, 26.iv.1972, Kunkel s.n. (G); Lanzarote, Malpais de la Corona, 150 m, 14.iii.1973, Kunkel s.n. (G). ITALY: Près de Napoles, 1860, Cappelli s.n. (G); Insula Inarime, Ischia, Casamicciola, ix.1875, Levier s.n. (P); Calabria, 19.iv.1877, Huter et al. s.n. (MPU, P); Lasla (prov. di Cosenza), Calabre, Italie, 13.vi.1893, St-Lager s.n. (MA 165674); Isola di Capri, vi.1905, M. Guadagno s.n. (MA 120948); Cava dei Tirreni (Salerno), vi.1913, Guadagno s.n. (MA 120949); Campania, ad muros humidis in umbrosis, Gragano (Napoli), 10.vii.1913, Pellanda s.n. (MA 120946); Salerno, Ravello, 28.vii.1921, Fiori s.n. (MAF 61063). MOROCCO: Tanger, vi.1839, Salzmann s.n. (COI); Djebel Amsiten, environs d’Agadir, 17.v.1877, Ibrahim s.n. (MPU); Djebel Habrid, prov. Chiadma, 1886, Ibrahim & Grant s.n. (MPU); Rabat, 1.iii.1888, Grant s.n. (MPU); Ceuta, v.1907, Vicioso s.n. (MA 120980); Environs de Larache, v.1910, Vaucher s.n. (G); Chaouia, Boulkaut in aridis, 15.v.1912, Pitard s.n. (G); Casablanca, Aui Seba, 19.v.1912, Pitard 1838 (P); Settat, El Bahloul, 8.vi.1912, Pitard 1837 (P); Telata de Reisana, v.1918, Dantín s.n. (MA 120939); Meknes, 12.vi.1918, Benoist s.n. (P); Volubilis, 23.iv.1920, s.d. (MPU); Marrakech, Djebel Guéliz, 525 m, 4.iv.1921, Rodié s.n. (MPU); Tetouan, v.1921, Alleizette s.n. (P); Xauen, 14.v.1921, Vidal López s.n. (MA 120941); Tetuán, v.1921, Pau s.n. (MA 120973); Tánger, 2.v.1921, Pau s.n. (MA 120970); Immouzer, Moyen Atlas, 1500 m, 19.vi.1923, Jahandiez s.n. (MPU); Larache, 28.vi.1923, Caballero s.n. (MA 163049); Boulhaut, gorges calcaires de l’oued Cherrat, 300 m, 21.iv.1924, Jahandiez s.n. (MA 120974); Berkane, Zegzel, 18.v.1928, Faure s.n. (G); pr. Xauen, in arenosis, 500 m, solo siliceo, 23.v.1928, Font Quer s.n. (MA 120981); Gurugú, coteaux, 100 m, Barranco del Lobo, 21.vi.1930, Sennen & Mauricio s.n. (MA 120967); Yacinem, v.1930, Mauricio s.n. (MA 120975); Segangan, Beni-Sidel, 17 et 24.v.1934, Sennen & Mauricio s.n. (MA 120977, COI); Base del Monte Tamarrut, Ifni, 700 m, 6.vi.1934 Caballero s.n. (MA REVISION OF CAMPANULA SUBGEN. SICYOCODON AND MEGALOCALYX Description: Annual 10–30 cm, hispid. Stem erect, simple or dichotomously branched in the upper part, or from base. Leaves 10–40(60) ¥ 4.2–25 mm, sessile, entire or nearly so, elliptical to oblong-lanceolate, obtuse. Flowers in shortly pedunculate dichasia. Pedicels hispid, hairs 0.9–2 mm long. Calyx 16– 23 ¥ 1.9–3 mm (26.8–29 ¥ 4–4.5 mm in fruit), hairy along the margin and on nerves, hairs 0.9–1.7 mm long; lobes 12–18 mm long (20–22 mm long in fruit), lanceolate, subacute; appendages 3.8–6 mm long (6.8– 7 mm long in fruit), ovate, obtuse, usually concealing ovary in fruit. Corolla 14–35 mm long, divided to 1/3–1/ 2, tubular-campanulate, glabrous or hairy on nerves outside, light violet; lobes 7–13 mm long, triangularovate, obtuse; tube 7–23 mm long. Stamen 6–8.4 mm long; filiform part of filament 0.8–1 mm long; base 1.5– 2.3 ¥ 1.2–2.2 mm, subpentagonal to suboblong, hairy on the abaxial surface and on the margin, hairs 0.1– 0.3 mm long; anthers 3–5.1 mm long; pollen 19–24 mm diameter, with a low density of spinules (0.70–0.75 spinules mm-2). Style 9–11.2 mm long, included. Stigma 1.7–2 ¥ 1–1.5 mm. Capsule 11–12.5 ¥ 9–10 mm, concealed by the acrescent connivent calyx lobes and appendages, sparsely hairy on the keels (hairs 0.6– 1.8 mm long). Seeds 1.2–1.5 ¥ 0.4–0.5 mm, yellowish. Chromosome number: 2n = 20 (Sugiura, 1940, 1942). Area: Northern Iran, northern Iraq and Turkey Notes: Damboldt (1978) subsumed this species in C. reuteriana because they only differ in the shape of the calyx lobes. However, in our opinion they can be separated via other useful characters. The main one is stamen shape: the length of filiform part of stamen is different, being 0.8–1 mm long in C. cecilii, while in C. reuteriana it is very short or absent (0–0.1 mm long). The shape of the stamen base is also different: winged only in C. cecilii (Fig. 20). Moreover, the size of corolla differs in both species (Fig. 9) and C. reuteriana presents two types of hairs in the pedicels and capsules, while C. cecilii only presents one. In addition, C. propinqua [unranked] grandiflora Milne-Redh. should be referred to Campanula cecilii. Representative specimens examined: IRAN: W Iran, in monte Takhi.Soleiman, v.1898, Strauss s.n. (W); Bakhtiari, Seghahan, 25.iv.1910, Schiman.Czeika 15010 (W); Zagros mountains, 51 miles W of Shirez, 23.iv.1973, Hewer 1919 (W); Luristan, 60 km W of Khorramabad, 30.iv.1966, Archibald 1622 (W); Kurdistan, 24 km N.W. of Karind, 13.v.1966, J.C. Archibald 1844 (W 17264); Kurdistan, 75 km W. N.W. of Sanadaj towards Marivan: valley of the Gul.i.Chida, 17.v.1966, Archibald 1981 (W); Lorestan, Pol.e Ma’luman c. 60 km from Khorramabad on road to Andimeshk, 5.v.1975, Wendelbo & Assadi 15576 (W). IRAQ: Gotvend, rio Karum, 5.vi.1889, Jiménez de la Escalera s.n. (MA 120997); Harran, Mesopotamia, 17.v.1865, Haussknecht s.n. (W); Assyria orientate: in montis Kuh.Sefin reg. Infer. Infra pagum Schaklava (Erbil.), 8.v.1893, Bornmüller 1554 (P, W); Distr. Sulaimaniyah (Kurdistan), Mt. Avroman ad confines Persiae, Tawilla, Sosakan, 15–18.vi.1957, Rechinger 10131 (G); near Silwan River, 7 km S of Derbenikhan, Sulaimaniyah, Liwa, 26.iv.1963, Barkley et al. 5110 (W); Diyala river, 30 km south of Durbendikan, Kirkuk, 9.iv.1964, Barkley 7356 (W). TURKEY: Kurdistania, Mardin, Senar, 6.vi.1888, Sintenis 921 (W); Siirt: Sirnak to Gizre, 14 km from Sirnak, 8.v.1966, Davis 42678 (W); NW of Mardin, Diyarbakir, 30.v.1983, Sorger 83162 (W); Excluded names C. bicolor Boiss., nom. nud., in sched. (G) C. valentina Pourr., nom. nud., in sched (MAF). C. obtusiuscula Pau, nom. nud., in sched (MA) Campanula afra f. cleystogama Font Quer, Iter Marocc. 1927, n∞. 636 (1928) nom. nud., in sched. C. dichotoma f. longipedunculata Font Quer, nom. nud., in sched. (BC) Habitat: Steppes, meadows. Phenology: Flowering March – May Illustration: Fig. 25 ACKNOWLEDGEMENTS This work was partially supported by a research grant ‘Flora iberica VIII’ (DGICYT PB 96–0849). L. Sáez has © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 215–241 Downloaded from https://academic.oup.com/botlinnean/article-abstract/141/2/215/2433659 by guest on 14 June 2020 Ind. loc.: ‘The several pot plants of this half-hardy annual were raised from seeds collected in Kurdistan, Iraq, in June, 1930.’ Lectotype: (here designated) Gardeners’ Chronicle Ser. 3, 89: 411 fig. 210. Epitype: (here designated) [Iraq], Assyria orientate: in montis Kuh-Sefin reg. Infra pagum Schaklava (Erbil), 8.v.1893, Bornmüller 1554 (P, W). Campanula cecilii Rechinger & Schimann-Czeika, in Flora Iranica 13: 12 (1965), nom. illeg. Ind. loc.: ‘It occurs from Kurdistan from 200 m to 800 m.’ Lectotype: (designated by Rechinger & SchimannCzeika, in Flora Iranica 13: 12, 1965): Bot. Mag. 157, 349 (1934). = C. propinqua [unranked] grandiflora Milne-Redh., Bot. Mag. 157: pl. 9349 (1934). Ind. loc.: ‘Armenia and Iraq (Kurdistan)’. Lectotype: (here designated) Bot. Mag. 157, 349 (1934). 239 240 L. SÁEZ and J. J. ALDASORO F A B 2 mm 1 mm 10 mm 2 cm 10 mm 10 mm D C Figure 25. Campanula cecilii. A–F: Iraq: in montis Kuh, sefin, infra pagum Schaklava (Erbil), 8.v.1893, Bornmüller 1554 (W). 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