Bulletin of The Natural History Museum Botany Series THE NATURAL HISTORY MUSEUM VOLUME 32 NUMBER 2 28 NOVEMBER 2002 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Prof. R. Bateman Editor of Bulletin: Mrs M.J. Stafford Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review before acceptance. 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Copyright © The Natural History Museum, 2002 ELECTRONIC ACCESS This journal is included in the Cambridge Journals Online service which can be found at: http://journals.cambridge.org For further information on other Press titles access http://uk.cambridge.org or http://us.cambridge.org World list abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) ISSN 0968-0446 The Natural History Museum Botany Series Cromwell Road Vol. 32, No. 2, pp. 61-156 London SW7 5BD Issued 28 November 2002 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Bull. nut. Hist. Mas. Land. (Bot.) 32(2): 61-123 Issued 28 November 2002 Studies in the genus Hypericum L. (Guttiferae) 4(2). Section 9. Hypericum sensu lato (part 2): subsection 1. Hypericum series 1. Hypericum NORMAN K.B. ROBSON tfSSfSSSL Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS 17 SEP ?nn* A. Sect. 9. Hypericum 61 Subdivision 61 Characters and variation 61 Distribution and evolution 62 Sect. 9. Hypericum subsect. 1. Hypericum series 1. Hypericum 62 Characters and variation 62 Hybrids 64 Distribution and evolution 65 Systematic treatment 66 Sect. 9. Hypericum 66 Sect. 9. Hypericum subsect. 1. Hypericum series 1. Hypericum 67 References ' 19 Systematic index 121 SYNOPSIS. The subdivision of the large sect. 9. Hypericum and four of its segregated sections having been treated in Part 4( 1 ), Part 4(2) is concerned with those members of sect. 9, viz. Hypericum sensu stricto, in which the stem internodes have glandiferous raised lines - subsect. Hypericum series Hypericum. The series comprises 1 1 species and can be divided into (i) a group centred in Europe and the Mediterranean (4 species) and (ii) one centred in north-east Asia (6 species), one species of which is confined to western North America and another has spread westward into Europe. The remaining species (H. perforation) is morphologi- cally and geographically intermediate between two taxa, one from each group, and behaves as an allotetraploid hybrid. It is divided into four subspecies: subspp. perforatum, songaricum (Ledeb. ex Rchb.) N. Robson stat. nov., veronense (Schrank) H. Lindb. and chinense N. Robson subsp. nov. The hybrids of H. perforatum are treated in detail and include H. x desetangsii nothosubsp. balcanicum N. Robson, nothosubsp. nov. (H. maculatum subsp. immaculatum x perforatum). Other new hybrids proposed are: //. x laschii nothoforma froelichii N. Robson, nothoforma nov. (//. maculatum subsp. obtusiusculum x tetrapterum), H. undulatutn x tetrapterum and H. elegans x perforatum. INTRODUCTION In Part 4( 1 ) the relationships of sect. 9. Hypericum sensu lato were discussed and it was shown that, although the whole group is apparently derived from sect. 7. Roscyna, it consists of six independ- ently derived sections, one from each of the three subspecies of H. ascyron and three from H. przewalskii (Robson, 2001: fig. 2). The latter three and sect. 9a. Concinna (related to H. ascyron subsp. gebleri) were treated in Part 4( 1 ), this part is concerned with most of sect. 9. Hypericum (related to H. ascyron subsp. ascyron), and Part 4(3) will contain the remainder of Sect. 9 and the American sect. 9b. Graveolentia (related to H. ascyron subsp. pyramidatum). Sect. 9. Hypericum Subdivision Sect. 9. Hypericum, which is distributed from the Azores through Europe, the Mediterranean and western and northern Asia to China, Japan and western North America, comprises two subsections: subsect. 1 . Hypericum and subsect. 2. Erecta. The latter is almost confined to Japan, Taiwan and eastern China, whereas subsect. Hypericum is widespread. In subsect. Hypericum the stems are almost always lined and the base is stoloniferous, whereas in subsect. Erecta the stems are terete and the base is woody or fibrous. Subsect. Hypericum, in turn, can be divided into two series: series 1 . Hypericum (with stem lines persistent and bearing dark glands) and series 2. Senanensia (with stem lines weak and eglandular or absent). Series 1 is widespread, series 2 confined to northern and central Japan. Characters and variation The two subsections of sect. Hypericum each have primitive characters that clearly relate them individually to H. ascyron subsect. ascyron, but neither can be derived from the other. Thus the primi- tive species of subsect. Hypericum (H. maculatum), along with the closely related H. undulatum and H. tetrapterum, has markedly 4- lined stem internodes, like H. ascyron; but the group as a whole is advanced in that the stem lines are glandiferous and the stems have creeping bases (i.e. they are stoloniferous). In contrast, the primitive © The Natural History Museum. 2002 62 N.K.B. ROBSON H. ascyron subsp. ascyron [W. Siberia to Japan, China] 2-4. H. undulatum group [Macaronesia, Europe Mediterranean, W. Asia] Spp. 20-42 [Sakhalin to Borneo] 1 . H. maculatum s.l. [Euro-Siberia] ser. 2. Senanensia Spp. 13-19 [Japan, Korea, C. China] 5. H. perforation [Eurasia, Med.] l.H. elegans [C. Siberia to C. Europe] 12. H. scouleri [W. North America to C. Mexico] 6. H. altenualum [C. Siberia,.China, Korea] 8-1 1 . H. yezoense group [Japan, Sakhalin, S. Kuriles] Fig. 1 Section 9. Relationships and distribution. taxa of subsect. Erecta (H. yamamotoi var. riparium Y. Kimura and H. ovalifolium Koidz.) have a branching woody stem base, like H. ascyron, but the stem internodes are slightly 2-lined or terete, the lines being eglandular. Also, the leaves in the H. yamamotoi and H. erectum groups of subsect. Erecta are frequently triangular-lanceo- late, like those of//, ascyron subsp. ascyron, whereas a comparable form of leaf occurs only apomorphically in subsect. Hypericum, i.e. only in some of the more advanced species. Distribution and evolution (Fig. 1) The distributions of subsect. 1 series 2 and subsect. 2 are wholly north and east Asian, but that of subsect. 1 series 1 extends from the Azores to Mexico. The latter series comprises a western group, based in south-eastern Europe on 1. H. maculatum and extending west to the Azores and east to central Siberia, and an eastern group with two centres, one in eastern Siberia and the other in western North America. The North American centre consists only of 12. H. scouleri, which has its most primitive form in the south-eastern United States and shows trends northward to central British Colum- bia and southward to Michoacan in central Mexico. The basic species of the east Asian centre is 6. H. attenuatum, which has its most primitive form in eastern Siberia and a range that extends westward to the Altai region and southward to south-central China. The basic species of the derivative Japanese H. yezoense group (H. tosaense) is morphologically very near H. attenuatum and has been included in it (Kitamura & Murati, 1962), but in my opinion it is specifically distinct. Hypericum elegans, a western derivative of H. attenuatum, extends the distribution of the eastern group as far as western Germany. The remaining species in series 1,5. H. perforatum, does not fit easily into the above scheme. Morphologically it is intermediate between 1 . H. maculatum and 6. H. attenuatum, and the areas of these species almost overlap in western Siberia (Maps 1, 16). Each of them is diploid (2n=16), whereas H. perforatum is tetraploid (2n=32); so it is conceivable that H. perforatum is an allotetraploid derivative of a cross between them, especially as H. perforatum behaves cytologically as a hybrid (see p. 64). However, the nearest subspecies of H. maculatum to H. perforatum morphologically is not the one that occurs in Siberia today (subsp. maculatum) but subsp. immaculatum, which is now confined to the Balkan region. The hypothesis must therefore be that subsp. immaculatum at some time was present in Siberia, where it hybridized with H. attenuatum. The chromosomes of the resultant diploid hybrid then doubled, giving rise to an allotetraploid species that is partially fertile (see p. 88). Subsequently the area of//, maculatum subsp. immaculatum became greatly reduced; but that of subsp. maculatum, which is apomorphic to it and thus a likely derivative of it, extended eastward into Siberia. Sect. 9. Hypericum subsect. 1. Hypericum series 1. Hypericum Characters and variation (Figs 2, 3) MORPHOLOGY. The two main divisions of series Hypericum could be separated on whether or not the subsidiary stem lines were present, i.e. whether the stem internodes were 4- or 2-lined, were it not for (a) 4. H. triquetrifolium, which has 'lost' them in the 4-lined group and (b) 5. H. perforatum, which, as has been explained, appears to be a hybrid with one parent from each group. Figure 3 shows that the 4-lined state is primitive in this series, as indeed it is in the whole genus. Intermediate states occur in hybrids of H. perforatum, but not in the species. The laminar leaf glands are absent or more rarely sparse in l.H. maculatum but more or less dense in the more advanced species. They are all or mostly pale except in 6. H. attenuatum, in which black glands often predominate. The sepals are broad with a rounded apex in 1 . H. maculatum and usually in 12. H. scouleri, as they are in the ancestral (?) H. ascyron subsp. ascyron (sect. 7. Roscyna). In all the other species they are acute and narrow (Spp. 2, 3 and 5) or acute and broad (6. //. STUDIES IN HYPERICUM 63 3 a. H. tetrupterum var. anagallidifolium 3b. H. tetrapterum var. corsicum ~~~~~~~~~~~, 3. H. tetrapterum var. tetraplerum 16 4. //. triquetrifolium 16 2b . //. undulalum var. boeticum Za. //. undulatum var. undulatum 16,32 la //. maculatum .ssp. immaculatum (16) \ 1 b. //. maculatum ssp. maculatum 16 1 c. //. maculatum ssp. obtusiusculum 32 5d. //. perforatum ssp. chinense Fig. 2 Section 9, subsect. 1 , series 1 . Relationships and 2n chromosome numbers. 12. //. scouleri 16 1 1 . //. yezoense 1 0. //. momoseanum I 9. H. iwatelittorale 6. //. attenuatum 5. //. perforatum 16,32f48 5a.//. perforatum ~ ssp. perforatum 5b. //. perforatum 32, 48 ssp. songaricum 5c. //. perforatum ssp. veronense 32 [16] -8. //. tosaense 1. H. elegans 32 attenuatum) or acuminate and broad (Japanese spp.) except 4. //. triquetrifolium, in which the more reduced forms have (secondarily) small rounded sepals. The apex is entire except in 2c. //. maculatum subsp. obtusiusculum and //. x desetangsii (2. //. maculatum x5.H. perforatum), where it is usually eroded-denticulate. The petal laminar glands are primitively pale and linear to striiform (la. //. maculatum subsp. immaculatum, 12. //. scouleri), as in sect. 7. Roscyna, and evolve in two directions: (a) the lines become interrupted, first toward the apex (Spp. Ib, 5, 6, 8-10, 12) and then throughout the lamina (Spp. 2-4, 7, 11) and sometimes disappear altogether (Spp. 2, 3); (b) some pale glands become black, but these usually remain in the minority except in 6. H. attenuatum and usually Ib. H. maculatum subsp. maculatum, where all glands become black. The vittae of the ovary and capsule are primitively all narrow and longitudinal, as in sect. 7. Roscyna; but the lateral vittae become oblique, in 4. H. triquetrifolium only sporadically, in others totally (Spp. 6-11). In 5. H. perforatum alone the oblique lateral vittae first become enlarged and then break up to form oval to globose vesicles. CYTOLOGY. Although the diploid chromosome number in sect. 7. Roscyna is usually 18, it ranges from 22 to 16 (Robson, 2001); and 2n=16 appears to be the basic number for the whole of sect. 9. Hypericum. In series \.Hypericum (Fig. 2), Spp. Ib, 2-4, 6 and 12 are diploid, Sp. la has been shown by indirect evidence from morphology and hybrids to be diploid, and the Japanese species (8- 11) seem likely to be so. Tetraploidy (2n=32) has been recorded in 2. H. undulatum but without reference to morphology, in Ic. H. maculatum subsp. obtusiusculum, which appears to be an autotetraploid of la. subsp. immaculatum, in 7. H. elegans, which is possibly a direct derivative of 6. H. attenuatum, and in 5. H. perforatum, which is apparently an allotetraploid from the cross la. H. maculatum subsp. immaculatum x 6. H. attenuatum. Hexaploids (2n=48) have also been recorded in H. perforatum, presumably resulting from the fertilization of an unreduced (apomictic) ovule by a normally reduced pollen grain (see p. 88 ). The diploids reported in this species must be dihaploids resulting from polyhaploidy, i.e. the secondary occurrence of the diploid number as a result of the development of an unfertilized reduced ovule, which is known to 3c sepal apex acute to \ rounded acuminate: to obtuse 2a black petal laminar glands absent ,- Fig. 3 Section 9, subsect. 1 , series 1 . Limits of some characters. 64 N.K.B. ROBSON 4. H. triquetrifqlium 16 2 * 3. H. undulatum * tetrapterum subsect. Hypericum subsect. Erecta Spp. 2(M1 3. H. tetrapterum .2. H. undulatum series Hypericum series Scnanensia Spp. 13-19 [Svars.J 16 ~~-:::; ___ [2 vars.] 16, 32 1 xa. H. x 'laschii 1 xb. //. * laschii ~~~ nf. laschii (16) \ 1 b. //. maculatum ssp. maculatum ,6 nf. froelichii (24) 12. //. scoukri 16 la. //. maculatum ssp. immaculatum (16) ^ lc. //. maculalum ssp. obtusiusculum 6. //. attenuatitm 16 5. //. perforatum 1 1 . //. . 10. H.^momoseanum 9. //. iwatelittorale 8. //. losaense 32 5xb. H. x desetangsii nssp. carinthiacum 24,40 V5xa. //. x desetangsii nssp. desetangsii "^ -"I" [4sspp.] [16] 32, 48 7.(//. elegans gsii -"~""\. / """"-•.>. ! 5xba. nf. maculatiforme 5xbb. nf. perforatiforme 24 40 5xc. //. x desetangsii 5 * 7. //. perforatum x elegans x^ 48 --'" nssp. balcanicum -~^ (32) 40 (24, 40) 5xca. [nf. a] 5xcb. [nf. b] (24) (40) Fig. 4 Section 9, subsect. 1, series 1 . Relationships ( ) and hybrids ( ), with 2n chromosome numbers (unverified numbers in parenthesis). occur in allopolyploids (see p. 88 and Robson, 1998). The report of a triploid number (n=12) by Gagnieu & Wilhelm (1965) is likely to be an error for n=16. A hypoploid number (2n=8, n=4) has been reported for 2. H. undulatum (Guillen et al., 1997), a species which is normally diploid (2n=16) or tetraploid (2n=32). Hybrids (Fig. 4) Natural hybridization is relatively rare in Hypericum (Robson, 1981: 167, fig. 55) except in series 1. Hypericum, where it is quite common largely on account of the floral biology of H. perforatum (see p. 88). This species behaves as a hybrid (high pollen sterility, lagging chromosomes at metaphase, 97% apomictic ovule develop- ment with pseudogamy). When crossed with diploids, its reduced pollen grains and ovules (n=16) produce triploid (2n=24) hybrids, while the fertilization of its unreduced ovules by a gamete of a diploid species produces pentaploid (2n=40) hybrids. With tetraploids such as H. maculatum subsp. obtusiusculum, of course, the hybrids are tetraploid (2n=32). Figure 4 (modified and expanded from Robson, 198 1 : fig. 55) shows all the hybrids that have been reported or inferred from series 1 . Hypericum. The status of some of the hybrids has been inferred from intermediate morphology, while several chromosome numbers have been inferred from those of the parents. Of the hybrids not involving H. perforatum, 2. H. undulatum x 4. H. tetrapterum, which is known only from plants with intermediate morphology, appears to be rare. As the tetraploid form of H. undulatum has been reported only once, the hybrid is likely to be diploid. 1. H. maculatum x 4. H. tetrapterum (H. x laschii) was described as intermediate between la. subsp. maculatum and tetrapterum and is known from the area of that subspecies only. It too is therefore likely to be diploid. On the other hand, the cross with (the tetraploid) lc. subsp. obtusiusculum, described by Frb'hlich (1911) and named here H. x laschii nothoforma froelichii, should prove to be triploid. The hybrids involving H. perforatum can be triploid to hexaploid, depending on the ploidy of the other parent and of the gametes of//. perforatum itself. Aneuploid numbers occur in experimental crosses (e.g. those of Lihova et al., 2000), but they have not been reported from wild populations. Noack (1939) found that, in hybrids involv- ing diploids, the triploids were intermediate between the parents whereas the pentaploids could not be distinguished from pure H. perforatum. Martonfi et al. ( 1 996/7), however, found that a pentaploid population of H. maculatum subsp. maculatum x perforatum from Slovakia did have some intermediate characters as well as the pentaploid number (2n=40). Tetraploid crosses with H. perforatum yield intermediates, at least in the first generation. Thus a possible hybrid, 5. H. perforatum x 1. H. elegans, is recorded here on the basis of a morphologically intermediate specimen from the Ukraine; and the cross H. maculatum subsp. obtusiusculum x perforatum also yields intermediate prog- eny in the Fr The relationship of the parents of this latter cross is very close, however, as subsp. obtusiusculum behaves cytologically and morphologically as an autotetraploid of subsp. immaculatum, not of subsp. maculatum, as I first thought (Robson, 1956; 1958a, 1975). This consanguinity facilitates back-crossing and results in the eventual occurrence of a complete series of intermediates in the wild (cf. Crackles, 1990). Counts of 2n=48 and 40 for this hybrid from cultivated plants (Robson, 1956; Robson & Adams, 1968) presumably result respectively from a cross with an unreduced perforatum gamete (32 + 16) and a backcross of this hybrid to either a 'normal' hybrid or to H. perforatum itself (24 + 16). The species 3. H. tetrapterum is diploid only, so its cross with //. perforatum (H. x medium), described as morphologically inter- mediate, is likely to be triploid; but the cross 1 . H. maculatum x 5. perforatum (H. x desetangsii) is much more complicated. The hybrid with the tetraploid lc. maculatum subsp. obtusiusculum (5xa. //. x desetangsii nothosubsp. desetangsii) is also usually tetraploid and has been considered above. That with the diploid la. maculatum subsp. immaculatum (5xc. //. x desetangsii nothosubsp. balcanicum) and Ib. maculatum subsp. maculatum (5xb. H. x STUDIES IN HYPERICUM 65 desetangsii nothosubsp. carinthiacum) occurs in two forms depend- ing on the sex of the gametes. With H. maculatum pollen, triploids (2n=24) are produced (e.g. 5xba nothoforma maculatiforme) whereas with perforation pollen, the progeny are pentaploid (2n=40) (e.g. 5xbb nothoforma perforatiforme). The corresponding hybrids with 1 a. H. maculatum subsp. immaculatum have not yet been recognized or named. Distribution and evolution (Fig. 5) As has already been explained (p. 61), sect. Hypericum is most closely related to H. ascyron subsp. ascyron (Sect. 7. Roscyna), and subsect. Hypericum is distributed over most of the Paleotemperate Region from the Azores to Mexico. The distribution of series Hypericum occupies most of this area except south-eastern China and south Japan. It is divided into two groups (i) mainly western and (ii) mainly eastern. Group (i), which extends from the Azores to central Siberia, includes the nearest taxon morphologically to series 2, la. H. maculatum subsp. immaculatum, which is restricted to the central Balkan region but differs little from the north Japanese H. kamtschaticum var. pibairense Miyabe & Y. Kimura (series 2), mainly in having constantly 4-lined gland-bearing stem internodes. Subsp. immaculatum appears to have given rise northward (group iii) to two subspecies: (a) by vicariance to Ib. subsp. maculatum (Atlantic coast to central Siberia) and (b) by autotetraploidy to Ic. subsp. obtusiusculum (=H. dubium Leers) (north-west Europe). Southward there is a derivative (group iv) that divided into a western (damp habitat) subgroup (Spp. 2-3) and an eastern (dry habitat) one (Sp. 4). In group (iii) the stems are 4-lined, the leaves broad and plane and the sepals broad and rounded (entire or not); in group (iv) the stem lines become broad (3. H. tetrapterum) or the subsidiary pair disappear altogether (4. H. triquetrifolium), the leaves become undulate (2a. H. undulatum var. undulatum) and narrow (4. H. triquetrifolium) or remain broad but revert to plane (2b. H. undulatum var. boeticum, 3. H. tetrapterum), whilst the sepals become narrow and acute but, in H. triquetrifolium, revert to rounded when small. Although a Portuguese specimen of H. undulatum was observed to have a habit and narrow leaves that approach those of H. triquetrifolium, these characters would seem best interpreted as a parallel development within H. undulatum. The first division in group (iv) would then have been between the western Mediterrean/ Atlantic wet-habitat H. undulatum and the east Mediterranean/W. Asian dry habitat H. triquetrifolium. The species 3. H. tetrapterum, with broad stem lines, broader plane leaves and a more congested inflorescence of smaller flowers subsequently evolved (in the south- western Mediterranean?) from H. undulatum and spread north as far as Scotland, Denmark and southern Sweden, north-east to Poland, Latvia and the western Ukraine, and east to Iran. Group (ii) divides immediately into an eastern, Old World sub- group (v) and a western, West American subgroup (vi), both of which start with a single species, respectively the eastern Siberian and Chinese 6. H. attenuatum and the British Columbia to Mexican 12. H. scouleri. Whilst none of the variation in H. scouleri has resulted in speciation, H. attenuatum has speciated eastward and westward. To the west it gave rise to 7. H. elegans, which inhabits steppe regions as far into Europe as south Germany (Saxony, with isolated stations further west). Eastward there is a group of species in Japan of which the two southern, 8. H. tosaense (Shikoku) and 9. H. momoseanum (C. Honshu) have at one time been included in H. attenuatum. They both occur in isolated localities, as does a close relative of H. tosaense, 9. H. iwatelittorale, in northern Honshu. On the other hand, the northern relative of H. momoseanum, 1 1 . H. yezoense, is widely spread in Hokkaido and northern Honshu and also occurs in southern Sakhalin. As we have seen, 5. H. perforatum has apparently resulted from a cross between one taxon from group (iii) (la. H. maculatum subsp. immaculatum) and one from group (v) (6. H. attenuatum). It was suggested that the cross occurred in western Siberia, and the present distribution and variation of//, perforatum supports this suggestion. Both parents have sessile herbaceous leaves, and the first division (vii) is between 5a. subsp. perforatum, with shortly petiolate herba- ceous concolorous leaves and 5b. subsp. songaricum, with sessile, often subcoriaceous and discolorous leaves. Subsp. perforatum has 11 Japan (N. Honshu, Hokkaido) Russia (S. Sakhalin, S. Kuriles) 3c Lebanon Levant Europe excl. N., W. Asia Algeria SW England, SW Wales, SW Ireland? Iberian Penin., NW Africa E. Med. to NW Iran 12 W. Canada, W. USA . & C. Mexico (v)\ 6 E. Siberia, Mongolia Korea, China NW Europe E. to W. Germany, Bohemia Styria & Hungary Ib N. & C. Europe Siberia E. to Angara-Sayan N. & C. Europe S. to N. Spain, N. Italy, NE Greece, NW Turkey Siberia E. to Angara-Sayan, NW Mongolia 5d ' China 10 Japan (C. Honshu) Japan (N. Honshu) Japan (Shikoku) W. Siberia to W. Europe 5b N. Kazakhstan & NW Xinjiang S. & W. to S. Ukraine \ 5c Macaronesia, Medit. & S. Europe to E. Kazakhstan & NW India Sudan (Jebel Marra), Saudi Arabia (Asir) Fig. 5 Section 9, subsect. 1, series 1. Distribution. 66 N.K.B. ROBSON spread westward throughout northern and north-western Europe and eastward into central Siberia. Beyond Siberia it recurs, morphologi- cally scarcely altered, in China to the south-east of the Mongolian desert and steppe region and is transformed gradually south-east- ward into a narrow-leaved form with long branches and congested inflorescences (subsp. chinense). Subsp. songaricum is present in the mountains of north-eastern Kazakhstan and adjacent Kyrghizstan and Xinjiang (China); then, after a gap of over 2500 km, it recurs in southern Russia (Astrakhan, Ciscaucasus) and Krym. There, with a small transitional zone, it is replaced by 5c. subsp. veronense, which appears to have spread through Turkey westward into southern Europe, the Mediterranean and Macaronesia, south to Saudi Arabia and the Sudan Republic (Jebel Marra - if truly native there, see Wickens, 1976a, \916b: 98), and east to the western Himalaya and Central Asia. Distributions of species and species groups in series Hypericum show few wide disjunctions, but those that do occur are remarkable. The widest is in group (ii), where the eastern Siberian centre of variation (i.e. primitive form) of H. attenuatum is now separated from that of H. scouleri in Arizona and New Mexico by the north Pacific Ocean. This distribution would seem to support the inference by Hui-lin Li (1952) that plants with eastern Asia - western North America disjunctions are predominantly herbaceous; and it is thus also likely to support his other inference, that such disjunctions are more recent than those involving eastern North America. Hyperi- cum scouleri is a member of the Arcto-Tertiary element of the Californian flora (Axelrod, 1959; Raven & Axelrod, 1978), where it occurs in the Yellow Pine (Pinus ponderosa Dougl. ex P. & C. Lawson) forests of the Sierra Nevada (Munz & Keck, 1959). It is possible, therefore, that H. scouleri has been in western North America since the Miocene. The other notable disjunction has already been mentioned. It would appear that H. maculatum subsp. immaculatum, now con- fined to the central Balkans, once had a distribution extending into western (or even central) Siberia, where it crossed with H. attenuatum and gave rise to H. perforatum. Its close resemblance to H. kamt- schaticum var. pibairense, of northern Japan, implies that subsp. immaculatum (or its immediate ancestor) migrated south-westward (without differentiating?) and eastward with minimal differentia- tion. It has so far not been possible to estimate the geological age of these disjunctions. SYSTEMATIC TREATMENT1 Sect. 9. HYPERICUM Perennial herbs or very rarely suffrutices up to 1 .2 m tall, with stems erect to prostrate, glabrous, without or with dark (black or very rarely reddish) glands; branching lateral, from upper or most nodes. Stems 4-2-lined or -winged when young, remaining so or becoming terete, eglandular or with glands confined to the lines or not. Leaves opposite or abnormally 3-whorled, decussate, sessile to shortly pseudopetiolate, free, persistent; lamina entire with venation pin- nate, closed, with tertiary reticulation dense to lax and obscure; laminar glands punctiform or rarely shortly striiform, equal or unequal; intramarginal glands black or occasionally also pale; ven- tral glands absent. Inflorescence \-c. 70-flowered with branching dichasial-monochasial from 1^ nodes, often with subsidiary branches from lower nodes; bracts and bracteoles foliar or reduced, persistent. Flowers stellate, homostylous. Sepals 5, free or united at base, persistent, erect to recurved in fruit, with margin entire or denticulate to glandular-ciliate; veins 3-5(-7); laminar glands pale and/or dark, linear to punctiform; marginal and/or intramarginal glands dark and/or very rarely pale or rarely absent. Petals 5, persistent, erect but not twisting after flowering, without apiculus; margin entire to crenate or eroded-denticulate to ciliate; laminar glands linear to punctiform, pale and/or dark; marginal glands dark, immersed to ± prominent, or absent. Stamen fascicles '3' (i.e. united 2+2+1), persistent, with stamens totalling 20-c. 100; filaments basally united; anther gland dark or rarely amber; pollen types IV?, X. Ovary with 3(4) completely or almost completely axile placen- tae, each oo-ovulate; styles 3(4), divergent from discrete bases; stigma narrowly capitate or narrow. Capsule 3(4)-valved, chartaceous, with valves longitudinally vittate or sometimes ± later- ally vitiate to vesiculate, pale. Seeds cylindric, not carinate or appendiculate; testa scalariform-reticulate to finely foveolate. BASIC CHROMOSOME NUMBER (x). 8; ploidy 2, 4 (rarely 3, 5, 6). HABITAT. Various; 0-3 1 50 m. DISTRIBUTION. Macaronesia, Europe (except extreme north), north-west Africa, south-west Asia (mesophytic regions) east to north-west India (Uttar Pradesh), central Asia, southern Siberia, north-west Mongolia, Korea, China (except the south-east, Tibet, Qinghai and E. Xinjiang), Japan, western Canada, western U.S.A., north-west to central Mexico. Represented in many other parts of the world by H. perforatum as an introduction. 42 species in 2 subsections, one with 2 series (for infraspecific taxa and hybrids, see subsectional descriptions). Key to subsections and series of sect. 9. Hypericum 1 Rootstock herbaceous, stoloniferous; stems persistently 2^4-lined or -winged, with glands on the lines and occasionally elsewhere or eventually terete and eglandular; black laminar leaf glands few or none (sometimes numerous in Sp. 6); capsule valves sometimes with lateral oblique vesicles (subsect. 1. Hypericum) 2 Rootstock woody or fibrous, not stoloniferous; stems always even- tually terete, eglandular (sometimes except Sp. 39); black laminar leaf glands usually present; capsule valves always longitudinally vittate subsect. 2. Erecta 2 Stems persistently 2^4-lined, lines glandiferous, at least one pair prominent (sometimes weak and eglandular in Sp. 12) series 1. Hypericum Stems eventually terete (except Sp. 13), eglandular, lines weak ... ... series 2. Senanensia Subsect. 1. Hypericum. Type: H. perforatum L. Stem internodes persistently 2^1-lined or narrowly 4-winged, the lines and sometimes elsewhere black- or rarely reddish-gland- dotted (very rarely terete and often eglandular in 12. H. scouleri). Leaves with laminar glands all pale or a few (sometimes more in 6. H. attenuatum) black, punctiform, or absent. Bracts and bracteoles entire. Petals with laminar glands pale to black, linear to punctiform, or absent; marginal glands immersed or sessile. Capsule valves (i) longitudinally vittate or with (ii) dorsal and lateral vittae or (iii) vesicles with or without dorsal vittae. Seeds linear-foveolate to finely foveolate. Species 1-19. 'An asterisk (*) before a locality or after a herbarium specimen symbol indicates that the specimen has not been seen by me. Type material seen by me is indicated by an exclamation mark(!). STUDIES IN HYPERICUM 67 Series 1 . Hypericum Hypericum sect. Euhypericum subsect. Homotaenium series Attenu- ata, series Kamtschatica pro parte quoad H. yezoense, series Acuta, series Quadrangula, series Elegantia, series Crispa Gorschk. in Shishkin & Bobrov, Fl URSS 15: 236-244 (1949), nom. invalid, omnia sine descr. lat. Hypericum sect. Euhypericum subsect. Heterotaenium series Per- /orato Gorschk. in Shishkin & Bobrov, Fl. URSS 15: 247 (1949), nom. invalid, sine descr. lat. Hypericum sect. Homotaenium series Quadrangula Y. Kimura in Nakai & Honda, Nova Fl. Jap. 10:136 (1951). Type: H. quadrangulum L., nom. rejiciendum (= H. tetrapterum Fr.). Hypericum sect. Homotaenium series Crispa Y. Kimura, loc. cit. (1951). Type: H. crispum L. (= H. triquetrifolium Turra). Hypericum sect. Homotaenium series Bilineata Y. Kimura, loc. cit. (1951). Type: //. attenuatum Fisch. ex Choisy. Stems persistently 2^-lined, lines glandiferous, at least one pair prominent (sometimes weak and eglandular in Sp. 12). Spp. 1-12. Key to sect. 9. Hypericum subsect. 1. Hypericum series 1. Hypericum 1 Stem internodes completely 4-lined or 4-winged; capsule valves longitudinally vitiate (rarely interrupted to vesiculate in Sp. 4) 2 Stem internodes partially 4-lined or 2-lined; capsule valves longitu- dinally vitiate to vesiculate 9 2(1) Sepals unequal, apex rounded to obtuse or subacuminate, entire to eroded-denticulate; leaves with venation ± densely reticulate and laminar glands absent or scattered, relatively large (1. maculatum) 3 Sepals subequal to equal, apex acute to acuminate, entire; leaves with venation ± laxly reticulate; laminar glands always present, varying in size 5 3(2) Stem internodes always with complete subsidiary lines; inflores- cence branches making narrow angle (c. 30°) with stem; sepals with apex entire; petals entire, with laminar glands black and dots to short streaks or linear and pale 4 Stem internodes with complete or incomplete subsidiary lines; inflorescence branches making a broad angle (c. 50°) with stem; sepals with apex usually finely eroded-denticulate; petals usually distally unilaterally crenate, with laminar glands black, mainly lines to streaks Ic. maculatum subsp. obtusiusculum 4(3) Sepals and petals with only pale glands; petal glands lines to streaks la. maculatum subsp. immaculatum Sepals and petals with mostly black glands; petal glands dots and short streaks Ib. maculatum subsp. maculatum 5(2) Stem internodes ± equally narrowly 4-winged; leaf margin usually crisped-undulate; inflorescence lax; flowers 12-17 mm in diam.; sepals nearly always with 3-28 laminar black glands; petals bright yellow, nearly always red-tinged dorsally (2. undulatum) 6 Stem internodes ± broadly 4-winged, the principal lines wider; leaf margin nearly always plane; whole inflorescence or partial inflores- cences dense; flowers 10-15 mm in diam.; sepals without or rarely with 1-2 laminar glands; petals rather pale yellow, not red-tinged (3. tetrapterum) 7 6(5) Petals tinged red dorsally; leaves elliptic to ovate, margin crisped- undulate; plant erect 2a. undulatum var. undulatum Petals not tinged red; leaves broadly elliptic to orbicular, margin entire; plant erect to decumbent .... 2b. undulatum var. boeticum 7(5) Inflorescence 10-c. 70-flowered from 1^ nodes; stems erect 3a. tetrapterum var. tetrapterum Inflorescence 1-3-flowered from terminal node; stems procumbent to ascending 8 8(7) Leaves elliptic to orbicular, mostly 0.5 mm petiolate, margin plane; inflorescence without flowering branches 3b. tetrapterum var. corsicum Leaves elliptic to triangular-ovate, subsessile to 0.5 mm petiolate, margin undulate; inflorescence sometimes with flowering branches from up to 5 nodes below 3c. tetrapterum var. anagallidifolium 9( 1 ) Leaf margin crisped-undulate, lamina usually triangular-lanceolate; stem widely branched for most of its length, forming broad pyramid; sepals 1-3 mm long 4. triquetrifolium Leaf margin plane, lamina ovate or oblong or elliptic to linear; stem narrowly branched rarely below middle, forming at most narrow pyramid; sepals (2.5)3 mm or more long 10 10(9) Stem lines (at least) with black glands, principal lines always present; petal margin usually distally black-glandular-crenate 11 Stem lines eglandular or very rarely with a few reddish glands, principal lines rarely absent (i.e. rarely terete); petal margin entire to distally black-glandular-subcrenate 12. scouleri 11(10) Subsidiary stem lines nearly always ± developed; sepals often eglandular-denticulate; other characters to varying degrees inter- mediate between 1. H. maculatum and 5. H. perforation (see key, p. 101) 5x. x desetangsii Subsidiary stem lines absent; sepals entire; other characters not so intermediate 12 1 2( 1 1 ) Capsule valves with lateral vittae or vesicles and often dorsal vittae (5. perforatum) 13 Capsule valves with longitudinal vittae only 16 13(12) Leaves (at least on main stem) sessile; petal laminar glands usually all pale 14 Leaves all petiolate; petal laminar glands pale and/or black 15 14(13) Capsule valves with lateral vittae linear or slightly swollen overall, not interrupted; sepals finely acuminate; leaves oblong to oblong- ovate, base ± shallowly cordate-amplexicaul 5b. perforatum subsp. songaricum Capsule valves with lateral vittae swollen at base and/or interrupted to short and irregular (vesicular); sepals acute; leaves usually nar- rowly triangular-lanceolate to linear or, if broader, then short (c. 5-10 mm long), base rounded to cuneate 5c. perforatum subsp. veronense 15(13) Inflorescence and/or partial inflorescences not usually congested, branches relatively short, straight; leaves usually oblong to ovate or elliptic; at least some petal laminar glands pale 5a. perforatum subsp. perforatum Inflorescence and/or partial inflorescences congested, branches relatively long, curved-ascending; petal laminar glands all black or absent 5d. perforatum subsp. chinense 1 6( 1 2) Sepals black-glandular-ciliate or with sessile black marginal glands 7. elegans Sepals entire or with apical black gland only, without or with immersed black marginal glands 17 17(16) Sepals and petals with pale and black laminar glands; stem often with scattered black or reddish glands; sepals acute to subacuminate ....6. attenuatum 68 Sepals and petals with pale laminar glands only; stem with black glands confined to raised lines or near them; sepals acute to abruptly acuminate 18 1 8( 1 7) Leaves without black laminar glands; inner petal margin crenulate (unknown in 9. iwatelittorale); stems usually erect, rather woody 19 Leaves usually with a few black laminar lands; inner petal margin ± plane; stems usually ± ascending, herbaceous 20 19(18) Leaves all sessile, mostly elliptic or oblong-elliptic to narrowly oblong: flowers 15-18 mm in diam.; sepals 2.5-3 mm long, gradually to abruptly acuminate, intramarginal glands always present, spaced 8. tosaense Leaves on lateral branches petiolulate, elliptic to oblanceolate; flowers 9-10 mm in diam.; sepals 3-4 mm long, acute to subacuminate. intramarginal glands few or absent 9. iwatelittorale 20( 1 8) Stems single. 0.25-0.65 m, erect or basally ascending; inflorescence 7- 1 1 -flowered from 2-3 nodes, usually with 1 -4 flowering branches below 10. momoseanum Stems usually caespitose, 0. 1 -0.35 m, erect to procumbent; inflores- cence 3-7(-9)-flowered from 1-2 nodes, with 0-1 flowering branches below 1 1 . yezoense 1. Hypericum maculatum Crantz, Stirp. austr. 2: 64 (1763), 2nd ed. 2: 98 (1769); Allioni, Fl. pedem. 2: 45 (1785), 3: t. 83 f. 1 (1785) pro parte, excl. syn. H. androsaemifolium Vill.; Schinz & Keller, Fl. Schweiz, 3rded., 1: 358 (1909); A. Frohl. mSitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 538 (1911), in Oesterr. hot. Z. 63: 18(1913); Thellung in Allg. Bot. Z. Syst. 18: 19(1912); Hayek, Prodr. Fl. Pen. bale. 1: 534 (1925); Hegi, ///. Fl. Mitt.-Eur. 5(1): 517 pro parte, excl. f. 2004 (1925); N. Robson in Bot. Soc. Brit. Isles Proc. 2: 237 (1957), in op. cit. 3: 99 (1958), in Davis, Fl. Turkey 2: 400 (1967) in adnot., in Tutin et al.. Fl. Europ. 2: 268 (1968); Zeleny et al. in Futak & Bertova, Fl. Slovenska 3: 302, t. 37 f. 1 (1982); A. Ramos in Trab. Dept. Bot. Univ. Complut. Madrid 12: 49, t. 3 f. 1 (1983), in Acta Bot. Malac. 11: 167, f. 6e (1986), in Castroviejo et al., Fl. Iberica 3: 168 (1989); Hagemann in Flora 173: 112-115, tt. 12-14, 39 (1983); N. Robson & Strid in Strid, Mm Fl. Greece 1: 607 (1986); N. Robson in Cullen et al., Europ. Gdn Fl. 4: 60 (1995); in Wisskirchen & Haeupler, Standardliste Farn-u. Bliitenpfl. Deutschl.: 269 (1998). Type: Austria, 'alpen\ 1760 (fl), Breyn in Crantz 828 (BPI-lectotype, N. Robson, selected here). Fig. 6, Maps 1, 2. Perennial herb 0.2-0.6 m tall, erect or ascending from creeping and rooting base, with stems numerous to few, initially unbranched below inflorescence, later branched above fore. 2/3 of their length. Stems 4- lined, the subsidiary ones (decurrent between leaves) often less prominent or even partially absent2, usually with black glands on the lines; internodes 1 0-50 mm, shorter than to exceeding leaves. Leaves sessile; lamina 1 5^0 x 1 0-20 mm, broadly to narrowly elliptic, paler beneath, chartaceous; apex rounded, margin plane, base rounded; venation: 3(2) pairs of main laterals from lower quarter to base of midrib, tertiary reticulation dense; laminar glands absent or rarely pale, scattered, sometimes also black, few, punctiform; intramarginal glands black, ± close, irregular in size. Inflorescence (2-)3-c. 15- flowered, from 1-3 nodes, with flowering branches narrowly or curved ascending from up to 5 nodes, the whole cylindric to broadly pyramidal or subcorymbiform; pedicels 1.5-4 mm; bracts and bracteoles up to 5 mm long, narrowly triangular-ovate to narrowly N.K.B. ROBSON elliptic, entire. Flowers (15-)20-25(-35) mm in diam., stellate to reflexed; buds broadly ellipsoid, rounded to obtuse. Sepals 5, ± unequal, 3.5^.5(-5) x 2-2.7(-3) mm, broadly elliptic to broadly ovate or ± broadly oblong, rounded to obtuse or subacuminate (varying in same flower), entire to eroded-denticulate, erect to recurved in bud, recurved in fruit; veins 5(-7), not or slightly branched; laminar glands pale and sometimes also black, punctiform and sometimes striiform; intramarginal glands pale or occasionally black or often absent. Petals 5, golden yellow, not tinged red in bud, (8-) 10- 15 x 4-7.5 mm, c. 3 x sepals, broadly elliptic to obovate or more rarely symmetrically oblanceolate, distally crenate or entire; laminar glands pale and/or black, punctiform and sometimes proxi- mally striiform to linear (in subsp. immaculatum almost all linear); intramarginal glands few, distal or usually absent. Stamens 50-80(- 100), '3 '-fascicled, longest 7-10 mm, c. 0.65-0.9 x petals; anther gland black. Ovary 3-locular, 2-A x 1 .5-2.5 mm, ± broadly ovoid to ovoid-ellipsoid; styles 3, free, 3-4 mm, l-1.5(-2) x ovary, rather narrowly spreading; stigmas narrowly capitate. Capsule 6-10 x 4-6 mm, c. 2-2.5 x sepals, ovoid-ellipsoid to ± broadly ovoid; valves with longitudinal vittae, linear to striiform. Seeds dark brown, 0.8- 1 .2 mm, cylindric, not carinate or appendiculate; testa finely linear- foveolate. 2n=16, 32 (see under subspecies). In usually rather damp habitats - woods, scrub, grassland, moor- land, meadows and ditches; 0 (W. Scotland) - 2650 m (SE Switzerland). Europe from Scotland (Ross-shire), northern Scandinavia (to Hinnoya (Lofoten Is.), 68°22' N), northern European Russia and western Siberia, south to northern Spain, northern Italy, northern Greece and the central Ukraine, and east to south-western Siberia (R. Ob) with eastern outlier near Krasnoyarsk). Introduced into Canada (British Columbia). Hypericum maculatum is the most primitive species in the (mainly European) series Hypericum, its nearest relative in the (eastern) series Senanensia (Spp. 13-19) being 13. H. kamtschaticum var. pibairense from Hokkaido (N. Japan). It comprises two diploid subspecies (subspp. maculatum and immaculatum) and a tetraploid subspecies (subsp. obtusiusculum) that, from chromosome mor- phology and synthesis by colchicine doubling of subsp. maculatum (Robson, 1958a), would appear to be autotetraploid. The latter has sometimes been treated as a species (H. dubium Leers); but (i) the autotetraploidy and (ii) the occasional difficulty in assigning a plant to one or other taxon indicate that subspecies is the most appropriate rank, pace Martonfi et al. (1999) (see p. 75). The diploid subspecies of H. maculatum comprise two popul- ations overlapping in distribution. One (subsp. immaculatum) is confined to the south Balkans and lacks black glands on the petals, the pale ones being linear and the leaves usually pale-gland-dotted. Its chromosome number has not yet been counted, but morphologi- cally it is very similar to subsp. maculatum, less so to subsp. obtusiusculum. The other population (subsp. maculatum), which is widespread and slightly overlaps in area that of subsp. immaculatum in a zone from Bosnia to southern Bulgaria, has petals with puncti- form to shortly striiform black glands, the pale (punctiform) glands being few, and the leaves usually lack pale gland dots. 1 a. Hypericum maculatum subsp. immaculatum (Murb.) A. Frohl. mSitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 547, f. 8 (1911), in Oesterr. Bot. Z. 63: 18 (1913), in Mitt. Naturwiss. Vereines Steiermark 51: 223 (1915); Hayek, Prodr. : Partial or complete absence of subsidiary lines may possibly indicate hybridi/.ation with H. perforation (cf. H. x desetangsii, p. 75). STUDIES IN HYPERICUM 69 h Fig. 6 A. H. maculatum subsp. maculatum: (a) habit; (b) sepal; (c) petal; (d) capsule. B. H. maculatum subsp. immaculatum: (e) petal. C. H. maculatum subsp. obtusiusculum: (f) habit; (g) sepal; (h) petal; (i) capsule (a, f x 2/3; rest x 4). A. (a) Duvigneaud 77; (b, c) Meinertzhagen s.n.; (d) Lacaita 6402. 70 N.K.B. ROBSON Fl. Pen. bale. 1: 534 (1925); Hegi, ///. Ft. Min.-Eur. 5(1): 518 (1925);Stjep.-Vesel. in Josifovid, Fl. Srbije3: 116 (1972). Type as for H. quadrangulum var. immaculatum Murb. Fig. 6, Map 1 . H. quadrangulum var. immaculatum Murb. in Acta Univ. Lund. 27(5): 152 (1891); Beck in Ann. K. K. Naturhist. Hofmus. 10: 182 (1895). Type: Bosnia, Hercegovina, Velezplanina, c. 1500 m, 27 July 1889, Murbeck s.n. (LDMectotype, selected here). H. quadrangulum subsp. immaculatum (Murb.) Wettst., Beitr. Fl. Alban. : 36 ( 1 892). Type as for//, quadrangulum var. immaculatum Murb. H. immaculatum (Murb.) Vierh. in Mitt. Naturwiss. Vereins Univ. Wien II, 4: 62 (1906), Aufz. Simony Pfl.: 27 (1906). H. maculatum subsp. immaculatum var. punctatum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math. Naturwiss. Kl. 120(1): 549 (1911). Type: not seen, no specimen cited. H. maculatum subsp. immaculatum var. epunctatum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math. -Naturwiss. Kl. 120(1): 549 (1911). Type: not seen, no specimen cited. H. maculatum var. immaculatum (Murb.) Gu§ul. & Nyar. in Savul., Fl. R. P. Roman. 4: 33 (1956); Stoj., Stef. & Kitanov, Fl. Bulg. 2: 721 (1967); Jordanov & Koz, in Jordanov, Fl. R. P. Bulg. 4: 260 (1970); Robson & Strid in Strid, Mtn Fl. Greece 1: 608 (1986); N. Robson in Cullen et al., Europ. Gdn Fl. 4: 60 ( 1 995), in Wisskirchen & Haeupler, Standardliste Farn-u. Bliitenpfl. Deutschl.: 269 (1998). Type as for H. quadrangulum var. immaculatum Murb. H.fallax subsp. immaculatum (Murb.) O. Schwarz in Drudea 5: 63 (1965). Stems always with complete subsidiary lines. Leaves with densely reticulate tertiary venation; laminar gland dots pale, usually ± dense, occasionally with a few black. Inflorescence branches narrowly ascending, making an angle of c. 30° with stem. Flowers 15-35 mm in diam. Sepals broadly ovate, rounded to obtuse or somewhat acute, entire or minutely denticulate; laminar glands pale, striiform and punctiform. Petals entire, without or rarely with 1-3 black marginal glands; laminar glands pale, linear to striiform. Styles equalling ovary. 2n=16? (see p. 63) Subalpine meadows; (SOO-)lOOO-c. 2200 m. Southern Romania (Carpathians), western and southern Bulgaria, northern Greece, Makhedonia, southern Serbia, Bosnia, Montenegro, Kosovo. ROMANIA. Gus.uleac & Nyarady (1956: 33) record 'var'. immaculatum from SE Transylvania (Odorhei, Codlea and Sibiu regions). BULGARIA. Mikhaylovgrad: Predbalkan, Ostabhang des Midshur [Midzhor], 4 September 1958 (fl), K. Meyer 1 140 (JE). Sofia: Cherni Vrukh [Mt Vito§a], c. 1 440 m, 23 July 1952 (fl), Vyhodcevski s.n. (H). Blagoevgrad: Rila Planina, in valle Ristritza, c. 1000-2000 m, 20 August 1907 (fl), Schneider & Bergmann 878 (BM, K); Pirin, above Bansko, above L. Okoto, 2130 m, 27 July 1993 (fl), Jury 12071 (BM, RNG*); Rhodopen, Snesanka, 1900 m, 8 October 1958 (fr), K. Meyer 1985 (JE). GREECE. Makedhonia: Drama, in monte Tsalakou ad septentriones ditionis 'Elatia', 1800 m, 18 August 1978 (fl), Greuter 16532 (BM, C); Serron, Mt. Vrondous, NE of summit, 1600m, 27 July 1981 (fl), Farsakoglou, Franzen & Strid 19538 (C). MAKHEDONIA. Sar Mtns, Mt. Scardos, 2000 m, 29 May 1983 (o. fr), McClintock s.n. (BM). SERBIA. Nis: ad [R.]Vlasina, July 1895 (fl), Adamovic s.n. (K).Kosovo: Bertiscus, in valle rivuli Docanska Bistrica, 1200-1300 m, 17-19 July 1933 (fl), Rechingerf. & Scheffer 1338 (BM). BOSNIA. Sarajevo, Trebevic, 1600 m, 30 June 1931 (fl), Gilliat-Smith 2922 (K); Velez Planina, 5 July 1912, Sagorski s.n. (JE). MONTENEGRO. Vasovavici [Vasojevicki], sub Varda, 19 July 1898 (fl), Baldacci It. Alban. VI 181 (BM, K); prope monast. Piva, July 1905, Rohlena s.n. (JE). Map 1 1 . H. maculatum. a. subsp. immaculatum A ; b. subsp. maculatum differentiate subspecies. ; other records O. Widespread in Germany , but the records do not STUDIES IN HYPER1CUM Of the two diploid subspecies of//, maculatum, subsp. immaculatum would seem to be the more primitive and to constitute a relict Balkan population that has given rise to both subsp. maculatum and (by autotetraploidy) to subsp. obtusimculum. It is also the nearest taxon morphologically to the H. tetrapterum group (Spp. 2-4) and would appear to be one parent of the allotetraploid 5. H. perforatum. The linear pale petal glands and usually glandular-punctate leaves both lead to the above conclusions, as do the punctate leaves and linear to striiform petal glands in H. perforatum (q.v.). A sheet from Greece (Makhedonia, Drama, Mt. Falakron, Stamatiadou 10131 (C)), with one plant with characters of subsp. maculatum among four plants of subsp. immaculatum, may indicate an independent development of punctate black laminar glands in the sepals and petals. Ib. Hypericum maculatum subsp. maculatum Fig. 6, Map 1. H.fallax Grimm in Nova Acta Phys.-Med. Acad. Caes. Nat. Cur. 3, App.: 362 (1767); O. Schwarz in Drudea 5: 62 (1965). Types: Germany, Sachsen, Eisenach, vor Winterstein, Grimm s.n. (JE?- syntype), auf der Schlotwiese, Grimm s.n. (JE?-syntype), unten dem Inselberg, Grimm s.n. (JE?-syntype), um Mosbach, Grimm s.n. (JE?-syntype). Syntypes not found in JE. H. delphineme Vill., Prosp. Hist, pi Dauphine: 44 (1779), Fl. delph.: 81 (1785), Hist. pi. Dauphine 1: 281, 294, t. 44 (1786), 3: 497 (1789). Type: France, Isere, Oysans [Oisans], La Moncherolle, 1789 (fl), Villars MNHGr 1837. 28503 (GRM-photograph!- lectotype, selected here). H. tetragonum Fr., Fl. hall: 124 (1818). Type: Sweden, Halland, 'copiose in pratis, aliud in Suecia non reperitur', Fries s.n. (UPS- holotype). Fries rejected the name H. dubium for his plant. H. dubium sensu Mauri, Roman, pi. cent, xiii: 27 (1820). H. quadrangulum var. [y] sensu Choisy, Prodr. monogr. Hyperic. : 47 (1821). H. quadrangulum sensu Fr., Novit.fl. suec.: 94 (1823); Spach, Hist, nat. veg., Phan. 5: 386(1836), in Ann. Sci. Nat. Bot. 5: 357 (1836); Lamotte in Bull. Soc. Bot. Fr. 21: 122 (1874); Lange in Willk. & Lange, Prodr. fl. hispan. 3: 591 (1878); J.D. Hooker, Student fl. Brit. Isles, 3rd ed.: 73 ( 1 884); Schinz in Bull. Herb. Boissier II, 3: 15 (1902), in Vierteljahrsschr. Naturf. Ges. Zurich 49: 231-241 ( 1 905); Tourlet in Bull. Soc. Bot. France 50: 307 ( 1 903); R. Keller in Engl. & Prantl, Nat. Pflanzenfam., 2nd ed. 21: 179 (1925); Druce, List Brit. pi. ,2nded.: 64(1928); Stefanoff inGod. Agr. les. Fak. Univ. Sofiya 10: t. 2 f. 18, t. 4 f. 30 (1932), 11: 175, 177 (1933), 12: 88 (1934), in Pflanzenareale 4: Karten 6a, 8 (1933); Gorschkova in Shishkin & Bobrov, Fl. U.R.S.S. 15: 242, t. 12 f. 3 (1949), etauct.; non L. H. quadrangulum var. [y] maculatum (Crantz) Choisy in DC., Prodr. 1 : 548 ( 1 824). Type as for H. maculatum Crantz. Choisy cited '//. maculatum Allioni', but Allioni (1785) cited Crantz. H. commutatum Nolle, Novit. fl. holstat.: 69 (1826); Rchb., Fl. germ, exc., II: 837, no. 5180 (1830-1832), Iconfl. germ. helv. 6: t. 345 (1844); Bonnet in Bull. Soc. Bot. France 25: 276 (1878), qua hybr.; Rouy in Rouy & Fouc., Fl. France 3: 335 (1896) qua hybr. Type: Germany, Schleswig-Holstein, in Due Lauenburg prope pagum Biichen ad fl. Stechnitz, Nolle s.n. (Bt-holotype). Frb'hlich (1911: 547) saw Nolte's type and identified it as H. maculatum subsp. typicum var. punctatum (Schinz) A. Frohl. H. quadrangulum var. [b] subcorymbosum Schur, Enum. pi. Transsilv.: 132 (1866). Type as for H. commutatum Nolte. H. quadrangulum var. [c] macrophyllum Schur, Enum. pi. Transsilv. : 132 (1866). Type: Romania, Transylvania, Kronstadt, July-Au- gust, Schur s.n. (BRNU? or LW?). 71 H. umbellatum Miel. ex Wohlf. in W.D.J. Koch, Syn. deut. schweiz. FL, 3rd ed. 1: 429 ( 1 892); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. I, 120(1): 557 (1911), pro syn.; nomen. H. quadrangulum var. genuinum Schinz in Bull. Herb. Boissier II, 3: 21 (1902), in Vierteljahrsschr. Naturf. Ges. Zurich 49: 240, 241 (1905) sub subsp. quadrangulum, pro parte excl. typum. Type as for H. quadrangulum auct. non L., i.e. H. maculatum Crantz. H. quadrangulum var. punctatum Schinz in Bull. Herb. Boissier II, 3: 20, 22 (1902), in Vierteljahrsschr. Naturf. Ges. Zurich 49: 240, 241 (1905) sub subsp. quadrangulum. Type: Switzerland, Graubunden, Oberengadin, Fornogletscher, Hegi s.n. (BR?- holotype). H. quadrangulum subsp. quadrangulum sensu Tourlet in Bull. Soc. Bot. France 50: 307 (1903); Schinz in Viertel-jahrsschr. Naturf. Ges. Zurich, 49: 231-241 (1905), pro parte excl. typum. H. maculatum subsp. eu-maculatum Schinz & Thell. in Schinz & Keller, Fl. Suisse: 381 (1909); Thellung in Allg. Bot. Z. Syst. 1912: 24 (1912); A. Frohl. in Oesterr. Bot. Z. 63: 18 (1913), in Mitt. Naturwiss. VereinesSteiermarkSl: 223 (1915); Hegi, ///. Fl. Mitt.-Eur. 5(1): 517 (1925). Type as for H. maculatum Crantz. H. maculatum subsp. typicum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 1, 120(1): 540 (191 1). Type as for H. maculatum Crantz. H. maculatum subsp. typicum var. genuinum (Schinz) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 1, 120( 1 ): 545 (1911). Type as for H. quadrangulum var. genuinum Schinz. H. maculatum subsp. typicum var. punctatum (Schinz) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 1, 120(1): 545 (1911). Type as for H. quadrangulum var. punctatum Schinz. Frohlich (p. 546) names forms glabrum and subnervosum (based on nervation) as well as rotundifolium and angustifolium (based on leaf shape) for both vars genuinum and punctatum; but giving the same epithet to infraspecific taxa within the same species is inadmissable (I.C.B.N. Art. 24, note 2). The epithet f. luteum, however, was given to subsp. typicum as a whole and would therefore appear to be valid. H. maculatum subsp. typicum forma luteum A. Frohl. in Sitzungsber. Kaiserl Akad. Wiss., Math.-Naturwiss. Kl. 1, 120(1): 546 (191 1). Type: Austria, Styria, Puntigamer Guns (bei Graz), 16 July 1910 (fl), Frohlich GZU 16228 (GZU!-lectotype, selected here). H. maculatum subsp. quadrangulum sensu Hayek, Prodr. Fl. Pen. bale. 1: 534 (1925). Hayek cited '//. quadrangulum Tourlet', i.e. H. maculatum Crantz not H. quadrangulum L. H. quadrangulum subsp. maculatum (Crantz) Goday & Carbonell in Anales Inst. hot. Cavanilles 19: 380 (1961). H.fallax Grimm subsp. fallax, O. Schwarz in Drudea 5: 63 (1965). H.fallax subsp. fallax Grimm var. fallax, O. Schwarz in Drudea 5: 63(1965). H. fallax subsp. fallax var. fallax forma punctatum O. Schwarz in Drudea 5: 63 (1965). Type not cited (JE). H. fallax subsp. fallax var. quadrangulare sensu O. Schwarz in Drudea 5: 63 (1965) pro parte, quoad descr. Schwarz cited '//. quadrangulare L. (1776)' as basionym. H. maculatum subsp. maculatum forma punctatum (Schinz) Stjep.- Vesel. in Josifovic, Fl. Srbije 3: 1 16 (1972). Icones: Rchb., Ic. fl. germ. helv. 6: t. 343 f. 5178 (1842^14); Gorschkova in Shishkin & Bobrov, Fl. U.R.S.S. 15:188, t. 12 f. 3 (1949); Zeleny et al. in Futak & Bertova, Fl. Slovenska 3: t. 37 f. 1 (1982). Stems always with complete subsidiary lines. Leaves with densely reticulate tertiary venation; pale glands usually absent but some- 72 N.K.B. ROBSON times few to numerous. Inflorescence branches narrowly ascending, making an angle of c. 30° with stem. Flowers 15-25 mm in diam. Sepals broadly ovate to broadly elliptic or elliptic-oblong, entire or more rarely slightly eroded-denticulate; laminar glands pale and black, punctiform. Petals entire, without or rarely with 1-3 black marginal glands; laminar glands all or mostly black, punctiform or proximally shortly striiform. Styles equalling ovary. 2n=16 (Noack, 1939; Robson, 1956, 1957; Schwarz, 1965; Reynaud, 1975; Dmetrieva, 1986; Parvenov & Dmetrieva, 1987), n=8 (Nielsen, 1924; Winge, 1925; Robson, 1956, 1957, 1981; Laane, 1969). Subalpine and alpine regions of the Pyrenees, Alps and Balkans; hilly regions further north (Massif Central, Vosges, Ardennes, Scot- land); 0-2650 m (lowland in the north). Distribution of the species except for lowland NW Europe (but with enclaves in the Ardennes and western and central Scotland) and valleys of the Alps east to Styria. SCOTLAND. W. Ross: Ullapool, by Ullapool R., 16 August 1996 (fl & e.fr), Kitchener s.n. (BM). Westerness: Sunart, Strontian, 13 July 1966 (bud), Cannon & Kendrick s.n. (BM). Argyll: Dunadd area, between Lochgilphead and Kilmartin, July 1 958 (fl), Kenneth s.n. (K). Arran: Lamlash, 10 August 1937, Mackechnie s.n. (BM). Lanark: Glenhove near Airdrie, 22 August 1951 (fl), Mackechnie s.n. (BM). Mid Perth: between Lawers and Feaman, 3 September 1913 (fr), Marshall in B.E.C. 3787 (BM, K). S. Aberdeen: Aboyne, 9 August 1968 (fl), Robson s.n. (BM). ENGLAND (introduced). Herts.: Gaddesden, The Hoe, 1 1 August 1962 (fl), J. & C. Dony & Allen s.n. (BM). THE NETHERLANDS. Very rare. Records from Gelderland (1954), south of Limburg (1917), and near Nijmegen ( 1924), fide Mennema, Quene- Broterenbrood & Plate (1980). BELGIUM. Liege: Recht, 10 August 1977 (fl), Duvigneaud 77 B 737 (H). Vim ur: Cul-des-Sarts, Le Marais, 5 August 1977 (fl & fr), Duvigneaud 77 B 704 (BM, H). Luxembourg: Vielsalm, bord de la Salm, 28 August 1935 (fr), Mosseray s.n. (BM, BR*). FRANCE. Ardennes: pres de Hulle, rive gauche de la Hulle, 14 August 1970 (fl), Duvigneaud 70 F 41 (H). Vosges: Ventron, Gerbamont, August 1 880 (Lfl), Pierrot (BM,K).Doubs: Mont d'Or, 6 August 1 85 \ (fl),Grenier s.n. (BM). Jura: Noirmont, pres de Rousses, 6 August 1856 (fl), Michalet fam. 2, 67 (K). Haute-Savoie: Vallee de 1'Arve, Vallon de Flaine/Cluses, 1700m, 16 July 1964 (fl),F. &J.-D.Bersier5 17 (G*, H). Savoie: Lanslebourg, 9 July 1884 (fl), Willmott (K). Isere: L'Oursiere, pres de Uriage, 1500-1600 m, 26 August 1880 (fl), TMet (BM). Hautes-AIpes: Lauteret, 18 July 1931 (fl), Pugsley s.n. (BM). Alpes-de-Haute-Provence: Goudeissant, pres Barcelonnette, 11 August 1854 (fl), Willmott s.n. (K). Loire: Mont Pilat, 1400m, 14 July 1881 (fr), Glastien s.n. (BM). Puy-de-D6me: leMont-Dore, plateau de Durbise, 20 August 1928 (fl), Fiton in Duffour 5662 (BM). Cantal: Puy Mary, NW of Murat, 12 September 1966 (fl), Summerhayes 4202 (K). Haute-Garonne: Luchon, 14 July 1889 (fl), Murray s.n. (BM). Ariege: Ax, chemin d'Orgeix, n.d. (fl), de Martrin s.n. (K). Pyrenees- Orientales: Mont-Louis, Ostrand der Wald von Font-Romeu, 7 August 1 926 (fl), Ronniger s.n. (W). Hautes-Pyrenees: Gavarnie, 1350-1500 m, 30 August 1932 (fl), Meinertzhagen s.n. (BM). ANDORRA. Ordino, 1250 m, 14 July 1962 (fl), Edwards & Perry 9 (BM). SPAIN. Gerona: [Santuario de] Nuria, Via Crucis, 3 August 1934 (fl), Dauder Rodes s.n. (BM). Lerida: Gorges de Llo, 1450 m, 29 August 1929 (fr), Sennen s.n. (BM). Teruel: Bronchales, Sierra Alta, 11 June 1962 (st), Kjellqvist & Love N588 (RNG). ITALY. Val d' Aosta: Val Ferret, SE of Mont Blanc, 1 2 July 1 854 (fl), Hort s.n. (BM). Novara: Val Formazza, Discesa da C. Cavala e S. Michele, 1 827- 1 250 m, 3 August 1 9 1 8 (fl), Boggiani s.n. (BM). Verona: Monte Baldo, 1 000 m, June 1902 (fl), Rigo s.n. (H). Bolzano: Pusteria med., 1860-1950 m, 29 August 1871 (e. fr), Huter s.n. (BM). Treviso: Bosco Cansiglio (?), 1000 m, 10 July 1922 (fl), Fiori Fl. Ital. Exs. Ill 2664 (BM, K). SWITZERLAND. Vaud: Alpen iiber Bex, 1832 (fl), Thomas in Reichenbach 1500 (BM, H, JE, WAG). Valais: Col du Lein, 1720 m, 29 July 1979 (fl), Lawalree 21846 (BM, BR*). Bern: Miirren, 1590 m, 2 September 1923 (fr), Lester Garland s.n. (K). Graubiinden: Tachitta Valley, nearPreda, Albula, 20 July 1928 (fl), Pugsley s.n. (BM). Glarus: Glarnisch Mtn, Gliiterbach, 1 1 50 m, 1 9 September 1 963 (fr), Robson 1 826 (BM). Appenzell: am Ga'bris ob Gais, July-August (fl), Schneider s.n. (BM). AUSTRIA. Vorarlberg: Ratikon, 19 July 1952 (fl), Jacobs 3577 (L). Tirol: prope Trins in valle Gschnitz, c. 1 100 m, n.d. (fl), Kerner in Fl. Exs. Austr.-Hung. 3647 (BM, H, JE, K). Salzburg: Radstadter Tauernpasshohe, 1700-1300 m, 13 August 1932 (fl), Hubl s.n. (BM). Karnten: Weissenfels Krain, September 1913 (st), Meebolds.n. (K). Steiermark: Schladming, 750 m, 3 July 1994 (fl), Townsend 94/34 (K). Niederosterreich: Monies Rax in valle Hollenthal, 600 m, August 1953 (fl), Patzak s.n. (H, W*). GERMANY. Bayern: Kampenwald, above Steinlinghutte, 26 July 1934 (fl), Willmott s.n. (BM); Oberostdorf, 25 July 1927 (fl), Mann s.n. (FR). Hessen: Wasserkuppe gegen den Pferdskopf, c. 900 m, 9 August 1903 (fl), Goldschmidt s.n. (FR). Thiiringen: Erfurt, bei Oberhof, Schmiicke, c. 900 m, 1 6 July 1952 (fl), Launert s.n. (BM); Suhl (?), Neuhaus (Reunsteig), 1 950, K. Meyer s.n. (JE). Sachsen: bei Dresden, n.d. (fl), Horn in Reichenbach, Exs. Fl. Germ. 1397 (BM, H, JE). Sachsen-Anhalt: Harz, Minfra nach Harlingerode, 1 858, (Collector!) (JE); Magdeburg, Werningerode, July 1 860 (fl), (Collector!) (FR). Brandenburg: fide Ascherson (1864: 113). Berlin: Spandau, Finkelhang, 28 July 1 867 (1. fl) Wagner s.n. (K). Mecklenburg- West Pomerania: Schwerin, prope Gustrowiam, 1830, Jahn! s.n. (JE); Riigen I., 1958, Inst. Excurs. s.n. (JE). DENMARK. Jylland: Frederikshavn, Pikkerbakken, 26 July 1968 (fl), Jensen, Nielsen & Pedersen in Fl. Jtl. Exs. 465 (BM, H, TAI). Fyn: Dyreborg Skov, Faaborg, 24 July 1952 (fl), Young 4482 (BM). Sjaelland: Dyrhavn, Klampenborg, 15 August 1947 (fl), Price 307(1) (K). FAEROE ISLANDS. Streymoy, Vagar (fide Johansen, 2000). NORWAY. Akershus: As, near Ski, c. 48km S. of Oslo, 29 July 1961 (fl), Goodfellow 61.1600 (BM, LIV*). Vestfold: Asgardstrand, 31 August 1934 (fr), Williams s.n. (BM). Vest Agder: Vigeland, Kristiansand, August 1909 (fl), Gamble 29773 (K). Rogaland: Stangaland, Eide, 14 August 1930 (fl), Rosseland s.n. (H). Hordaland: Hardanger Fjord, Utne, S. Bergenhus, 16 August 1934 (fr), Williams s.n. (BM). Nordland: Holandsfjord, c. 200 m, 22 July 1950 (fl), Boyd in Durham U. Exped. 346 (BM); Lofoten Is., Svolvaer, Svolvaergjeita, 27 July 1939 (fl), Blakelock 73 (K). SWEDEN. Blekinge: Rodeby, Rodebyholm, 11 July 1989 (fl), Hull s.n. (H). Gotland: Oland I., Bredsatra sn., Bredsatra by, 15 August 1957 (fl), Saarsoo 7684 (H). Kronsberg: Soraby s. n. 6. om Stavsakra, 4 August 1957 (fl), Saarsoo 7684 (H). Jonkoping: Korsberga, Molebo, 3 August 1908 (fl), Stalin s.n. (H). Skaraborg: Skara, 15 July 1908 (fl), Jacob & Jacobson s.n. (BM). Alvsborg: Skene, c. 700 m, nordlich Berghem station, 13 August 1 965, Buttler & Gaubl 7941 (FR). Orebro: Narke, Svennevad, Gropen, c. 75 m, 4 August 1950 (fl), Kjellmart s.n. (H). Stockholm: Stockholm, 23 July 1858 (fl), Nyman s.n. (BM). Uppsala: prope Upsaliem, July 1869 (fl), Ahlberg s.n. (BM). Vastmanland: Arboga, Hamnen, 19 July 1951 (fl), Fridell s.n. (H). Kopparberg: Leksands sn., Bjorkberg, 17 September 1956 (fl & fr), Norrman s.n. (BM). Gavleborg: Gavle, Lovudden, 20 August 1964 (fl & fr), Nannfeldt s.n. (BM). Vasternorrland: Alno, 15 September 1909 (fl 6 fr), Ronnblad s.n. (H). FINLAND. Oulu: Kn. Kajaania, Likapuro, n.d., Heikkinen s.n. (JE). Keski-Suomi: Laukaa, Leppavesi Linnasaari, 30 July 1965 (fl), Valovirta s.n. (H). Kuopio: Savonia borealis, lisalmi, lima'ki, 6 August 1975 (fl), Tallgren s.n. (BM). Pohjois-Karjala?: Karelia austr., par. Vehkalahti, Pyhalto, 7 July 1961 (fl), Fagerstrom s.n. (BM). Turku Pori: Turku, Ruissalo, Laurila, 18 July 1968 (fl), Tallgren s.n. (BM). Aland: Hammarland, Nafsby, 20 July 1968 (fl), Haakana s.n. (BM). RUSSIA. Ladoga-Ilmen: Leningradsky obi., Lupskiy r-n., Merabush- Lupesfka, 13 July 1969 (fl), Nepli 18 (K). Dvina-Pechora: Archangelsk, Beresink, 1919 (fl), Grantham s.n. (BM). Upper Volga: Prov. Mosqua, prope urbe Noginsk, 30 June 1968 (fl), Samozvon 158 A (H). Volga-Kama: Prov. Kostroma, distr. Kineschma, prope Pawlowskoje, 4 July 1903 (fl), Klementz \ 970 (H). Upper Dnieper: Smolensk obi., Dukhovshchina distr., prope pag. Ovsianka (10 km NW of Preczistoe), 8 August 1974 (fl & fr), Makarov & Bazhenova s.n. (H). Middle Dnieper: fide Gorschkova (1949: 243). Volga- Don: Orel obi., Orlovsk, Ostrovakami, n.d. (1. fl), Tarachkova & Poganki 3 1 7 (H). Transvolga:/kfc Gorschkova (1949: 243). Upper Tobol: Chelyabinsk, Ural, in vallibus montis Jurma, July 1844 (fl), [Buhsel] (Herb. Buhse). Ob: south, fide Gorschkova (1949: 243). Angara-Sayan: Ibryul' village, fide Gorschkova (1949: 243). STUDIES IN HYPERICUM 73 ESTONIA. Ingria, Hietamaki, Nutty, 20 July 1915 (fl), Jamalainen s.n. (H); Dago I. [Hiiumaa], Emmaste, 1872, Winkler 335 (JE). LATVIA. Krashavskiyr-n.,Zabludovka, 13 July 1967(fl),Ma/tarovas.n. (H). LITHUANIA. Throughout the Territory (Vilkeviciute, 1971). BELORUS. NiaDkow, distr. Nowogrodek [Novogrudok], 28 - 1898 (fl), Dybowski in Woloszczak, Fl. Polon. Exs. 715 (BM). UKRAINE. *Rossia Subcarpatica: monies Vysoke Poloniny, polonina Rovna, c. 1300 m, 8 July 1938, Jirdsek s.n. (PR). *Carpatorossia: Corna Hora, in ... montis Hoverla, 1 800-2056 m, July 1935, M. Deyl in PR 164845 (PR). Bucovina: in summis alpinis ... alpis Tschumaled, July 1833 (fl), Herbich s.n. (W). POLAND. Jelenia Gora: Sudet, Hirschberg, 1 July 1958, Kohler & Zippold s.n. (JE). Walbrzych: Sla^sk Dolny, Spalona pow. Bystrzyca Kl, 25 July 1958 (fl), Gotowin 157 (H). Wroclaw?: Breslau, Strachate, 28 July 1888, Kionka s.n. (JE). Bielsko Biala: distr. Wadowice, infra cas. Potrqjna, 24 July 1938 (fl), tancucka in PI. Pol. Exs. 329a (BM, K). Tatry: Tatra, Zakopane, Dol. Maty Zaly, 11 July 1958, Kohler & Zippold s.n. (JE). Krosno?: E. Carpath., Breskul, ± 1450 m, 29 August 1938 (fl), Kosij in PI. Pol. Exs. 329b (BM, K). CZECH REPUBLIC. Bohemia: Monies Sumava, p. p. Hamny, 3 August 1962 (fl), Hosticka s.n. (H); *distr. Cheb, in valle R. Libocky polock sub vico Opalov, ad Dolni mlyn, c. 560 m, 18 Seplember 1950, Kldstersky in PR 165161 (PR). Moravia: Weisskirchen bei Hrubuska, July 1911 (fl), Petrak 1 142 (BM); *chalel Bunc near Salas, 6 July 1987, Mdrtonfi 459 (KO). SLOVAKIA. Michalovce, Vihorlal, oberhalb Remelske Hamre, c. 500 m, 23 July 1974 (fl), F.K. & J. Meyer 1 1006 (JE); *Bukovske vrchy Mis, near Runina, 3 1 July 1 995, Mdrtonfi 2022 (KO); *pasture 'Voniarky' near Dobsina, c. 915 m, 13 Augusl 1991, Mdrtonfi 976 (KO). HUNGARY. Absent? SLOVENIA. Bled [Veldes], 1924 (fl), Leathes s.n. (BM). CROATIA. Widespread (Schlosser & Vukolinovi, 1869: 382). SERBIA. North: Fruska Gora. Central: Suva planina, Stara planina, Tara planina, Zlatibor, Kopaonik (Stjepanovi-Veselichi, 1972). ROMANIA. Transsilvania: Cri§ana, Bihor, ad Balneas Slana de Vale, c. 1 100 m, July 1936 (fl), Borza in PI. Exs. Rom. 1520c (H, K); in monle Bulka- Kandrenulni ad Dorna-Kandreni [Dornis,oara], July 1838 (fl), Herbich s.n. (W); *Munlii Bihorlui Mis., above Scarijoara, c. 1400 m, 16 July 1995, Mdrtonfi 1985 (KO). Moldavia: Comm. Darmane§li, monle Nemira, c. 1500 m, 14 July 1970 (fl), Baraba$ & $ova 158 (H); *Distr. Turda-Aries,, Valea Morii prope oppid. Cluj, c. 600-650 m, 28 July 1923, E.I. Nydrddy in CL 444000 (CL). Walachia: Oltenia, dislr. Gilorl, monl. Paring, ad MusStoiu, c. 1540 m, 30 Seplember 1960 (e. fr), Buia el al. in Fl. Ollen. Exs. 52 (BM, H). BULGARIA. Jordanov & Kozhukharov (1970: 260) record var. maculatum from N. and W. Slara planina, the Znepolski region (Golo birdo), W. Granichni planina, the Vilosa region (Vilosa), Slavianka and Rila. 1 c. Hypericum maculatum subsp. obtusiusculum (Tourlet) Hayek, Sched.fl. stiriac. 23-24: 27 (1912); Thellung in Allg. Bot. Z. Syst. 1912: 24 (1912); A Frohl. in Oesterr. Bot. Z. 63: 13, 18 (1913), in Mitt. Naturwiss. Vereines Steiermark 51: 223, 224 (1915); Schinz & Keller, Fl. Schweiz, 4th ed. 1: 451 (1923); Hegi, ///. Fl. Mitt.-Eur. 5(1): 517 (1925); N. Robson in Bot. Soc. Brit. Isles Proc. 2: 237 (1957), in op. cit. 3: 99 (1958), in Tutin et al., Fl. Europ. 2: 268 (1968), in Wisskirchen & Haeupler, Standardliste Farn- u. Bliitenpfl. Deutschl: 269 (1998). Type as for H. quadrangulum subsp. obtusiusculum Tourlet=//. desetangsii var. imperforatum Bonnet. Fig. 6, Map 2. H. dubium Leers, Fl. herborn.: 165 (1775); Sm., Engl. Bot.: 296 ( 1 796), Fl. brit. 2: 802 ( 1 800); Bab. in Trans. Bot. Soc. Edinburgh 1: 88 (1841), Man. Brit, hot.: 57 (1843); Bellynk, Fl. Namur. 52 (1855); Syme, Engl. Bot., 3rd ed. 2: 151 (1864); Schinz in Vierteljahrsschr. Naturf. Ges. Ziirich49: 23 1-241 (1905); Martonfi et al. in Preslia 71: 337 (1999). Types: Germany, Hesse, 'ad sepes am Sieghaus', Leers s.n. (?-syntype); 'am Homberg', Leers s.n. (?-syntype). According to Stafleu & Cowan (1979), Leers' herb- arium was taken to Russia (early in the nineteenth century?) and subsequently partly destroyed and partly lost. His description (apart from the ovate entire sepals) and the location of Herborn are both consistent with his plant's belonging to subsp. obtusiusculum; and broad entire sepals are not unknown in subsp. obtusiusculum, having been found in Kirkcudbrightshire, SW Scotland (see record, p. 75), and they were recorded by Tourlet (1903). They also occurred in a plant of subsp. maculatum in which the chromosomes had been doubled by colchicine treat- ment (Robson, 1956, 1957, 1981). Schinz, in the paper cited above, discussed the name H. dubium but treated it as a synonym of H. quadrangulum in his sense, i.e. H. maculatum Crantz. Frb'hlich (1911) also regarded H. dubium as a synonym of H. maculatum (i.e. subsp. typicum). H. obtusum Moench, Methodus: 129 (1794), nom. illegit. Type as for H. dubium Leers. H. leersii C.C. Gmel., Fl. bad. 3: 253 (1808); 5: 575 (1826), nom. illegit. Type as for H. dubium Leers. H. quadrangulum var. [p] sensu Choisy, Prodr. monogr. Hyperic.: 47(1821). H. quadrangulum var. [[3] dubium (Leers) Choisy in DC., Prodr. 1: 548 (1824), pro parte excl. syn. H. delphinense Vill. H. dubium var. [p1] maculatum sensu Bab., Man. Brit, hot., 3rd ed.: 58 (1851), i.e. sensu H. dubium Bab. in Trans. Bot. Soc. Edin- burgh 1: 88 (1841), Affl/i. Brit, hot.: 57 (1843), 2nded.: 60(1847). H. dubium var. [a] genuinum Syme, Engl. Bot., 3rd ed. 2: 151 (1864). Holosepalum dubium (Leers) Fourr. in Ann. Soc. Linn. Lyon, N.S. 16:352(1868). IHypericum quadrangulum var. [b] ambiguum Schur in Verh. Naturf. Vereins Briinn 15(2): 158(1 877). Type: Czech Republic, Moravia, oberhalb der Mazocha bei Blansko, August 1871, Schur s.n. (BRLU? or LW?). H. desetangsii var. [|3] imperforatum Bonnet in Bull. Soc. Bot. France 25: 277 (1879); Schinz in Bull. Herb. Boissier II, 3: 14 (1902), in Vierteljahrsschr. Naturf. Ges. Zurich 49: 241 (1905). Type: France, Seine-et-Oise, Meudon, 11 August 1878 (1. fr), Bonnet in Soc. Dauphinoise 1998 (P-lectotype, selected here; CLF!, FR!, K!-isolectotypes). My preliminary selection of lectotype for this variety, Lloyd's specimen from Loire-Inferieure [Loire- Atlantique], was published by Martonfi et al. (1999: 339) as 'Robson, in prep.'. Neither they nor I have seen this specimen, and the more widely distributed Bonnet exsiccata cited above would seem to be a more appropriate choice. H. quadrangulum var. occidentale Franch., Fl. Loir-et-Cher. 97 (1885); Rouy in Rouy & Fouc., Fl. France 3: 335 (1896). Types: France, Loir-et-Cher, environs de Mondoubleau, 16 July 1867 (fl), Legue s.n. (P! -lectotype, selected here); bois de Roquelane, August 1864 (fl), Legue s.n. (P!-syntype); Onzain, petit bois de Mare, Monin s.n (?-syntype, not lectotype as cited in Martonfi et al., 1999: 339). The last specimen has not been found in P. H. acutum subsp. desetangsii var. imperforatum (Bonnet) Rouy in Rouy & Fouc., Fl. France 3: 337 ( 1 896), comb, illegit. Type as for H. desetangsii var. imperforatum Bonnet. H. quadrangulum subsp. obtusiusculum Tourlet in Bull. Soc. Bot. France 50: 307 (1903). Type as for H. desetangsii var. imperfor- atum Bonnet. Tourlet describes H. quadrangulum subsp. quad- rangulum on the same page in the sense of H. tetrapterum Fr. H. quadrangulum subsp. obtusiusculum var. [a] imperforatum (Bonnet) Tourlet in Bull. Soc. hot. France 50: 308 ( 1 903). Type as for H. desetangsii var. imperforatum Bonnet, i.e. this is subsp. obtusiusculum Tourlet var. obtusiusculum. H. quadrangulum subsp. obtusiusculum var. [pV| perforatum Tourlet 74 N.K.B. ROBSON Map 2 Ic. H. maculatum subsp. obtusiusculum •. other records O. French and German records probably incomplete, as are the Irish ones. in Bull. Soc. Bot. France 50: 308 (1903). Type: France, Indre-et- Loire, vallon de la Desmee, [Les] Hermites, Doucet s.n. (G?- holotype). H. maculatum var. [|3] babingtonii H. & J. Groves, Man. Brit. hot. 9th ed.: 74 (1904). Type as for H. dubium var. maculatum sensu Bab., i.e. H. dubium Leers. H. quadrangulum subsp. erosum var. epunctatum Schinz in Vierteljahrsschr. Naturf. Ges. Zurich 49: 240, 241 (1905), nom. illegit. (Art. 52.1). Type as for H. desetangsii var. imperforatum Bonnet. Schinz, in order not to provide an unnecessary synonym and in support of the Rules of Nomenclature, named this form [sic] epunctatum. H. quadrangulum subsp. erosum var. punctatum Schinz in Vierteljahrsschr. Naturf. Ges. Zurich 49: 240, 241 (1905), nom. illegit. (Art. 52.1). Type as for H. quadrangulum subsp. obtusiusculum var. perforatum Tourlet. H. maculatum subsp. erosum sensu Schinz & Keller, Fl. Suisse: 381 (1909); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl., 120(1): 550(1911). H. maculatum subsp. erosum var. imperforatum (Bonnet) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120( 1 ): 553 (1911). Type as for H. desetangsii var. imperforatum Bonnet. H. maculatum subsp. erosum var. perforatum (Tourlet) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss. Math.-Naturwiss. Kl. 120(1): 553 (1911). Type as for H. quadrangulum subsp. obtusiusculum var. perforatum Tourlet. H. maculatum subsp. erosum var. imperforatum forma latisepalum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 554 (1911), in Mitt. Naturwiss. Vereines Steiermark 51: 221 (1915). Type: Austria, Styria, bei Doblad, 22 August 1909 (fr), Frohlich GZU 16202 (GZUMectotype, selected here). Note: none of Frohlich's combinations of his formae under subsp. obtusiusculum in 1915 (p. 221) were effective. H. maculatum subsp. erosum var. imperforatum forma latisepalum [sub]forma nigrum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl, 120(1): 554 (1911), in Mitt. Naturwiss. Vereines Steiermark 51: 22 1 ( 1 9 1 5). Type: Austria, Styria, Ragnitz bei Graz, July 1909 (fl), Frohlich GZU16194 (GZUMectotype, selected here). H. maculatum subsp. erosum var. imperforatum forma latisepalum [sub]forma lucidum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss, Math.-Naturwiss. Kl. 120(1): 554 (1911), in Mitt. Naturwiss. Vereines Steiermark 51: 221 (1915) sub var. imperforatum (Bon- net) Frohl. Type: Austria, Styria, bei Stattegg, nach Graz, 10 July 1910 (fl), Frohlich GZU 16200 (GZUI-lectotype, selected here). H. maculatum subsp. styriacum A. Frohl. in Mitt. Naturwiss. Vereines Steiermark 51: 222, 224 (1915); Hegi, ///. Fl. Mitt.-Eur. 5( 1 ): 5 1 8 (1925). Type: Austria, Styria, ad vicum Maria-Trost prope urbem Graz, c. 430 m, July 1911 (fl), Frohlich in Hayek, Fl. Stir. Exsicc. 1198 (GZUI-lectotype, selected here, BM!, GB!, H!; K!-photo- graphs; see also Martonfi et al., 1999: 340). H. maculatum subsp. obtusiusculum var. perforatum (Tourlet) A. Frohl. in Mitt. Naturwiss. Vereines Steiermark 51: 243 (1915). Type as for H. quadrangulum subsp. obtusiusculum var. perforatum Tourlet. H. maculatum subsp. obtusiusculum var. imperforatum (Bonnet) A. Frohl. in Mitt. Naturwiss. Vereines Steiermark 51: 243 (1915). Type as for H. desetangsii var. imperforatum Bonnet. H. dubium var. perforatum (Tourlet) Pugsley in J. Bot. 78: 35 (1940). Type as for H. quadrangulum subsp. obtusiusculum var. perforatum Tourlet. H. erosum (Schinz) O. Schwarz in Drudea 5: 63 (1965) pro parte excl. typum. Icones: Syme, Engl. Bot., 3rd ed. 2: t. 269 (1864); Ross-Craig, Draw. Br. PL 6: t. 8 (1952). STUDIES IN HYPERICUM Stems with subsidiary lines present or sometimes partly (or wholly?) absent. Leaves with rather dense tertiary venation (somewhat laxer than in subsp. maculatum); laminar gland dots pale, dense to very sparse or absent. Inflorescence branches widely ascending, making an angle of c. 50° with stem. Flowers 25-30 mm in diam. Sepals broadly to narrowly ovate, with apex finely eroded-denticulate or rarely entire. Petals usually distally unilaterally crenate with mar- ginal black glands in depressions?; laminar glands pale and black or rarely only pale, linear to striiform and sometimes punctiform. Styles 1-2 x ovary. 2n=32 (Robson, 1956, 1957, 1958a). Montane regions and valleys in the Alps, lowland further north, in meadows, rough pasture and damp woods; 0-c. 850 m. North-west Europe east to western Germany and Bohemia (except most of western Scotland), valleys of the Alps east to Styria, western Hungary. Introduced into Canada (British Columbia). SCOTLAND. E. Sutherland: Lairg, 24 July 1961 (fl), McCallum Webster 5726 (K). Easterness: Inverness, Clashnaharry, 3 October 1975 (fr), McCallum Webster 18700 (BM). Moray: Brodie Castle, 3 August 1961 (fl), McCallum Webster 5848 (K). S. Aberdeen: Drum, R. Dee by Old Manse, 15 August 1956 (fl & fr), Robson s.n (BM). W. Perth: Callander, 16 July 1989 (fl), N.F. Stewart s.n. (BM). Mid Ebudes: Mull, between Glen Forsa Lodge entrance and Forsa bridge, July 1984 (fl), Stirling s.n. (BM). Kintyre: near Fairy Isles, 28 July 1979 (fl), Kenneth s.n. (BM). Selkirk: banks of R. Ettrick near Selkirk, July 1926 (fl), Hayward s.n. (BM). Renfrew: near Gourock, 17 July 1899 (fl), Pugsley s.n. (BM). Dumfries: Applegarth, Perchall, 8 August 1957 (fl), Bangerter & Collett 381 (BM). Kirkcudbright: Rockcliffe, Au- gust 1999, Tregale s.n. (Herb. Tregale). ENGLAND. Westmorland: Tebay, Low Borrowbeck, 31 July 1991, Halliday 72/91 (BM). N.B. The records of subsp. maculatum in Cumbria (Halliday, 1997: 578) all proved to be subsp. obtusiusculum. NW Yorks.: near Richmond, 22 August 1 842 (fl & fr), J. Ward s.n. (BM). W. Lanes.: near Whittingham, Coney Garth Lane, September 1982 (fr), L. & P. Livermore s.n. (LIV). Cheshire: below Overton Scar, 15 July 1909 (fl), Wolley Dod s.n. (BM). Leicester: Ulverscroft, 10 August 1890 (e. fr), A.B. Jackson s.n. (BM). N. Lines.: Woodhall Spa, 6 August 1945 (fl), Alston s.n. (BM). Shropshire: near Ludlow, Bringwood Chase, August 1935 (e. fr), Ross s.n. (BM). Warwicks.: Brandon, 17 July 1925 (fl), Riddelsdell s.n. (BM). Hereford: near Ross, Chase Wood, 19 July 1915 (fl), Adamson s.n. (BM). E. Gloucs.: Brimscombe, 3 August 1953 (e. fr), Townsends.n. (K). Bedford: Potsgrove, 7 July 1 945 (fl),Dony s.n. (BM). W.Norfolk: Dersingham, 16 July 1949(fl), Libbey 3229 (BM). Oxford: near Hempton, 30 October 1 930 (fr), Riddelsdell s.n. (BM). N. Essex: near Quendon, 25 July 1942 (fl), Pugsley s.n. (BM). W. Kent: near Ide Hill, 13 July 1939 (fl), C.E. Britton 4322 (BM). W. Sussex: Midhurst, West Lavington, 7 July 1934 (fl), N.D. Simpson 34220 (BM). Dorset: between Crendell and Daggins, 2 August 1934 (e. fr), Good 484 (BM). E. Cornwall: near Rilla mill, 6 September 1 948 (fl), Alston s.n. (BM). WALES. Glamorgan: near Lisvane, 3 July 1968 (fl), Pinkard s.n. (H). Pembroke: Caneston wood, 11 August 1931 (fl), Alston s.n. (BM). Merioneth: Dolgelly, Bontnewydd, 8 August 1923 (fr), Barton s.n. (BM). Radnor: Llangullo station, 5 August 1956 (fl), P.C. & J. Hall s.n. (BM). Denbigh: Llanrwst, 9 August 1930 (fl), Holder 464 (LIV). Anglesey: Benllech, 5 July 1911 (fl), Cooper s.n. (BM). IRELAND. Kerry: Killarney, 30 July 1885 (fl), Linton s.n. (BM). Clare: 0.8 km W. of Ballynalacken Castle, 1 August 1962 (fl), Moore 459 (BM). Galway: Co. Galway, July 1854 (fl), More s.n. (BM). Wicklow: near Roundswood, 31 August 1962 (fl), McCallum Webster 7981 (K). Antrim: Belfast, Shaw's Bridge, 24 August 1872 (fr), Stewart (BM). Fermanagh: Inver Lough, 19 July 1957 (fl), Meikle 2127 (K). NETHERLANDS. Overijssel: near Mokkelengoor, S. of Ijpelo, 14 June 1947 (fl), van Royen s.n. (H). Gelderland: s' Hertogenbosch, Meteren, Tielenwaard, 22 July 1974 (1. fl), Hekking s.n. (BM). BELGIUM. Oost Vlanderen: Hamme Mille (?), 11 July 1953 (fl), Pierart s.n. (K). Liege: Spa, September 1876 (fr), C.H. Wright s.n. (K). Hainault: bois d'Obourg, 4 July 1 867 (e. fr), Martinis in PI. rar. crit. Belg. VI, 269 (BM). Namur: Crupet, pres du confluent du Crupet et du Bocq, 8 August 1978 (fl), Lawalree 21100 (BM, *BR). Luxembourg: La Roche [-en- Ardennes], August 1960 (1. fl), Dubois 1635 (BM, *BR). 75 FRANCE. Pas de Calais: Hersin[-Coupigny], 15 July 1917 (1. fl), G.C. Brown s.n. (BM). Aisne: la foret de Retz pres Villers-Cotterets, 3 September 1 878 (e. fr), Bonnet s.n. (K). Seine-et-Oise: Bois de Meudon, pres 1'Etang de Colm Porche, 19 August 1877 (fl), Delacour s. n. (K). Loire-Atlantique: bord de la Divatte, 5 August 1886 (fl & e. fr), Lloyd s.n. (BM). Orne: Alengon, route de Fresnay, 19 July 1891 (1. fl), Beaudoin in Soc. Dauph. II 599 (BM). Allier: Cresset, bord du Sichon, 27 July 1 867 (e. fr), Genevier s.n. (BM). Doubs: Mouthier, 15 July 1927 (fl), Williams s.n. (BM). GERMANY. Hessen: Konigstein/Taunus, NW der Stadt, 20 August 1961 (fl), Schaarschmidt 257 (FR); Kreis Limburg-Weilburg, Weilburger Lahntal, Weilburg, 260 m, 22 July 1978 (fl), Kalheber 78-527 (H). Thuringen: Weimar, Ettersburg, September 1923, Bornmiiller s.n. (JE). Baden- Wiirttemberg: Schwarzwald, Heselbach, 9 August 1931 (fr), Williams s.n. (BM). Bayern: Schwaben, Leipheim/Donau, Donauried, 7 July 1961 (fl), Doppelbaur4&8 (H); Oberbayern, nahe dem Westiiferdes Staffelsees, 650 m, 1 1 August 1957 (fl), Launert s.n. (BM). SWITZERLAND. Valais: Col de Balme, 30 August 1868 (fr), G. Engelmann s.n. (WIS). Zurich: oberhalb Oberglatt, c. 420 m, 3 September 1936 (fr), W. Koch s.n. (H). St. Gallen: Fiirstenland, Gem. Waldkirch, Tannenberg, c. 850 m, 31 July 1948 (fl), W. Koch 48/473 (H). AUSTRIA. Steiermark: ad vicum Maria-Trost prope urbem Graz, c. 430 m,July 1911 (f\),Frohlich in Hayek, Fl. Stir. Exs. 1198 (BM,GB photograph, GZU, H, K photograph). Karnten: fide Frohlich (1915: 223). CZECH REPUBLIC. Bohemia: *Doupovske Hory Mts, E. from Karlovy Vary, near Brazec, c. 685 m, 12 July 1996, Mdrtonfi2\25 (KO). HUNGARY. Veszprem: *Montes Bakony, inter stationem ferroviae Porva- Csesznek ad pagum Borzavar, c. 400 m, 9 July 1934, Polgdr in BP 261467 (BP); *in fossis Nadaser pr. villa Dobospuszta iuxta pag. Saska, 27 July 1954, Szalay s.n. (ZIRC). CANADA (introduced). British Columbia: Prince Rupert, September 1 939 (fl), tfmgUBC 21 544 (UBC);Bumaby Mountain, 1 August 1990(l.fl), Lomer UBC 202196 (UBC); Pemberton, 2 August 1946 (fr), Eastham UBC 14970 (UBC); North Vancouver, Capitano R. estuary, 8 August 1990 (fl), Lomer UBC 202195 (UBC). The presence of linear glands (sometimes pale) in the petals and the greater frequency of pale glands in the leaves suggest that subsp. obtusiusculum has evolved from subsp. immaculatum, not, as I thought from my own observations and experiment, from subsp. maculatum. The plant obtained by colchicine doubling of the chro- mosomes (Robson, 1957, 1958a) had broad leaves, broad entire sepals and black petal laminar glands, as was to be expected starting from var. maculatum; but the inflorescence branching was wide and the petal glands (as far as I can remember in the absence of a specimen) were mostly linear. The presence of broad entire sepals in this experimental plant, in Leers's (lost) type of H. dubium and in a collection from Applegarth, Dumfries, SW Scotland (Bangerter & Collett 381 (BM)), suggests that the frequent occurrence today of narrow oblong sepals and incomplete subsidiary stem lines may have resulted from recent introgression from H. perforatum. Schwarz (1965) recorded 2n=16 for 'H. erosuni1 (= subsp. obtusiusculum) and 2n=24 for its hybrid with H. perforatum, adding that all his material gave the diploid number. Unless his results are duplicated in the future (which would necessitate a revision of the view adopted here), one must assume that his material was misidentified. Ix. Hypericum x laschii A. Frohl. in Mitt. Naturwiss. Vereines Steiermark 51: 244, 245 (1915), in Preslia 32: 97-99, ff. 1, 2 (1960); Gu§ul. & Nyar. in Savul., Fl. R. P. Roman. 4: 46 (1956). Type as for H. tetraptero-quadrangulum Lasch. 1. H. maculatum Crantz x 3. H. tetrapterum Fr. H. tetraptero-quadrangulum Lasch in Linnaea 4: 414 (1829). Type: Germany, Brandenburg, Driesen, 1829 (fl & fr), Lasch s.n. (B|- holotype). H. maculatum Crantz x H. acutum Moench sensu A. Frohl. in 76 N.K.B. ROBSON Sitzungsber. Kaiserl. Akad. Wiss., Math-Naturwiss. Kl. 120(1): 584(1911). In comparison with H. tetrapterum: Stems relatively narrowly 4- winged. Leaves usually rather densely veined, with pale glands numerous, some large and some small. Sepals relatively broad, obtuse to rather acute. Flowers about twice as large. For distribution see under nothoformae. The name H. x laschii applies to all hybrids between H. tetrapterum and H. maculatum, but only crosses involving subspp. maculatum and obtusiusculum are recorded. It therefore occurs in diploid (2n=16) and triploid (2n=24) forms, depending on which subspecies of H. maculatum is involved. Nearly all records are of crosses involving subsp. maculatum (diploid), but Frb'hlich (1911) described one that appears to involve subsp. obtusiusculum (i.e. it would be triploid). The syntype populations of H x desetangsii Lamotte from near Troyes (Aube, France) appear to be an example of this latter cross (nothoforma froelichii), not, as all authors after Bonnet ( 1 879) have assumed, of H. maculatum x perforatum. Bonnet, himself, stated that it was 'intermediate entre la suivante [H. tetrapterum] et I'//, quadrangulum [H. maculatum]' .In the interests of nomenclatural stability, however, I have retained the current usage. The type of//. intermedium Bellynck has also proved to belong to //. x laschii nforma froelichii. A proposal to conserve the name H. x desetangsii with a new type will therefore have to be made. Such a proposal would also conserve it against possible threats from H. mixtum Du Moulin and H. commutatum Nolte (see p. 104). 1 xa. Hypericum x laschii nothoforma laschii 1 a H. maculatum subsp. maculatum x 3. H. tetrapterum Fr. H. maculatum Crantz subsp. typicum A. Frohl. x acutum Moench sensu A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl. 120(1): 582, f. 12(1911). Icon: Rchb., Ic. fl. germ. helv. 6: t. 344, no. 5178 (1842-1844) [quadrangulo-tetrapterum] . Intermediate in height and habit between the parents. Leaves with tertiary venation usually relatively densely reticulate. Sepals c. 0.5- 0.75 x petals, obtuse to acute, with pale striiform laminar glands. Petals entire, with pale or pale and black punctiform and striiform laminar glands. FRANCE. Cher: Acheres, 20 July 1909 (fl), Imbault s.n. (FR). Rhone: Amas, August, 1878 (fl), GandogerP}. Gall. Exs. 407 (H). Tarn: Angles, 23 July 191 1 (fl & fr), d'Alleizette s.n. (CLF). SWITZERLAND. Schwyz (Einsiedlen), Zurich? (Geissboden), fide Frohlich (1911). AUSTRIA. Salzburg (Salzburg), Oberosterreich (\sch\),fide Frohlich (1911). CZECH REPUBLIC. Moravia: Bad Luhaovice,/kfe Frohlich (1960). SLOVAKIA. Bad Kovaova (near Z\o\en),fide Frohlich (1960). GERMANY. Bayern: Kreis Augsburg, ostlich Reinhartshausen, 5 Au- gust 1966 (fl), Petermann s.n. (H). Thiiringen: Feun (with parents), August 1977 (fl), Schulze s.n. (FR). Berlin: Bredower Forst bei Berlin, August 1864 (fr), O. Kuntze (BM). POLAND. Jelenia Gora: Schoenau [Zwierzawa], Polnisch-Hundorf, 4 June 1897 (fl), Scholz s.n. (H). Zielona Gora: Griinberg [Zielona Gora], Lindenbusch, 3 1 August 1 892, Hellwig in Collier 576 (JE). DENMARK. Fyn: Svendborg Fioniae, July 1872 (1. fl), Lange s.n. (K). SWEDEN. Malmohus: Helsingborg, 1 September 1897 (fl & fr), Strandmark s.n. (H): also several other collections from extreme SE Skane. This hybrid may be expected to occur anywhere within the wide area of overlap of the distributions of the parents; but, on account of the differing habitat requirements of the parents, it is rare. Ixb. Hypericum x laschii nothoforma froelichii N. Robson, hybr. nov. Type: Austria, Styria, Ragnitz bei Graz, c. 400 m, 3 1 July 1909 (fl & fr), Frohlich GZU16255 (GZU!-holotype). Ic. H. maculatum subsp. obtusiusculum x 3. H. tetrapterum Fr. H. quadrangulum sensu Des Etangs in Mem. Soc. Agric. Dep. Aube 10:24(1841). ?//. tetrapterum var. [0] intermedium Coss. & Germ., Fl. descr. anal. Paris: 64 (1845). This variety is described as intermediate between H. tetrapterum and H. quadrangulum (i.e. H. maculatum), of which the only subspecies in the Paris region is subsp. obtusiusculum. H. intermedium Bellynck, Fl. Namur. 31 (1855); Gren., Fl.jurass. 1: 154 (1863); Nyman, Consp. fl. eur.: 133 (1878); non Steud. (1847). Type: Belgium, Namur, Wepion, Dave s.n. (BR!- holotype). H. maculatum Crantz subsp. erosum sensu Schinz & Keller, x H. acutum Moench sensu A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 588, f. 13 (1911). Intermediate in height (c. 0.6 m) and habit between the parents. Leaves with tertiary venation laxly reticulate. Sepals c. 0.5 x petals, broadly to narrowly oblong or narrowly elliptic (sometimes in same flower), obtuse to acute or subacuminate, entire to apiculate-eroded, with pale punctiform and striiform and single black linear laminar glands. Very rare, probably as a result of the differing habitat requirements of the parents. Austria. AUSTRIA. Steiermark: Ragnitz bei Graz, 15 July 1910 (fl), Frohlich GZU 16256(GZU). This hybrid was known from the type locality only (but probably includes the types of//, x desetangsii and H. intermedium). Frohlich (1911) described it as agreeing almost completely with the var. maculatum hybrids, but that (i) it was growing crowded together with both putative parents, so that cross-pollination was quite possible, (ii) it had characters intermediate between them, and (iii) its flowering dates were also intermediate. The latest of Frohlich's collections of this hybrid (26 July 1914) was identified by him as //. maculatum subsp. styriacum x acutum. 1 . Hypericum maculatum Crantz x 5. H. perforatum L. (see 5x, p. 102). 2. Hypericum undulatum Schousb. ex Willd., Enum. pi. Berol.: 810 (1809); Spach in Ann. Sci. nat. Bot. 5: 357 (1836); Bab., Man. Brit, hot.: 67 (1843), in J. Bot. 2:: 97, t. 16 (1864); Archer Briggs in / Bot. 2: 45 (1864); Hance in J. Bot. 5: 299 (1867); Nyman, Consp. fl. eur. 1: 133 (1878); Lange in Willk. & Lange, Prodr.fl. hisp. 3: 59 1 ( 1 878); Bonnet in Bull. Soc. Bot. France 25: 28 (1879); A. Frohl. in Sitzungsber. Kaiserl Akad. Wiss., Math.- Naturwiss. Kl. 120(1): 583 (1911); Ross-Craig, Draw. Br. PL 6: t. 10 (1952); N. Robson in Tutin et al., Fl. Europ. 2: 268 (1968); Franco, Nova Fl. Portugal 1: 453 (1971); A. Ramos in Trab. Dept. Bot. Univ. Complut. Madrid 12: 49 (1983), in Acta Bot. Malac. 11: 167, f. 6d (1986), in Castroviejo et al., Fl. Iberica 3: 167 (1989); Valdes, Talavera & Galiano, Fl. Vase. Andal. Occ. 1: 3 16 + map & fig. (1987); B. Tebbs in Press & Short, Fl. Madeira: 222 (1994); N. Robson in Cullen et al., Europ. Gdn Fl. 4: 60 (1995). Type: Morocco, Tangier, Schousboe s.n. (C-holotype; G- BOISSI-isotype). Fig. 7, Map 3. H. quadrangulum var. [e] undulatum (Schousb. ex Willd.) Choisy in DC., Prodr. 1: 548 (1824); Colm., Apunt.fl. Castillas: 37 (1849). STUDIES IN HYPERICUM 77 m Fig. 7 A. H. undulatum van undulatum: (a) habit: (b) stem section; (c) upper leaf; (d) sepal; (e) petal. B. H. tetrapterum var. tetrapterum: (f) habit; (g) stem section; (h) sepal; (i), petal; (j) capsule. C. H. triquetrifolium: (k) habit: (1) sepal; (m) petal (a. c, f, k x 2/3; b, g x 2; rest x 6). A. (a, b) Go\der & Jury 545; (c) Reverchon 238; (d, e) Davis 54961. B. (f, g) Gadeceau s.n.; (h, I) Granvik s.n.; (j) Coste 801. C. Davis & Coode 39246. 78 H. decipiens H. Watson in London J. Bot. 3: 588 (1844). Types: Azores, Flores (fl), H. Watson s.n. (KMectotype, selected here); Fayal (st), J.C. Hunt s.n. (K!-syntype). H. quadrangulum sensu Colm., Recuerd. hot. Galicia: 10 (1850); Lowe, Man.fl. Madeira: 79 (1868) pro parte excl. typum. H. boeticum sensu Syme, Engl. Bot., 3rd ed. 2:153, t. 270bis ( 1 864); H. Wats, in Godman, Nat. hist. Azores: 143 (1870); Bull in J. Bot. (Lond.) 13: 296 (1875); Trelease in Rep. Mo. hot. Gdn, no. 8: 100 ( 1 897); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10; t. 4 f. 27 (1932), 11: 176 (1933), 12: 88 (1934), in Pflanzenareale 4(1): Karte6a(1933). H. tetrapterum var. [|3] undulatum (Schousb. ex Willd.) Cout. in Bolm Soc. Brot. 12: 21-22 (1895), comb, illegit. (Art. 58.2). H. acutum subsp. undulatum (Schousb. ex Willd.) Rouy in Rouy & Foucaud, Fl. France 3: 337 (1896); Cout., Fl. Portugal: 404 (1913), 2nd ed.: 482 (1939); comb, illegit. (Art. 58.2). H. desetangsii sensu Lloyd, Fl. Quest France, 5th ed.: 72 (1898), non Lamotte (1874). H. acutum var. undulatum (Schousb. ex Willd.) Pau in Bol. Soc. Aragonese Ci. Nat. 8: 118 (1909), comb, illegit. (Art. 58.2). H. quadrangulum var. tetrapterum forma undulatum (Schousb. ex Willd.) Borg, Descr.fl. Malt, isi: 248 (1927). H. maculatum subsp. undulatum (Schousb. ex Willd.) P. Fourn., Quatrefl. France: 453 (1946). H. tetrapterum subsp. undulatum (Schousb. ex Willd.) P. Silva in Agron. Lusit. 12(2): 271 (1951) comb, illegit. (Art. 58.2). H. quadrangulum subsp. undulatum (Schousb. ex Willd.) Goday & Carbonell in Anales Inst. Bot. Cavanilles 19: 380 (1961). Icones: Bab. in J. Bot. (Lond.) 2: 97, t. 16 (1864); Ross-Craig, Draw. Br. PL 6: t. 10(1952). Perennial herb 0.015-0. 12 m tall, erect to decumbent from creeping and rooting base, with stems numerous to few, initially unbranched below inflorescence, later branched above. Stems narrowly and ± equally 4-winged, with black glands on the wings and sometimes elsewhere; internodes 12-65(-80) mm, longer than leaves. Leaves sessile; lamina 6-40 x (3-)7-14(-20) mm, elliptic or narrowly ovate to narrowly oblong or rarely suborbicular, paler beneath, thickly to thinly chartaceous; apex rounded to obtuse or rarely subacute, margin usually crisped-undulate, base rounded to cordate- subamplexicaul; venation: 3(4) pairs of main laterals from lower quarter of midrib, tertiary reticulation rather dense; laminar glands pale, dense, medium to small; intramarginal glands black, close, irregular in size. Inflorescence 1-c. 40-flowered, from 1-3 nodes, with flowering branches ascending to horizontal from up to 6 nodes, the whole cylindric to subcorymbiform, lax; pedicels 1-2.5 mm; bracts and bracteoles 2-4 mm long, lanceolate, entire, with (0-)2-8 black glands. Flowers 12-17 mm in diam., stellate; buds ellipsoid, obtuse. Sepals 5, subequal, 3-5.5 x 1-1.8 mm, narrowly elliptic to lanceolate, acute to acuminate, entire, erect in bud and fruit; veins (3)5-7, when 5 the outer pair branched; laminar glands punctiform, all pale or nearly always (3-)6-20(-28) black; intramarginal glands black, few or absent, rarely with terminal black gland. Petals 5, bright yellow, nearly always tinged red dorsally, (5-)7-10 x 3-4 mm, c. 2-2.5 x sepals, obovate-oblanceolate, entire; laminar glands black, punctiform, few or absent; marginal glands black, up to 6, scattered, or absent. Stamens c. 25^0, '3 '-fascicled, longest (4.5-) 5-8 mm, c. 0.75-0.9 x petals. Ovary 3-locular, 2-3.5 x 1-1.5 mm, narrowly ovoid to narrowly ellipsoid; styles 3, free, 3-4 mm, 1-1 .3 x ovary, spreading; stigmas narrowly capitate. Capsule 5-8 x 3.5^1 mm, c. 1 .65 x sepals, ovoid; valves with longitudinal linear vittae. Seeds yellow-brown, 0.6-0.8 mm, cylindric, not carinate or appendiculate; testa linear-foveolate. 2n=16, 32 (ploidy apparently N.K.B. ROBSON not correlated with morphology). The record of 2n=8, n=4 for this species by Guillen Oterino et al. (1997) is difficult to interpret. The count appears to be genuine, being vouched for and illustrated by drawings; but no other report of such a hypoploid number has been made elsewhere in the genus. Non-calcareous marshes, mildly acidic bogs; 0-2700 m. Extreme western Europe and North Africa: England (Cornwall, W. Devon), Wales (Pembroke, Cardigan, Merioneth), Ireland (W. Cork?, see below), France (W. Brittany, see below), central and W. Spain, Portugal, W. Algeria, N. Morocco, Madeira, Azores. Hypericum undulatum occurs as two very unequal varieties. The vast majority of the species (var. undulatum) has relatively narrow undulate leaves and red-tinged flower buds; but a population in the south of Spain (Sierra Nevada) has broader plane-margined leaves and untinged flower buds. This population, which Boissier named Hypericum boeticum, appears to be intermediate in some characters between 2. H. undulatum and 3. H. tetrapterum', and since there are some plants with intermediate characters elsewhere in the Iberian Peninsula, some (particularly Spanish and Portuguese) botanists reduced these taxa to subspecies. The real 'link' between these species, however, would seem to be the plant from southern Italy and Sicily known as H. neapolitanum Ten., which is the most luxuriant form of H. tetrapterum. Hypericum boeticum, on the other hand, has flowers and inflorescence typical of H. undulatum in size and shape; and the broad leaves (which are linked in the population with ones more typical of H. undulatum) should be interpreted as a development in that species parallel with the broadening of the leaves in more advanced forms of H. tetrapterum. The plane mar- gins and plain yellow petals are thus a reversion, not an intermediate condition. Populations that are truly intermediate between these species and apparently hybrid in origin occur in scattered localities in the Iberian Peninsula (see below). It seems appropriate, then, to treat the Sierra Nevada population as a variety (var. boeticum). 2a. Hypericum undulatum var. undulatum Plant erect. Leaves (3-)4—14(-18) mm broad, elliptic to ovate, margin crisped-undulate. Petals red-tinged dorsally. 0-1750 m. Distribution of the species except extreme south Spain. The record for Ireland (Co. Cork) is based on a specimen in K and that for France (Brittany) on specimens of a Lloyd collection in BM and ANG. The former, from Glengarriff, could just possibly be due to mislabelling, but the latter appears to be a correct record. Both collections were misidentified, the former as H. tetrapterum and the latter as H. x desetangsii. Lloyd, in Fl. Quest France 5th ed., cites several localities round Brest for H. x desetangsii, his description of which agrees with H. undulatum, not H. x desetangsii. It seems, therefore, that the 'Atlantic' distribution of H. undulatum includes (or included) the extreme 'Atlantic' part of France and possibly SW Ireland. WALES. Merioneth: Morfa Arthog (Arthog Bog), 2 August 1953 (fl), Benoit s.n. (K). Cardigan: 0.4 km W. of Trawscoed Station, 1933, Salter (Salter, 1935). Pembroke: between St. David's and Solva, 8 August 1931 (fl), Alston s.n. (BM); near Narberth, 1885 (fl), Barrett s.n. (BM). ENGLAND. W. Cornwall: Penzance, Tremethick Moor, 7 August 1911 (fl), Barton s.n. (BM, K); N. of Camborne, Reskadinnick, 4 August 1971 (fl), Murphy s.n. (BM). E. Cornwall: Tywardreath Marsh, 23 July 1935 (fl), Grose 1278 (BM); near Falmouth, Penjerrick, August 1871 (e. fr), T.A. Briggs s.n. (BM). S. Devon: Morwellham, 9 August 1901 (fl), Hume 3944 (BM); Egg Buckland, Common Wood, 3 1 August 1 872 (1. fl), T.A. Briggs s.n. (BM). N. Devon: Torrington, 9 September 1935 (fl & fr), Chappie s.n. (BM). STUDIES IN HYPERICUM 79 Map 3 2. H. undulatum. a. var. undulatum • , other records O; b. var. boeticum A. 2. H. undulatum x 3. H. tetrapterum IRELAND? W. Cork: Glengariff, 18 August 1908, C. Bucknall s.n. (K). See Robson (1958b); but this record is not mentioned by Perring (1996). FRANCE. Finistere: environs de Brest, end July 1886 (fl), Blanchard in Lloyd s.n. (ANG, BM); cult, du jardin d'un pied de Brest, end July 1890 (fl), Lloyd s.n. (ANG). SPAIN. Pontevedra: Pontevedra, 27 July 1930 (fl & fr), Buch s.n. (H). Coruna: La Coruna, near Coruna, August 1888 (fl), Guardia s.n. (BM). Oviedo: gorge of Pena de la Paloma, near Tineo, 16 July 1927 (fl), Wilmott s.n. (BM). Cantabria: Santander, Picos de Europa, Fuente De, 1200 m, 1 August 1969 (fl), Grimes 229 (BM). Leon: El Bierzo, Villanueva to San Adrian, 900m, 10 July 1933, Rothmaler 41 1 (JE). Soria: Vinuesa, Santa Ines, in loco Gargantillas, 1450 m, 15 July 1989 (fl), Segura Zubizarreta in Soc. Exch. PI. Vase. Eur. Bass. Med., Fasc. 23, 14273 (RNG). La Rioja: Valle del Rio Iregna, between Pantano Ortigosa and Lumbreras, 1 July 1958 (bud), Sandwith 5265 (K). Avila: 25 km SE of Avila, 3 August 1962 (fl), Scott 359 (BM). Zaragoza: Catalayud, Sierra de Vicort, 23 July 1910 (fl), Viciosa in Sennen 1122 (FR, H, JE). Tarragona: Cambrils, acequia 'Canaleta del Civil', Partida de 1'Ardiaca, 12 August 1926 (fl), Teodora s.n. (BM). Guadalajara: Aldeanuevade Atienza, 1 1 September 1965 (fl), Silvestre s.n. (RNG, SEV*). Huelva: between Palos de la Frontera and Punta Arenilla, 8 October 1968 (fl & fr), Bramwell & Valdes 173 (RNG). PORTUGAL. Throughout, but rarer in SE (Franco, 1971: 453). Minho: Viana do Castelo, fl. Minii prope Seixas, 29 July 1938, Rothmaler s.n. (JE). Tras-os-Montes: Vila Real, Chaves, prope Sapiaes, 800 m, 31 July 1938, Rothmaler 1 4002 (JE). Douro Literal: S. of Oporto, Granja, 3 1 July 1910 (fl), Johnston s.n. (BM). Beira Vita: Guarda, Serra da Estrela, 5 km from Pousada de Sao Louren?o to Gouveia, 1 360 m, 20 August 1 982 (fl), Goyder &Jury 545 (BM, RNG). Beira Eaixa.fide Ramos (1993). Beira Literal: Coimbra, 20 September 1 950 (e. fr), /. & A. Matos s.n. (BM). Estremadura: Lisboa, Torres Vedras prope Puteiro, 3 July 1936, Rothmaler s.n. (JE). Ribatejo, Alta Alentejo, Baixa Alentejo: /?<&> Ramos (1993). Algarve: Faro, ribeira de S. Cristovao, August 1 882 (e. fr), Peres in Fl. Lusit. (4° anno) 562 (BM). MOROCCO. Tanger: Tetuan, Cap Spartel, 200 m, 3 July 1989 (fl), Garcia, Silvestre & Talavera 110/89 (SEV). Rif: W. side of Ketama, 1500- 1600m, 8 July 1973 (f\),Davis 54961 (BM, E*). Moyen Atlas: RasChedaya, Bab Bouidir, 1400 m, 2 August 1968 (fl), Fry' 193 (BM). ALGERIA. Oran: Gambetta, 9 June 1913 (fl), Faure s.n. (H). MADEIRA. S. of Ribeiro Frio, 3 km N. of Poiso, 1 100 m, 3 1 July 1979 (fl), Jarvis & Wild 820 (MO); Queimadas, 900 m, 17 August 1961 (fl), Coleridge 6 (BM; K); Santa Madalena, 4 July 1957 (fl), Malmberg s.n. (H); Passa d'Area, 12 August 1855 (fl), Lowe s.n. (BM). AZORES. Santa Maria: Ribeira da Vila, 50 m, 2 May 1972 (st), Goncalves 3947 (BM). Sao Miguel: Seara near Sete Cidades, Lagoa Azul, W. bank, 243 m, 26 July 1970 (fl), Dolman 329 (BM). Terceira: Terreiro dos Prados, 450 m, 10 July 1971 (fl), Goncalves 3148 (BM). Sao Jorge: Camolico do Pinheiro, 400 m, 4 September 1971 (e. fr), Goncalves 3442 (BM). Pico: Madalena, 9 July 1938 (fl), Cederkreutz s.n. (H). Faial: Caldeira, 500 m, 7 October 1971 (fl & e. fr), Convolves 3652 (BM). Flores: Fazenda, Santa Cruz to Lajes, 200 m, 3-8 September 1985 (fl), Parry 103 (BM). Corvo: Weather station, 200 m, 3 September 1967 (fl), Pryor 170 (BM). 2b. Hypericum undulatum var. [pV| boeticum (Boiss.) Lange in Willk. & Lange, Prodr.fl. hisp.: 591 (1878); N. Robson in Tutin et al., Fl. europ. 2: 268 (1968) in adnot. Type as for//, boeticum Boiss. Map 3. H. boeticum Boiss., Elench. pi. nov.: 25 (1838), Voy. hot. Espagne 1: t. 34 (1839), 2: 1 14(1 839); Nyman, Consp.fl. eur. 1: 133(1878); Burdet, Charpin & Jacquem. in Candollea 39: 788 (1984). Types: Spain, Granada, prope Yunquera prov. Malacitanae, 1 837, Boissier & Renter s.n. (G-BOISS!-lectotype, N. Robson, 22 June 1955); prope Alhaurin, Carratraca et thermas Vilo, Boissier & Reuter s.n. (G-BOISS!-syntype);secusfluviumJenil inter Granada etGuejar, Boissier & Reuter s.n. (G-BOISS !-syntype); inter Berja et Adra in ditione Alpujarras dicta, Boissier & Reuter s.n. (G-BOISS!- syntype). 'In humidis mont. Regn. Granat. vulg., Aest. 1837' (fl), Herb. Boissier (K) is likely to be a duplicate of one of the above collections. H. tetrapterum var. [f3] rotundifolium Willk. in Willk. & Lange, 80 N.K.B. ROBSON Prodr.fl. hisp. 3: 591 (1878). Type: Spain, Granada, Barranco de Gualnon, Dehesa de S. Geronimo, 1800 m, July 1844, Willkomm s.n. (G-lectotype, selected here; BM!-isolectotype); Monachil, Funk s.n. (G-syntype). Willkomm's original (unpublished) name for this taxon would appear to have been var. nivale, according to the label on Willkomm, PI. Hisp. Exsicc. 1844-6 no. 358 (BM). H. tetrapterum var. boeticum (Boiss.) Cout. in Bolm Soc. Brot. 12: 21-22(1895). H. acutum var. [p] rotundifolium (Willk.) Schinz in Bull. Herb. Boissier II, 3: 16 (1902); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120: 582 (1911), comb, illegit. H. acutum subsp. boeticum (Boiss.) Cout., Fl. Portugal: 404 (1913), 2nd ed.: 482 (1939), comb, illegit. H. baeticum sensu auct. plur., sphalm. Icon: Boiss., Voy. hot. Espagne 1: t. 34 (1839). Plant erect to decumbent. Leaves (4-)8-20 mm broad, broadly elliptic to orbicular, margin plane. Petals not red-tinged dorsally. 1500-2700 m. Spain (Andalucia). SPAIN. Granada: barranco del no Segura, 1500 m, July 1906 (e. fl), Reverchon 238 p.p. (BM, FR, H); Capileira, c. 1750 m, 18 June 1962 (st), Rovainen s.n. (H). Malaga: Sierra de Yunquera, 3 July 1 890 (fl), Reverchon 238 p.p. (JE, K). When H. boeticum is regarded as an extreme development of H. undulatum, it becomes evident that Willkomm's var. rotundifolium is not part of//, tetrapterum but forms an extreme development of//. undulatum var. boeticum. Indeed there seems to be no break in variation between typical var. boeticum and the higher-altitude 'var.' rotundifolium, which differs in its decumbent habit, smaller leaves and fewer smaller flowers. Examples of this form (additional to the syntypes cited above) are: SPAIN. Granada: Sierra Nevada, pass between Borreguiles de San Juan and Penones de San Francisco, 2700 m, 23 August 1948 (e. fr), Heywood & Davis 983 (BM); Rio Monachil, 2300 m, 28 July 1950 (fl), Rovainen s.n. (H). 2. H. undulatum x 3. H. tetrapterum Map 3. The following specimen has characters intermediate between 2a. //. undulatum var. undulatum and 4a. H. tetrapterum var. tetrapterum and is assumed to be a hybrid between these taxa: SPAIN. Viscaya: Bilbao mont., 20 July 1905 (fr), Elias 801 (BM). An intermediate specimen from Andalucia, however, is probably a hybrid involving 2b. H. undulatum var. boeticum: SPAIN. Granada: Andalucia, nearTrevelez, Puerta de Granada, 1 260 m, 4 August 1969 (fl), Brinton-Lee 940 (BM). 3. Hypericum tetrapterum Fr., Novit. fl. suec.: 94 (1823), ed. altera: 236 (1828); Spach, Hist. nat. veg., Phan. 5: 387 (1836), Ann. Sci. Nat. Bot. 5: 357 (1836); W. Koch, Syn.fl. germ.: 134 (1837); Rchb., Ic.fl. germ. helv. 6: t. 344, f. 5179 (1844); Godr. in Gren. & Godr., Fl. France 1: 34 (1 847); Bellynck, FL Namur: 31 (1855); Syme, Engl. Bot. 3rd ed. 2: 152, t. 270 (1864); Boissier, Fl orient. 1: 805 (1867); Bab., Man. Brit, hot., 6th ed.: 66 (1867); Lange in Willk. & Lange, Prodr. fl. hisp. 3: 591 (1878); Bonnet in Bull. Soc. Bot. France 25: 279 (1879); Schinz & Keller, Fl. Schweiz: 326 (1900); Coste, Fl. France 1: 259 (1901); Stefanoff in God. Agr.-les. Fak. Univ. Soflya 10: t. 4f. 26 (1932), 11: 175 (1933), 12: 87 (1934), in Pflanzenareale 4(1): Karte 6a (1933); Mansfeld in Repert. Spec. Nov. Regni Veg. 47: 278 (1939); Pugsley in J. Bot. 78: 35 (1940); Ross-Craig, Draw. Br. PI. 6: t. 9 (1952); N. Robson in Davis, Fl. Turkey 2: 399 (1967), in Tutin et al., Fl. Europ. 2: 268 (1968), in Rech.f., Fl. Iranica, Guttif.: 17 ( 1968), in Townsend & Guest, Fl. Iraq 4: 378 (1980); in Cullen et al., Europ. Gdn Fl. 4: 60 (1995), in Wisskirchen & Haeupler, Standardliste Farn- u. Blutenpfl. Deutschl.: 270 (1998); Pignatti, Fl. Italia 1: 177 (1982); Zeleny et al. in Futak & Bertova, Fl. Slovenska 3: 306, t. 37 f. 2, map 57 (1982); Hagemann in Flora 173: 115, tt. 5B, 15-18, 39 (1983); A. Ramos in Trab. Dept. Bot. Univ. Complut. Madrid 12: 49 (1983), in Acta Bot. Malac. 11: 167, t. 6c (1986), in Castroviejo et al., Fl. Iberica 3: 168, t. 46 (1989); Azadi in Fl. Iran 27: 9 + 1. + map (1999). Type: Sweden, Skane, in silva Resten, 1819 (?), Fries s.n. (UPS-holotype). Fig. 7, Map 4. H. quadrangulum L., Sp. pi: 785 (1753); Huds., Fl. angl: 292 (1762); Crantz, Stirp. austr. 2: 65 (1763); Leers, Fl. herborn., 2nd ed.: 169 (1789); Vill., Hist. pi. Dauphine 3: 497 (1789); Sm., Engl. Bot.: t. 370 (1797), Fl. Brit. 2: 801 (1800); DC., Fl. France 4: 862 (1805); Choisy, Prodr. monogr.fam. Hyperic.: 47 (1821), in DC., Prodr. 1: 548 (1824); Bab. in Trans, hot. Soc. Edinburgh 1: 83 (1841),Man. Brit, tor.: 57 (1843); Syme,Eng/. Bot., 3rded. 2: 152, t. 270 (1864); Rendle & Britten in J. Bot. 45: 436 (1907); Grande in Boll. Orto hot. Regia Univ. Napoli 8: 74 (1926); Pugsley in J. Bot. 78: 25 (1940); N. Robson in Greuter, Burdet & Long, Med-Checklist 3: 27 1 ( 1 986); nom. rejic. (see N. Robson in Taxon 39: 135 (1990) and Brummitt in Taxon 43: 114 (1994)). Type: Netherlands (?), Hon. Cliff. 380 (BM!-holotype). H. quadrangulare sensu L., Diss. Hypericum: 5 (1776), Amoen. Acad. 8: 322 (1785); Murray, Syst. veg., 14th ed.: 584 (1784); Lam.,Dict. 4: 163 (1797); Willd., Sp. />/. 3: 1459(1802), sphalm. Type as for H. quadrangulum L. H. acutum Moench, Methodus: 128 (1794); Beck, Fl. Nieder- Osterreich 2( 1 ): 530 ( 1 892); Rouy in Rouy & Fouc., Fl. France 3: 335 (1896); Burnat, Fl. Alpes marit. 2: 28 (1896); Schinz in Bull. Herb. Boissier II, 3: 16 (1902), in Vierteljahrsschr. Naturf. Ges. Zurich 49: 241 (1905); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 580(1911);ThellunginA//g. Bot. Z. Syst. 1912: 19(1912); Coutinho, Fl. Portugal: 404 (1913), 2nd ed.: 48 1 ( 1 939); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 179 (1925); Hegi, ///. Fl. Mitt.-Eur. 5(1): 520 (1925); Briquet, Prodr.fl. corse 2(2): 144 (1936); Gorschkova in Shishkin & Bobrov, Fl. URSS 15: 241 (1949); Stjep.-Vesel. in Josifovic, Fl. Srbije 3: 1 16, t. 33 f. 2 (1972); nom. illegit. (Art. 52). Type as for H. quadrangulare L. [H. alatum Retz., Obs. hot. : 1 8 ( 1 8 1 0); Rchb. in Flora 5: 530 ( 1 822). Type: ex America septentrionale, Collector? (LU). Wahlenberg (Fl. suec.: 476 (1826)) makes this a synonym of his H. quadri- alatum along with '//. quadrangulare Sm.'and H. tetrapterum Fr.; but the original description (e.g. ... caule ... subtomentoso, calyc- ibus serrulato-glandulosis) clearly applies to quite another plant.] H. quadratum Stokes, Bot. Mat. Med. 4: 99 (1812), nom. illegit. (Art. 52). Type as for H. quadrangulum L. H. quadrangulum var. [8] sensu Choisy, Prodr. monogr. Hyperic.: 47 (1821). H. quadrangulum var. [8] confertum Choisy in DC., Prodr. 1: 548 ( 1 824). Type unlocalized, collector unrecorded (G-DC!-lectotype, selected here); England, 1816, Dawson Turner s.n. (G-DC!-syn- type). Although Choisy cites only 'Anglia' for the provenance of his var. Sin 1 82 1 , the Dawson Turner specimen is H. perforatum. The other specimen, however, is H. tetrapterum, and its dense inflorescence would appear to be the origin of the epithet confertum. SI H. quadrangulare var. [a] hum He Boenn., Prodr. fl. monast. westphal.: 221 (1824). Type: Germany, none stated. H. quadrangulare var. [[3] alatum (Retz.) Boenn., Prodr. fl. monast. westphal.: 227 (1824) pro parte excl. typum. See H. alatum Retz. above. H. quadrangulare var. [y] patulum Boenn., Prodr. fl. monast. westphal. : 227 ( 1 824). Type: Germany, none stated. H. quadrialatum Wahlenb., Fl. suec.: 276 (1826), nom. illegit. (Art. 52.1). Type as for H. tetrapterum. H. neapolitanum Ten., Ind. Sent. Hort. Neap. ann. 1829: 19 (1829); Guss., Fl. sic. syn. 2: 379 (1843); Lojacono-Pojero, Fl. sicula 1(1): 188 (1886^88). Type: Sicily, sine loc., in pratis paludosis, pre May 1830 (fl), Tenore s.n. (NAP-holotype; K!-isotype). Gussone gives a valid description but attributes the name to Tenore. I have not been able to check that Tenore himself pro- vided a valid description. H. confertum (Choisy) G. Don, Gen. Syst. 1: 606 (1831), non Moench (1794) nee Choisy (1821), nom. illegit. (Art. 52). Based on H. quadrangulum var. confertum Choisy. H. neapolitanum var. confertum (Choisy) Guss., Fl. sic. syn. 2: 379 (1843); Lojacono-Pojero, Fl. sicula 1: 188 (1886-1888). Type as for H. quadrangulum var. confertum Choisy. Holosepalum quadrangulum (L.) Fourr. in Ann Soc. Linn. Lyon, N.S. 16:352(1868). Hypericum rotundatum Schur in Verh. naturf. Vereins Briinn 15(2): 159 (1877). Type: Romania, Transylvania, 'Auf Moorwiesen, an Graben auf den Burgenwiesen bei Kronstadt [Bra§ov]', July 1854, Schur s.n. (FR!-holotype). H. borbasii Formanek in Verh. naturf. Vereins Briinn 29: 1 43 ( 1 89 1 ). Type: Turkey: Istanbul (Asia), Alemdag ('Alemda'), July-August 1890 (fl), Formanek s.n. (BRNMl-holotype). H. quadrangulum var. acutum (Moench) Fiori in Fiori & Paol., Fl. Italia 1: 387 (1898), Nuovofl. Italia 1: 521 (1924), comb, illegit. (Art. 58.2). H. quadrangulum subsp. quadrangulum sensu Tourlet in Bull. Soc. Bot. France 50: 307 (1903); Schinz in Vierteljahrsschr. Naturf. Ges. Zurich 49: 231-241 (1905), pro parte uterque quoad typum. H. tetrapterum forma stigmaphyllum Woron. in Kuzn., Busch & Fomin, Fl. Cauc. crit. 3(9): 43 (1906). Type: Russia, Ciscaucasia, Stavropol, Norman s.n. (LE). H. tetrapterum var. [b] neapolitanum (Ten.) N. Terrace, in Nuovo Giorn. Bot. Ital. 14: 135 (1907). H. acutum forma humile (Boenn.) Hegi, ///. Fl. Mitt.-Eur. 5(1): 521 (1925); Gu§ul. in Savul., Fl. R. P. Roman. 4: 26 (1956) ['Boenningh. ex Hegi']. H. acutum formapatulum (Boenn.) Hegi, ///. Fl. Mitt.-Eur. 5(1): 521 (1925). H. acutum subsp. eu-acutum Hayek, Prodr. Fl. Pen. bale. 1: 534 (1925), nom. illegit. (Arts 21.3, 52, 58.2). H. maculatum subsp. quadrangulum (L.) Hayek, Prodr. Fl. Pen. bale. 1: 534 (1925), pro parte quoad typum. H. quadrangulum var. tetrapterum (Fr.) Borg, Descr. Fl. Malt. isl. : 248(1927). H. acutum subsp. tetrapterum (Fr.) Maire in Jahandiez & Maire, Cat. pi. Maroc: 483 (1932), comb, illegit. (Art. 58.2). H. acutum var. typicum Gus,ul. in Savul., Fl. R. P. Roman. 4: 885 (1956). Type as for//, acutum Moench=//. quadrangulum L.=H. tetrapterum Fr. On p. 26 this combination is given as 'f. nova' , but on p. 885 (with other nomenclatural novelties) as 'var. nova'. The latter would seem to have been intended. H. quadrangulum subsp. acutum (Moench) Goday & Carbonell in Anales Inst. Bot. Cavanilles 19: 380 (1961), comb, illegit. (Art. 58.2). H. fallax subsp. fallax var. quadrangulare (L.) O. Schwarz in Drudea 5: 63 (1965) pro parte quoad typum. Icones: see under varieties. Perennial herb (0. 1 5-)0.2-0.7(-l .2) m tall, erect to ± decumbent or rarely ascending to procumbent from creeping and rooting base, with stems numerous to few, initially unbranched below inflores- cence, later branched above for half to most of their length. Stems ± broadly 4-winged, the subsidiary wings narrower or rarely with wings equally narrow, with black glands on the lines and sometimes elsewhere; internodes 7-50 mm, shorter than to exceeding leaves. Leaves sessile or rarely (especially the lower) up to 0.5 mm peti- olate; lamina (6-)9-40 x (4-)7-24 mm, ovate or elliptic or triangular-ovate to orbicular, somewhat paler beneath, thinly chartaceous; apex rounded to very rarely apiculate-obtuse, margin plane or rarely ± undulate, base rounded to cordate-amplexicaul; venation: 1-3 pairs of main laterals from base of midrib; tertiary reticulation dense, rather obscure; laminar glands pale, dense, small; intramarginal glands black, rather close, uniform. Inflorescence (1-) 10-35(-c. 70)-flowered, from l-2(-4) nodes, with flowering branches ascending from up to 10 nodes, the whole cylindric to corymbiform, dense or with partial inflorescences dense; pedicels 1-2 mm; bracts and bracteoles 2-3 mm long, lanceolate to linear, entire, with 0-4 black glands. Flowers 10-15 mm in diam., stellate; buds ellipsoid, subacute. Sepals 5, equal, 3-5 x 1-1.5 mm, lanceo- late to narrowly oblong, acute to acuminate, entire, erect in bud and fruit; veins 3 with outer pair branched or 5 unbranched; laminar glands striiform to punctiform or occasionally basal linear, all pale or 1-2 black; marginal glands absent or rarely up to 7, black, immersed. Petals 5, rather pale yellow, not tinged red, 5-8 x 2-3(- 4) mm, c. 1 .4 x sepals, oblanceolate, entire or distally unilaterally crenate; laminar glands absent or rarely few, punctiform, black and pale; marginal glands 1-4 or usually absent. Stamens 30-40(-60), '3'-fascicled, longest c. 4.5-7.5 mm, c. 0.9 x petals. Ovary 3- locular, 2-2.5 x 1-1 .5 mm, narrowly ovoid; styles 3, free, 2-3.5 mm, 0.8-1 .4 x ovary, widely spreading. Capsule 5-7 (-8) x 2.4-4 mm, c. 2 x sepals, narrowly cylindrical-ellipsoid to ovoid; valves with longitudinal linear vittae. Seeds dark brown, 0.6-0.8(-1.1) mm, cylindric, not carinate or appendiculate; testa finely foveolate. 2n= 1 6 (Stenar, 1938; Noack, 1939; Polya, 1950; Robson, 1956; Gadella & Kliphuis, 1966; Zhukova, 1967), n=8 (Nielsen, 1924; Winge, 1925; Chattaway, 1926; Noack, 1939; Sugiura, 1944). In wet habitats - marshes, streamsides, open ditches, wet meadows, springs; 0-1300 m (Switzerland, Valais), -1900 m (Corsica), -2000 m (Turkey). Europe from Scotland (Orkney and Sutherland southward), Den- mark, S. Sweden (Skane), Poland, the SW Ukraine and Transcaucasia south to Spain, Algeria, Sicily, Greece, Turkey, N. Israel, N. Iraq and NW Iran. The variation in H. tetrapterum is wide but almost continuous, from a tall, branching, relatively narrow-leaved form in southern Italy and Sicily ('//. neapolitanum') to short unbranched forms in northern Europe. In two areas, however, decumbent small-leaved few-flowered high-altitude forms have developed - Corsica and the Levant. Although in neither area have field studies been made to decide how distinct these forms are from the adjacent typical form, they both seem worth recognizing at least until such studies are made. Hvpericum tetrapterum is related to 2b. H. undulatum var. undulatum, differing from it in the more broadly winged stem internodes, the usually relatively broader leaves with plane margin. 82 N.K.B. ROBSON and the denser inflorescence with smaller flowers, sepals with fewer black laminar glands, and paler petals that are not tinged red. The form most similar to var. undulatum occurs in southern Italy and Sicily ('//. neopolitanum Ten.'). 3a. Hypericum tetrapterum var. tetrapterum H. tetrapterum subsp. tetrapterum, Zangheri, Fl. Ital.l: 396 (1976). Icones: Rchb., Ic. fl. germ. helv. 6: t. 344, f. 5179 (1844); Ross- Craig, Draw. Br. PL 6: t. 9 (1952); Ramos in Castroviejo et al., Fl. Iberica 3: t. 46(1989). Stems erect, to 1.2 m tall. Leaves sessile or lowermost to 0.5 mm petiolate, 9^0 mm long, ovate or elliptic to orbicular, margin nearly always plane. Inflorescence 10-c. 70-flowered, from 1—4 nodes, with flowering branches from up to 10 nodes below. Lowland to montane zones; 0-1380 m. Distribution as for species, except for the alpine zone ( 1 380-2000 m). SCOTLAND. Orkney: Hoy, North Walls, Lyness, 16 July 1956 (fl), Sinclair 8872 (K). East Ross: Dingwall, 25 July 1961 (fl), McCallum Webster 5788 (K). Moray: near Elgin, Lochinver Farm, 1 3 August 1960 (fr), McCallum Wester 3833 (K). Kincardine: St. Cyrus, 6 August 1881, Nicholson s.n. (JE). Mid Perth: Crieff, near Comrie, July 1880 (fl) Bennett s.n. (BM). W. Lothian: near Kirkliston, 1 August 1905 (fl), Bennett s.n. (BM). E. Lothian: Aberlady, 15 August 1933 (fl & fr), Taylor s.n. (BM). Lanark: Carluke, meeting of Cauldron Gill and Yieldshiel Burn, 11-15 July 1962 (fl), Phillips 176 (BM). Wigtown: Wigton, July 1895 (fl), McAndrew s.n. (BM). Clyde Isles: Arran, Whiting Bay, Glenashdale, 26 July 1953 (fl), Robson 24 (BM). Argyll: near Crarae, 20 July 1936 (fl), A.B. Jackson s.n. (BM). S. Ebudes: Islay, Bowmore to Laggan Bridge, 28 August 1955 (e. fr), Hillcoat s.n. (BM). ENGLAND. Westmorland: Arnside, 5 August 1896 (fr), Worsdell s.n. (K). Durham: Wynch Bridge, 18 June 1934 (fl), Taylor s.n. (BM). NE Yorks.: near Scarborough, Trontedale, 20 July 1939 (fl), Evans s.n. (BM). S. Lanes.: Freshfield Dunes, 26 July 1929 (fl), Holder 461 (LIV). Notts.: E. of Blaxton on B 1396, 25 July 1963 (fl), Bowden & Hillman 359 (BM). S. Lines.: near Fenhouses, 31 July 1945 (fl), N.D. Simpson s.n. (BM). Worcs.: Wyre Forest, 11 August 1915 (fl), Rea s.n. (BM). E. Gloucs.: Shuthanger Common, 22 September 1973 (fl), Fowler s.n. (BM). Huntingdon: Eynesbury, August 1944 (fl), Peck s.n. (BM). W. Suffolk: Mildenhall, July 1908 (fl), Adamson s.n. (BM). Oxford: King's Sutton, 1 August 1961 (fl), Sheasby60\ (BM). Middlesex: Finchley Common, 12 September 1925 (fr), Moring 282 (BM). Surrey: Horley, Langshott Wood, July 1948? (fl), Morgan s.n. (BM). E. Sussex: Pett, 12 September 1938 (fl & fr), Bangerter s.n. (BM). Isle of Wight: Freshwater, 1 August 1953 (fl), Hancock s.n. (BM). S. Wilts.: Donhead St. Mary, Alec's Shade, 4 August 1947 (fl), Wilmott 1 102 (BM). N. Devon: Braunton Burrows, 15 July 1957 (fl), Matthews s.n. (BM). W. Cornwall: St. Ives, 17 June 1952 (fl), Edwards 57 (BM). WALES. Glamorgan: Kenfig Dunes, 1 5 July 1 934 (fl), N.D. Simpson s.n. (BM). Carmarthen: Pembrey, September 1900 (fl), Riddelsdell s.n. (BM). Pembroke: Sagerton Common, 18 August 1957 (fr), Powers 5 (BM). Radnor: near Rhos-goch, 18 August 1885 (fl), A. Ley s.n. (BM). Caernar- von: Bardsey I., August 1933? (fl), Butler (BM). Flint: Bodlewyddan Park, 12 August 1941 (fl), A.B. Jackson s.n. (BM). IRELAND. N. Kerry: Lough Leane, E. side of Ross I., c. 20 m, 20 July 1973 Jermy 10391 (BM). W. Cork: near Toormore Mizen Head, near Ballyrigode House, 1 July 1961 (fl), James 60 (BM). Waterford: c. 2.4 km S. of Tramore, near road to Great Newton Head, 18 August 1967 (fl), Ferguson 2039 (BM). Dublin: How the Hill, August 1877 (e. fr), Gamble 26303 (K). Longford: 8 km SW of Granard, 17 August 1961 (fl), Summerhayes 3599 (K). S. Donegal: Ballyshannon, 1992 (fl & fr), Wyse Jackson s.n. (Herb. M.W.J.). CHANNEL ISLANDS. Guernsey: Petit Bot Bay, 22 July 1950 (fl), F. Balfour-Browne s.n. (BM). Sark: near harbour, 10 July 1953, Sowerby 134 (BM). Herm: N. of island, 13 August 1952 (fl), Hancock s.n. (BM). Alder- ney: Clanque Bay, 16 August 1932 (fl), A.B. Jackson 579 (K). Jersey: no loc., 21 July 1900 (fl), Lester-Garland s.n. (K). Map 4 3. H. tetrapterum. a. var. tetrapterum •, other records O; b. var. corsicum A ; c. var. anagallidifolium • (among the Lebanon records). STUDIES IN HYPER1CUM 83 THE NETHERLANDS. Zuid-Holland: Rozenburg I., 'De Beer', 26 August 1 948 (e. fr), Maas Geesteranus 432 1 (K, L*). Utrecht: N. of Utrecht, Molenpolder, 19 August 1953 (fr), Leeuwenberg s.n. (H). BELGIUM. Antwerpen: entre Lierre [Lier] et Emblehem, 25 July 1920 (eAr), Lambert s.n. (BM).Namur: Wavreville, June \9\4(f\),Boulengers.n. (EM). FRANCE Oise: pres Senlis, Foret de Hallare, 18 August 1834 (fl), Bentham s.n. (K). Seine: bois de Meudon, 26 August 1889 (fl & fr), Gadeceau s.n. (BM). Finisterre: Morlaix, August 1883 (fl), Miciol s.n. (BM). Loire-Atlantique: ruisseau d'Escoublac, 1887 (fr), Gadeceau s.n. (BM). Deux-Sevres: Saint-Genard, pres de Chancelles, August 1951 (fl), Contre in Soc. Fr. Exch. PI. Vase. B. de Retz 5 (K). Cher: Saulzait-le-Potier, 22 July 1859 (e. fr), Deseglise s.n. (BM). Haut-Rhin: Pfetterhausen, 9 August 1 869 (fl & fr), Gadeceau 217ter (BM). Doubs: Besan^on, marais de la Malcombe, n.d. (fl), Grenier 2036bis (BM, JE). Ain: Cascade de Glossdieu, 9 August 1891 (fl). Borel s.n. (K). Rhone: ad Ainas, 6 August 1874 (e. fr), Gandoger401 (BM, K). Savoie: Saint Bezon, 7 August 1892 (fl), Borel s.n. (K). Puy-de-D6me: Puy St Gulunier?, 7 August 1928 (fl), Elussl 355 (BM). Cantal: Ydes, 8 August 1874 (fr), Heribaud- Joseph s.n. (BM). Aveyron: Saint-Paul-des-Forets, August 1904 (fr), Coste s.n. (BM). Basses-Pyrenees: Harambetz, vid Chapelle St Nicolas, 15 August 1968 (fl), Sonck s.n. (H). SPAIN. Guipuzcoa: Vascongadas, San Sebastian, M. Castelli, 80 m, September 1940, Rothmaler 16415 (JE). Huesca: Castejon de Sos, valley of Rio Esera, c. 950 m, 27 July 1955 (fl), Sandwith s.n. (K). Barcelona: Montserrat, 24 August 1941 (fl & e. fr), Collector! (RNG). Zaragoza: Catalayud, 17 July 1912 (fl), Vicioso s.n. (BM). Avila: Sierra de Credos, Rio Alberche S. of St. Martin del Pimpollar, c. 1450 m, 23 July 1974 (fl), Brenan 13036 (K). Castellon: Segorbe, 18 July 1922 (fl), Pau in Sennen PI. Esp. 4620 (BM, RNG). Almeria: Laujarde Andarax, 8 August 1978 (fl), Roivainen s.n. (H, RNG). ALGERIA. Atlas Tellien: Medea, gorges de la Chiffa ((fide Quezel & Santa, 1963: 682); pres Blidah [Blida], Ravin de 1' Oued-el-Kebir, 13 July 1854 (fl), Cosson s.n. (G-BOISS). Petite Kabylie: Djudjura (fide Quezel & Santa, 1963: 682). CORSICA. Stagno di Palo, 20 July 1933 (fl), Aellen 1655 (K). SARDINIA. Tempio, 7 August 1882, Reverchon s.n. (JE). SICILY. Messina: Messina, Haeckel s.n. (JE). Collector! : [Boscovi Penzzubaltone della] Castagnera, July 1879 (fr), Tacono s.n. (FR). Palermo: Palermo, June 1900 (fl & fr), Ross s.n. (BM). ITALY. Venezia Giulia: bei Trieste, Monfalcone, September 1921 (fr), Meebold s.n. (K). Trentino-Alto Adige: Bozen [Bolzano], pre 1877 (fl), Hausmann 1377 (BM). Piemonte: Casale Masferrato, 15 July 1957 (fl), Vila & Cedercreutz s.n. (H). Massa-Carrara: Massa, August 1952, Pellegrini s.n. (H). San Marino: Carrepa, 7 September 1920 (e. fr), Pampanini s.n. (K). Campania: between Majori [Maiori] and Capo d'Orso, 29 June 1883 (fl), Lacaita 6400 (BM). SWITZERLAND. Geneve: Geneva, July 1859, Huet de Pavilion s.n. (JE). Vaud: Nyon ad pagum Trilia, 9 August 1 876 (fl), Elfving s.n. (H). Bern: Radelfingen, 1 August 1879 (fl), de Rutte s.n. (K). Luzern: Lucerne, Sonnenberg, 24 July 1 930 (fl), Rathbone s.n. (BM). Appenzell: am Ga'bris ob Gais, July-August, Schneider s.n. (BM). AUSTRIA. Tirol: Oberinntal, Vols bei Innsbruck, c. 600 m, 8 August 1980 (fl), Polatschek s.n. (BM). Salzburg: near Salzburg, Gaisberg, n.d. (e. fr), Hinterhuber s.n. (K). Karnten: Klagenfurt, 1864 (fl), Kokeil s.n. (K). Steiermark: Lemsitz, Rosegg Ettendorf bei Stainz, 1 4 August 1 924 (fl & fr), Trover s.n. (H). Oberosterreich: bei Aishersheim, August 1861, Keck s.n. (JE). Niederosterreich: Modling, Vorderbriihl, August 1919 (fr), Fenz! 82 (K). Burgenland: Podersdorf, Zitzmandorfer Wiesen, 1-9 August 1966 (fl), Excurs. Bot. Inst. Ffin. s.n. (FR). GERMANY. Baden: Baierthab, 18 August 1883 (fl), Druce s.n. (BM). Wurttemberg: Oberamt Reutlingen, Kirchentellinsfurt, 31 July 1991?, Lindberg s.n. (H). Bayern: Oberbayem, Haspelmoor, 17 July 1918? (fl), Oberneder 3261 (BM). Hessen: Bickenbach, bei Darmstadt, 10 September 1961 (fl & fr), Schaarschmidt96 (FR). Thiiringen: Souderbei Schlobheim, 25 July 1952 (fl), Launerts.n. (BM). Brandenburg: Lychen, bei Kustmichener Bach, 20 July 1900 (fl), Heiland s.n. (H). Schleswig-Holstein: Kaltenhofer, 25 July 1893 (fl), Simmons s.n. (H). DENMARK. Jutland: between Lonstrup and Hirtshals, 17July 1967(fl), Holm-Nielsen & Jeppesen in Fl. Jutl. Exs. 347 (BM, H, TAI); Fiinen: Potenden to Sonderso, 6 August 1883 (fl), Mortensen s.n. (H). Falster: Hannenov Skov, D. 37, 17 July 1942 (fl), Dahl s.n. (BM). Zealand: W. of Vrangeskov, N. of Ringsted, 24 July 1969 (fl), Jacobsen & Svendsen in PI. Dan. Exs. 93 (H). SWEDEN. Skane: Benstadt, 2 October 1899 (fr), Sjoshol s.n. (H); Par. Everod, Lyngby, 31 August 1933 (fl), Hasslow s.n. (BM). POLAND. Siedlce: Sedletz, ad pag. Jagodne prope urbem Lukow, 28 July 1902 (fl), Puring 1971 (H). Wroclaw: Stawno apud Milicz, ad ripam stagni 'Golica', 5 August 1963 (fl), Kotowicz & Koziol 857 (H). Bielsko-Biala: Wadowice, prope Andrychow, 6 August 1938 (fl), tancucka PI. Polon. Exs. 330(BM,JE). UKRAINE. West and south-west regions (fide Karnaukh, 1955: 307); Krym: Oreanda, 29 July 1987 (fl), Golde s.n. (LE). CZECH REPUBLIC. Moravia: bei Schnobolin n. Olmiitz, June 1908 (fl), Laus s.n. (BM, JE, K); Brno, Bilovice, ad Ricmanice, c. 230 m, 27 July 1925 (1. fl), Mrkos 243a (H, K). SLOVAKIA. Throughout (see Zeleny et al., 1982: map 57). ROMANIA. Transylvania: Cluj, ad pagum Feleac super oppidum Cluj, c. 650 m, 28 July 1923 (fl & fr), Gurtler & Nydrddy 1521b (H); Oreada, inter oppida Bucea et Grossi, 900 m, 30 July 1971 (fl), Cernoch 22232 (BM, H). Walachia: Muntenia, distr. Prahova, Prahova rly stn, 5 August 1923 (fl & fr), Grintescu 1 521 a (H). Also in Moldavia (/We Gus.uleac & Nyarady, 1956:26). BULGARIA. Sofia: prope Sofia, ad radicem m. Vitosa [Cherni Vrtikh], 5 August 1939 (fl), Lindberg s.n. (H). SLOVENIA. Bled [Veldes], 1924 (fl), Leathes s.n. (BM); Carniolia, ad pagum St. Margarethen prope Rudolfswert, 200 m, August (1. fl), Paulin, Fl. exs. Carniol. 326 (BM). SERBIA, bei Bor, June 1917, Koppe s.n. (JE). BOSNIA. Sarajevsko Polji, prope Ilidza, c. 500 m, 5 August 1902 (fr), Malz s.n. (K). MAKHEDONUA. *L. Ochrid [Ohrid], W. of Ochrid, 25 July 1976, Akeroyd, Mellors & Preston (CGE). ALBANIA. North: Tepalens [Toplane?] bei Luzat, c. 250 m, 9 September 1964 (fr), Meyer 6029 (JE); Dardha, Weg zum Qani i Dardhes, c. 800 m, 1 August 1959 (fl), Meyer 4569 (JE). South: Berat, c. 60 m, 16 August 1935 (fr), Alston & Sandwith 2514 (BM, K). GREECE. Thrace: Emirli, 100 m, 24 June 1931 (fl), Tedd 742 (K). Makedonia: Dramas, vill. Katafyton (SE foot of Mt. Orvilos), road to Ano Vrondou, 740 m, 23 July 1977 (fl & fr), Strid & Georgiadou 13052 (C); Grevenon, Mt. Smolikas, 2-3 km S. of Samarina towards Armata, 1275- 1350 m, 27 August 1975 (1. fl), Hartvig & Seberg 4816 (C). Ipiros: Pindus, Smokovo to Rentino, 4 August 1937 (fl), Balls & Gourlay B. 37 1 1 (BM, K). Corfu: Limni Korisson, near Linia, 17 July 1972 (fl), Davis 54547 (BM). Thessalia: Mt. Olympos, S. side, Leptokarya to Karya, 22.8 km, 770 m, 27 August 1975, Strid & Hansen 9555 (C). Sterea Ellas: Attica, prope Kephissiam [Kifisia], ad radices mentis Pentelici, 195-300 m, August (1. fl). Heldreich s.n. (BM). Akhaia: Ep. Kalavryton, Mt. Chelmos (east), c. 2 km SSW of Zarouchla, 1400-1450 m, 22 August 1982 (fl), Hartvig, Franzen & Cristensen 10296 (C). Argolos: prope Lerni, 13 July 1930 (fl), Guiol 1518 (BM). Arkhadia: Kinouria/Lakedhemonia, Mt. Parnon, Ajios Petros, Vitilithra, 1300 m, 13 July 1971 (fl), Samatiadou 13225 (BM). Evvoia: Mt. Dirphys, 24 July 1932 (1. fl), Guiol s.n. (BM). Karpathos: 0.25 km SW of Avlona, 300 m, 8 April 1992 (st), Chilian & Turland 285 (BM). Lesvos: Agios Anagari, c. 3 km S. of Sikounta, 100-150 m, 9 September 1992 (e. fr), Hansen & Nielsen 8030 (C). TURKEY. Istanbul (Europe): Belgrad Forest, 3 1 July 1959 (fl), Yallirik ISTO 135 1 (E). Istanbul (Asia): Aydos, 20 September 1950 (fr), Baylop (& Berk) 3603 (BM, ISTE*). Kocaeli: Izmit, Izmit to Sapanca, c. 20 km, Masukiye, 20 July 1981 (fl), Kurtto 3088 (H). Kastamonu: N. side of Ilgaz Dag, 1500 m, 28 July 1962 (fl), Davis, Coode & Yaltirik D. 38288 (E, K). Giresun: above Giresun, 900 m, 24 August 1962 (fl), Furse 4137 (K). Trabzon: Of, s.l., 5 August 1957 (fl & fr), Davis & Hedge D.20823 (BM, E, K). Coruh: Chope [Hopa], 6 August 1917 (fr), Shishikin s.n. (LE). Manisa: Manisa, 3 September 1932 (fl), Kotte 319 (K). Kutahya: Gediz, §aphane, 1 200 m, 27 August 1 950 (fl), Davis 1 8505 (E). Afyon: Afyon to Sandikli, 3 km from Sinkanli [Sinanpa§a], 28 August 1 97 1 (fl), Fraser Jenkins 2 1 64 (BM, E). Sivas: 1 1 km N. of Sivas, 1300 m, 10 September 1977 (fr), Sorger 77-109- 8 (Herb. Sorger). Antalya: SW of Elmali, W. of Gombe, 1200 m, 10 July 1968 (fl & e. fr), So/gev68-25a-2 (BM); Gebiz, Bosburun Dag at Panargazu yayla, 600 m, 23 July 1949 (fl), Davis 15492 (K). Hatuy: Amanos daglari, Dortyol Pinarbasi, 22 September 1967 (e. fr), Akman 136 (ANK, BM). 84 SYRIA. Damascus, n.d. (fr), Gaillardot 910 (JE); Mt. Hermon, in valle Orny 1500 m, 2 July 1855 (fl). Kotschy 218 (BM, K, UPS). LEBANON. Coast: Saida, au dessus du moulin de 1'Eniir, 4 August 1854 (fl), Gaillardot 911 (G, JE). Lower mountains: Becharre [Bcharre], 18 August 1945 (fr). Davis 10087 (K). Al Biqa: Bekaa, Malaka, July 1853 (fl), Gaillardot s.n. (JE). IRAQ.Amadiyah:Sersang, 1050m, 14 July \955 (fl), Haines W.534(K). Rawanduz: Pushtashan. 1 5 km NE of Rania, Qandil Range, 1 1 50 m, 29 July 1957 (1. fl), Rawi NHI 23897 (K). Sulaymaniya: in Mt. Avroman humid., 1200-1800 m, July 1867 (fr), Haussknecht s.n. (JE). IRAN. Gilan: Deylaman to Lanak, 600-1600 m, 9 July 1972 (fl & fr), Tenne 34320 E (BM, EVIN). Mazanderan: Astara to Bandar-Pehlevi, 15 km, 1 8 August 1 963 (fl & fr), Jardine 92 1 (E, W). Arak: in monte Raswend, August 1989 (fl), Strauss s.n. (JE). Gorgan: Ketul, June 1948 (fl), Sharif in Herb. Min. Agr. 359 (W). AZERBAIJAN. Lenkoranskiy rayon, Port Il'ish, 17 July 1963 (fl), Bobrov & Tzvelev 284 (LE). ARMENIA. South-east: recorded from Dzhermuk, Shake and Goris (fide Takhtajan, 1966: 13). GEORGIA. Batumi, Strandpromenade, 5 August 1977 (fl), Marstaller s.n. (JE): Sukhumi. E. of the town, 27 August 1957 (fr), Meyer, Lippold & Kohler 835 (JE). Gudauta. NW bei Xirnema, 13 July 1969 (fl), Lepper, FwcAeretal.5299(BM,JE). RUSSIA. Stavropol: Nevinnomyssk, bei der Kuban-Briicke, 18 August 1957 (fr), Meyer. Lippold & Kohler 18 (JE). 3b. Hypericum tetrapterum [var.] ycorsicum (Steud.) Boiss., Fl. orient. 1: 816 (1867) pro parte quoad typum. Type as for H. corsicum Steud. Map 4. H. tenellum Tausch in Flora 14: 2 1 1 ( 1 83 1 ); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120: 584 (191 1), non Solander ex Clarke (1812). Type: Corsica, in montibus Corsicae, Tausch s.n. (PRC-holotype). H. corsicum Steud., Nomencl. 2nd ed., 1: 787 ( 1 840); Godr. in Gren. & Godr., Fl. France 1: 315 (1847); Bonnet in Bull. Soc. Bot. France 25: 280 (1878); Rouy, ///. pi. Eur., fasc. 5: 35, t. 106 ( 1 896). Type as for H. tenellum Tausch. H. repens Duby ex Nyman, Consp.fl. eur. 1: 133 (1878), nomen. H. tetrapterum var. [[3] tenellum (Tausch) Parlat, Fl. ital. 5: 518 (1875); Cesati, Passerini & Gibellini, Comp.fl. ital: 758 (1884). H. acutum subsp. corsicum (Steud.) Rouy in Rouy & Fouc., Fl. France 3: 338 (1896); Hayek, Prodr. Fl. Pen. bale. 1: 534 (1925) pro parte quoad typum; Briquet, Prodr.fl. Corse2(2): 145(1936); comb, illegit. (Art. 58.2). H. quadrangulum subsp. corsicum (Steud.) Fiori in Fiori & Paol., Fl. Italia 1:387 (1898). H. insulare Foucaud & Mandon in Bull. Soc. Bot. France 47: 89, t. 2 (1900). Types: Corsica, entre le col de Sorba et Ghisoni, Mandon & Foucaud s.n. (?); Cascade de la fontaine de Puzzichello, pres de Ghisoni, Mandon, Rotges & Foucaud s.n. (?). H. acutum var. corsicum (Steud.) Halacsy, Consp.fl. graec. 1: 281 (1900), comb, illegit. (Art. 58.2). H. acutum var. insulare (Foucaud & Mandon) Briq. in Annuaire Conserv. Jard. Bot. Geneve 5: 95 (1901), comb, illegit. (Art. 58.2). H. acutum subsp. corsicum var. rotundifolium sensu Briq., Prodr.fl. Corse 2(2): 143 (1936), non H. acutum var. rotundifolium (Willk.) Schinz (= H. undulatum var. boeticum). H. tetrapterum subsp. corsicum (Steud.) Zangh., Fl. Ital., 1: 196 (1976). Icones: Rouy, ///. pi. Eur., fasc. 5: t. 106 ( 1 896); Foucaud & Mandon in Bull. Soc. Bot. France 47: t. 2 (1900). Stems procumbent to ascending, up to 0.2 m long. Leaves mostly 0.5 N.K.B. ROBSON mm petiolate, 6-12 mm long, elliptic to orbicular, margin plane. Inflorescence l(2-3)-flowered. Montane to alpine zones; 1000-2000 m. Corsica. The Cretan record (Bonnet, 1879) is an error (see Greuter, 1974: 156). Bonnet's record from Spain (loc. cit.) would appear to refer to small plants of 2b. H. undulatum var. boeticum. CORSICA. Monte d'Oro, 2000 m, 6 August 1 826? (fl), Soleirol 1 1 2 (BM, K, WAG); Licano, 12 July 1912 (fl), Lacaita s.n. (BM); Bastelica, Monte Renoso, 25 August 1878 (fl & fr), Reverchon s.n. (BM). 3c. Hypericum tetrapterum var. [[}] anagallidifolium Boiss., Fl. orient. 1: 806 ( 1 867); Rech.f. in Ark. Bot. 5: 294 ( 1 960). Type as for H. anagallidioides Jaub. & Spach. Map 4. H. anagallidioides Jaub. & Spach, ///. pi. orient. 1: 47, t. 24 (1842), non H. anagalloides Cham. & Schltdl. (1828). Type: [Turkey (Amanus)], in oriente, Labillardiere s.n. (Fl-holotype, photo- graph!). H. anagallidifolium Boiss., Fl. orient. 1: 806 (1867), in synon. H. acutum var. anagallidifolium (Boiss.) Post, Fl. Syria, 2nd ed. 1: 233 (1932), comb, illegit. (Art. 58.2). H. acutum var. anagallidioides (Jaub. & Spach) Zohary, Fl. Palaestina 1: 223 (1966), comb, illegit. (Art. 58.2). Icon: Jaub. & Spach, ///. pi. orient. 1: t. 24 (1842). Stems ascending, slender, c. 0. 1 5 m long (habit ofAnagallis arvensis L.). Leaves subsessile to 0.5 mm petiolate, c. 7-12 mm long, elliptic to triangular-ovate, margin undulate. Inflorescence 1-3-flowered, from one node, sometimes with flowering branches from up to 5 nodes below, then totalling 20-25 flowers. Stream banks; 1300-2000 m. Turkey (Amanus), Lebanon, Israel. TURKEY. Hatay: Ziaretdag, 18 September 1884 (1. fl), Post s.n. (BM); Monts Amanus, 1200-1800 m, September 1913 (fl), Haradjian 4635 (E, G). LEBANON. Lebanon: montagne entre Saida et Deir Mekhalta pres du Djoume, n.d. (fl), Gaillardot s.n. (JE); Tripoli, Vallee de Hasroun, 7 July 1 866 (fl), Blanche s.n. (JE). ISRAEL. Upper Galilee (fide Zohary, 1966: 223). 4. Hypericum triquetrifolium Turra, Farsetia: 12 (1765); Post, Fl. Syria, 2nd ed. 1: 233 (1932); Briquet, Prodr.fl. corse 2(2): 148 (1936); Blakelock in Kew Bull. 3: 404 (1948); Quezel & Santa, Nouv. Fl. Algerie: 682 (1963); Zohary, Fl. Palaestina 1: 233, t. 328 (1966); N. Robson in Davis, Fl. Turkey 2: 400 (1967), in Tutin et al., Fl. Europ. 2: 269 (1968), in Meikle, Fl. Cyprus 1: 300, t. 16 (1977), in Townsend & Guest, Fl. Iraq 4: 379 (1980); Mouterde, Nouv. Fl. Liban Syrie 2: 527, t. 228 ( 1 970); Tackholm, Students ' Fl. Egypt, 2nd ed. : 1 50 ( 1 974); Ali & Jafri, Fl. Libya 2: 2, f. 1 (1976); Pott.-Alap., Fl. Tunisie: 508 (1979); Hagemann in Bot. Jahrb. Syst. 102: 247, f. 9 (1981), Hagemann & Meusel in Flora 175: 385^05, ff. 1A, 2-8, 9B, 9C, 10A, 11 (1984), Hagemann in op. cit. 183: 284, f. 57 (1989); Demiri, Fl. Ekskurs. Shquiperise: 314 (1981?); Pignatti, Fl. Italia 1: 350 (1982); Ramos in Acta Bot. Malac. 11: 167, f. 6b (1986), in Castroviejo et al., Fl. Iberica 3: 167 (1993); Boulos, Fl. Egypt2: 156 (1999). Type: 'Habitat in Grecia, Sicilia, Calabria'. Turra's herbarium was in the Museo civico, Vicenza, but was destroyed during the second World War (Stafleu & Cowan, 1986). Fig. 7, Map 5a, b. H. crispum L., Mant. pi: 106 (1767); Choisy, Prodr. monogr.fam. Hyperic.: 49 (1821), in DC., Prodr. 1: 549 (1824); Sibth. & Sm., STUDIES IN HYPERICUM 85 O Map 5a 4. H. triquetrifolium, west area • , other records O . Map 5b 4. H. triquetrifolium, east area •. 86 N.K.B. ROBSON Fl Graeca 8: 55, t. 776 (1833); Spach, Hist. nat. veg., Phan. 5: 385 (1836), in Ann. Sci. nat., Bot. 5: 357 (1836); Unger & Kotschy, Ins. Cypern: 353 (1865); Boiss., Fl. orient. 1: 806 (1867); Rouy in Rouy & Fouc., Fl. France 3: 350 (1896); Haldcsy, Consp. fl. graecae 1: 283 (1900); Woron. in Kuzn., Busch & Fomin, Fl. Cauc. Crit. 3(9): 45 (1906); Hand.-Mazz. in Ann. Naturhist. Mus. Wien 27: 60 (1913); Holmboe, Veg. Cypr.: 127 (1914); R. Keller in Engl. & Prantl, Nat. Pflanzenfam., 2nd ed. 21: 179 (1925); Nabelek in Spisy Pri. Fak. Sci. Masarykova Univ. 35: 53 (1932); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: t. 4 f. 28 (1932), 11: 176 (1933), 12: 88 (1934), in Pflanzenareale 4: karte 6a (1933); Rech. f., Fl. Aegaea: 263 (1943); H. Lindb. in Acta Soc. Sci. Fenn., Ser. B. 2 (no.7): 23 (1946); Gorschkova in Shishkin & Bobrov, Fl. URSS 15: 244 (1949); Tackholm, Students' Fl. Egypt: 223 (1956); Edgecombe, Weeds of Lebanon, 3rd ed.: 252 + t. (1970); nom. illegit. Type as for H. triquetrifolium Turra. Icones: Zohary, Flora Palaestina 1: t. 328 (1966); N. Robson in Meikle, Fl. Cyprus 1: t. 16 (1977); Cadete in Valdes, Talavera & Galiano, Fl Andalucia Occid. 1: 315 (1987). Perennial herb 0. 15-0.55(-0.77) m tall, erect to decumbent, rooting at the base, the roots bearing shoots, with stems solitary or few, widely branched along most of length, usually forming pyramid. Stems 2-lined, with black glands on the lines and sometimes else- where; internodes 10-30 mm, longer than leaves. Leaves sessile; lamina 3-20 x 1-9 mm, triangular-lanceolate or rarely narrowly ovate to linear-oblong, concolorous, sometimes glaucous, chartaceous; apex acute, margin crisped-undulate, base cordate- amplexicaul; venation: 2-3(^1) pairs of main laterals from base or near base of midrib, tertiary reticulation usually obsolete; laminar glands pale, ± dense, very small; intramarginal glands black, spaced. Inflorescence l-7(-ll)-flowered, from 1-2 nodes, with flowering branches ± horizontally spreading from all other nodes, sometimes excepting 1-6 immediately beneath apical one, the whole broadly pyramidal; pedicels 1-5 mm; bracts and bracteoles 1.5-2.5 mm long, lanceolate, entire or rarely apically denticulate. Flowers 8- 12(-14) mm in diam., stellate; buds broadly ellipsoid to globose, rounded. Sepals 5, subequal to unequal, l-2.2(-3) x 0.5-1 mm, oblong to ovate-oblong or lanceolate, acute to rounded-apiculate or rounded, entire or eroded-denticulate, erect in bud and fruit; veins 3, unbranched; laminar glands pale, 2, linear, distally interrupted; intramarginal glands absent. Petals 5, bright? yellow, not red-tinged dorsally, 4.5-7 x 1-1.5 mm, c. 3.5-5 x sepals, linear-oblanceolate, entire; laminar glands pale, striiform and punctiform, and occasion- ally black, solitary; marginal glands absent or rarely black, few. Stamens 15^0, '3'-fascicled, longest 2.5-5 mm, c. 0.6-0.8 x petals; anther gland black. Ovary 3-locular, 1 .5-2 x 0.5-1 mm, ellipsoid to broadly ovoid; styles 3, free, 2.5-3 mm, 1.7-3 x ovary, spreading; stigmas narrow. Capsule 3-5 x 2-3.5 mm, 2.5-3 x sepals, ovoid; valves with longitudinal linear vittae and occasionally a few lateral vesicles. Seeds darkish brown, 1.5-1.8 mm, subcylindric, not cari- nate or appendiculate; testa finely foveolate to scalariform-foveolate. 2n=16(Reynaud, 1973, 1975). In dry open stony ground and cultivated fields; 0-1350 m (Israel), - c. 2 100m (Iraq). S. Spain (Malaga, Andalucia), Gibraltar, Balearic Is., S. France, S. Italy, Sicily, Malta, Tunisia, Algeria, Libya (Cyrenaica), Montenegro, Albania, Greece (mainland and islands), Crete, Cyprus, Turkey, Levant south to Egypt (Sinai) and Jordan, Iraq, NW Iran. Probably not native west of Sicily and Cyrenaica or in Montenegro. SPAIN. Malaga: Malaga, cerca del aerodrome, May 1965 (fl), Borja s.n. (BM); San Julian, 1 8 May 1 952 (fl), Roivainen s.n. (H). Andalucia: Andevalo, fide Ramos (1987). GIBRALTAR. Near First River Ferry, 26 June 1912 (fl), Wolley-Dod 1478 (BM). BALEARIC IS. Minorca: Alayor, naturalized (fide Knoche, 1922: 188). FRANCE. Bouches du Rhone: Les Camoins, between Neoules and Marseilles, 10 July 1968 (fl), Blaize s.n. (H); Aix-en-Provence, July 1891, Bruyas s.n. (JE). Herault: Montpellier, Port Juvenal, July 1 932 (fl), Auersmitl s.n. (BM). ITALY. Puglia: Apulia, circa Manfredonia, 3-15 m, 28 June 1875 (fl), Porta & Rigo 241 (BM, FR, JE, K); Taranto, Leucaspide, 30 June 1908 (fl), Lacaita 291/08 (BM). Calabria: Reggio, Pellaro, 13 July 1898 (fl & e. fr), Rigo 472 (H). Campania: Naples, n.d. (fl), Lindley s.n. (K). SICILY. Messina: 10 km NE of Taormina, Capo San Alessio, 100 m, 17 July 1 983 (fl), Akeroyd, Jury & Rumsey 3572 (RNG). Catania: Catania, June 1885, Ross s.n. (JE). Siracusa: Siracusa, Roman amphitheatre, 100 m, 17 July 1983 (fl & fr), Akeroyd, Jury & Rumsey 3564 (RNG). Caltanisetta: Caltanisetta, August 1 898, Ross 220 (JE). Agrigento?: Palermo a Camastra, June (fl & fr), Todaro 1240 (BM, JE, K). Enna: ad Simethi prope Regalbuto, 28 June 1840 (fl), Heldreich s.n. (BM). MALTA. No precise locality, 1926 (fl), Bankart s.n. (BM). TUNISIA. Kroumerie: Ghardimaou, 22 April 1938 (fl), Simpson 38.360 (BM). Dorsale: Zaghouan, 250 m, 19 July 1969 (fl), Tomkinson 37 (BM). Kairouane: Bou-Arada, September 1898 (fl & fr), Le Testu s.n. (BM). Centrale: Dougga, 300 m, 16 June 1969 (fl), Tomkinson 29 (BM). ALGERIA. Constantine: Oued Zenati (fide Quezel & Santa, 1 963: 683). LIBYA. Cyrenaica: Gibel Akhdar, between Al-Marj and Beida, 25 April 1968 (fl), Boulos 3070 (BM). MONTENEGRO. S. end of Petrovak (between Budva and Bar), Cona Gora, 12 July 1976 (fl), Halliday 88/76 (LANC). ALBANIA. Present (fide Demiri, 1983: 315). GREECE. Makedhonia (Khalkidhiki): 8 km NE of Nea Moudania (ancient Olynthos), 100 m, 9 September 1981 (fl & fr), Hussein, Jury & RutherfordlS (BM, RNG*); Nea Potidaia, 6 August 1971 (fl), WestfieldColl. Exped. 315(BM).Thessalia: inter Larissa[Larisa]etTyrnaios, 31 July 1885, Haussknecht s.n. (JE). Sterea Ellas: S. of Athens on coastal road to Cape Sounion, 24 September 1962 (fr), Bourden 117 (BM); prope Athenas, Skaramanga, 28 May 1939 (fl), Lindberg s.n. (H). Argolis: in ruinis arcis Mykenai [Mycenae], 250 m, 26 June 1971 (fl), Cernoch 22070 (BM); excavations of ancient Epidaurus, 19 October 1975 (fl), Kramer 5521 (H). Messinia: near Kalamata, 100 m, 10 July 1975 (fl), Nydeggers.n. (BM, H). W. Aegean Is. : Idhra, Kamisi, 2 June 1968 (fl), N. Post 3260 (BM). Kikladhes: Naxos, inter Naxos et Chalki, 28 June 1932 (fl), Rechingerf. 2289 (BM). E. Aegean Is.: Khfos, Campos, 17 July 1939 (fl), Plan 362 (K); Kalimnos, ad Myrthies, c. 200 m, 3 June 1035 (st), Rechingerf. 7843 (BM); Samos,Vathy, 16-23 June 1932 (fi), Rechingerf 1915(BM).Dhodhekanisos: Rodhos, pres Salakos, 16 June 1870 (fl), Bourgeau 16 (BM, E, G, K); Rodhos, N. of Charakion, near ruins of Fernkios, 3 May 1952 (st), Fagersten s.n. (H). Karpathos: Avlona, 300 m, 24 July 1950, Davis 18024 (K). KRITI. Khania: La Cannee, 2 July 1883 (fl), 3 August 1883 (fr), Reverchon 17 (BM, FR, H, JE, K); Kissamos [Kisamou], 20 July 1938 (fl & fr), Davis 290 (BM). Rethimnon: Kesantopelta, 19 July 1976 (fl & fr), Raithalme s.n. (H). Iraklion: Knossos, ruins of Knossos Palace, 1 8 July 1079 (fl), Lassig 7914 (H). Lasithi: between Gournia and Tolotti, facing Psira I., 31 May 1966(fl), Whitefoord 11 (BM). CYPRUS. Paphos: Yeroskipos, 20 June 1939 (fl), Lindberg s.n. (H, K). Limassol: Prodhromos, 1350 m, 9 September 1955 (fl), Atherton 588 (K). Larnaca: prope Athienu [Athienou], 26 May 1862 (fl), Kotschy 972 (JE, K). Nicosia: Nicosia, English School, 150 m, 5 October 1958 (fr), Oswald 13 (K). Kyrenia: Kythraea, June 1880 (fl), Sintenis & Rigo 581 (BM, K). TURKEY. Europe - Tekirdag: Gallipoli, Kilia, n.d. (fl), Ingoldby 585 (K). Istanbul: KucukcekmecetoBiiyukc.ekmece,50-100m, 12 August 1962 (fl), Davis (& Coode) 39246 (BM, E, K). Asia - Canakkale: Umurbey, Umurbey Cayi, 22 July 1981 (fl), Kurtto 3306 (H). Kocaeli: Karamursel to Yalova, above Gemlik, 70 m, 8 July 1978 (fl), Uotila 27133 (H). Amasia: ad pagum Sana, 350-500 m, 10 July 1889 (fl), Bornmuller 1489 (BM, JE, K). Tokat: Tokat, n.d. (fl), Wiedermann 35 (JE, K). Izmir: Zeytindag civari, 18 September 1 954 (fr), Baytop 2654 (BM, ISTE*). Ankara: Ankara, 575 m, 10 July \939(fl),Frodin 199(U).Maras,:Elbistandistr.,Kapidere,31 July 1952 (fl), Davis (Dodds & £etik) 20419 (BM, E, K). Elazig: 60 km E. of Elazig, 1 STUDIES IN HYPERICUM September 1954 (fl), Davis (& Polunin) 24813 (BM, E, K). Bitlis: Bitlis Gorge S. of Kambos Dag, 1250 m, 16 August 1956 (fl), McNeill 598 (E, K). Mugla: Dat^a to Marmaris, 4 km, 10 m, Dudley in Davis 35439 (E, K). Denizli: Pamukkale, 400 m, 28 June 1967 (fl), Sorger 67-40-1 8 (BM, WU*). Antalya: Manavgat to Alanya arasi, 24 July 1957 (fl), Baytop 5043 (BM, ISTE*). I^el: Silifke-Mut main road, 19 June 1971 (fl), Ayanoglu, Sezile & fubukfu 1111 (BM). Gaziantep: Aintab [Gaziantep], 600 m, 16 June 1865 (fl), Haussknecht 8 1 2 (BM, E, JE, K). Mardin: Kischemill, 23 July 1 888 (fl), Stapf'm Sintenis 1343 (JE, K). SYRIA. North: prope Aleppo [Halep], 1771? (fl), Russell s.n. (BM); 30 km due W. of Balad Sinjar, 7 June 1934 (fl), Field & Lazar 685 (K) (or in Iraq?). South: Mt Hermon [Jebel esh Sheikh], 1 855 (fl), Kotschy 706 (H-M). LEBANON. Coast: bei Ghazir [Jdaidet Ghazir], c. 400 m, 19 June 1934 (fl), Busujan 220 (H-M, JE); Saida, pres de Darbesine, 28 June 1853 (fl), Blanche 950, 950a (JE, K). Lower mountains: Ain Zhalta and higher, 1950 m, 28 August 1954 (fl), Trott 2041 (K). ISRAEL. Samaria: 1 .6 km outside Nazareth on Tiberias road, 15 August 1962 (fl), Curie 61 (K); Jabal Qulayb [Mt Camel?], 1350 m, 12 June 1937 (fl), Myers & Dinsmore 12090 (H). Judaea: Jerusalem, Mt Scopus, 13 August 1933 (fl & fr), Amdursky 262 (H-M); Sheikh Nuran, May 1918 (fl), White s.n. (K); Ghaza, Abasan, 1 1 November 1964 (fl & fr), Boulos s.n. (K). JORDAN, near Amman, Al-Jubaiha, University Campus, 22 July 1974 (fl), Boulos 7300 (BM); 5 km S. of Shaubak, 29 June 1974 (fl), Jallad, Lahham & Hanania 609 (H); Khanazareh [Khanasira?], 800 m, 6 October 1950 (fl & fr), Chapman 51 (K). EGYPT. Sinai: Rajah, 4 June 1945 (fl), Davis 10426 (K). IRAQ. Amadiyah: Zawitah, 900 m, 27 July 1933 (e. fr), Guest in Rustam 3754 (K). Rawanduz: just below Chaklawa, Erbil liwa, 28 August 1962 (fl), Barkley & Asker 3413 (K). Sulaimaniyah: Kirkuk liwa, Jarmo, 750 m, 22 May 1955 (fl), Haines W.226 (K). Mosul: N. of Mosul, October 1929 (fl & fr), Dimmock in Rogers 0418 (BM, K). Nineveh: 50 km from Mosul to Dohuk, 410 m, 11 June 1958 (fl), Chapman in NHI 26253 (K). Arbil: 5 km N. of Erbil, 25 August 1962 (fr), Barkley 3355 (K). Kirkuk: Kor Mor, 30 km N. of Tuz, 9 June 1929 (fl), Rogers 0310A (BM, K). DiySla: Naft Khana [Naft Khaneh], June 1929 (fl), Rogers 0320 (K). IRAN. Kurdistan: Palangan, 1800-2 100m, 9 August \962(f\),Transhalr & Terme 12298E (W). Kermanshah: Ridjab, 24 July 1962 (fl), Transhalr & Terme 12300E (W). Ears: Kotal Pirjan, June 1960 (fl), Boroumand 5820E (EVIN*, W). Lorestan: Kouhdasht, Synareh?, 850 m, 13 August 1973 (fl), Moussari 34325E (BM, EVIN*). Hypericum triquetrifolium is the eastern Mediterranean counterpart of the Atlantic 2. H. undulatum; but it has quite a different habitat. The habit, however, although very different at first glance, is ap- proached by some widely branching forms of H. undulatum; and some of these (e.g. Valorado s.n. from Portugal, sine loc. (BM)) have rather similar but larger leaves. Hagemann & Meusel (1984) de- scribe it as a derived xerophilous species 5. Hypericum perforatum L., Sp. PL: 785 (1753), Syst. Nat. 12th ed.: 170 (1767); Choisy, Prodr. monogr. Hyperic.: 50 (1821), in DC., Prodr. 1: 549 (1824); Ledeb., Fl. altaic. 3: 364 (1831), FL ross. 1: 447 (1842); Spach, Hist. nat. veg., Phan. 5: 388 (1836), in Ann. Sci. nat. Bot. 5: 357 (1836); Rchb., Ic.fl. germ. helv. 6: t. 343, f. 5177 (1844); Boiss., FL orient. 1: 809 (1867); Maxim, in Bull. Acad. Imp. Sci. Saint-Petersbourg 27: 432 ( 1 882), Melanges Biol. Bull. Acad. Imp. Sci. Saint-Petersbourg 11: 166 (1882); S. Watson in Proc. Amer. Acad. Arts 17: 329 ( 1 882); Coulter in Bot. Gaz. 11: 107 (1886), in A. Gray, Syn.fl. N. Amer. 1: 288 (1897); Hemsley in /. Linn. Soc., Bot. 23: 74 (1886); Rouy in Rouy & Fouc., Fl. France 3: 332 ( 1 896); Reiche inAnales Univ. Chile 93: 561 (1896), Fl. Chile 1: 269 (1896); Diels in Bot. Jahrb. Syst. 29: 476 (1900); R. Keller in Bot. Jahrb. Syst. 33: 554 (1904); Schinz in Vierteljahrsschr. Naturf. Ges. Zurich 49: 241 (1905); Woron. in Kuzn., Busch & Fomin, Fl. Cauc. Crit. 3(9): 49 (1906); H. 87 Leveille in Bull. Soc. Bot. France 54: 595 (1908); Pampanini in Nuovo Giorn. Bot. Ital. 17: 672 (1910); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 520 (1911), in Mitt. Naturwiss. Vereins. Steiermark 51: 240 (1915); Thellung mAllg. Bot. Z. 1912: 19(1912); Rehder in Sargent, PL wilson. 3: 452 ( 1 9 1 7); Urban, Symb. antill. 9: 406 ( 1 925); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 180 (1925); Hegi, ///. Fl. Mitt.-Eur. 5(1): 526 (1925); Hayek, Prodr. Fl. Pen. bale. 1: 533 (1925); Fedtch. & Shishkin, Fl. Yugo-vost3: 711 (1930); Hand.- Mazz., Symbol, sin. 7: 404 (1931); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: t. 4 f. 32 (1932), 11: 176 (1933), 12: 88 ( 1 934), in Pflanzenareale 4: Karte 7 ( 1 933); Grossg., Fl. Kavk. 3: 68 (1932), 2nd ed. 6: 175, karta 194 (1962); Gorschkova in Shishkin & Bobrov, Fl. URSS 15: 248, t. 12 f. 1 (1949); Y. Kimura in Nakai & Honda, Nova Fl. Jap. 10: 131 (1951); Ross- Craig, Draw. Brit. PL 6: t. 7 (1952); Gu§ul. & Nydr. in Savul., Fl. R. P. Roman. 4: 29 (1956); Gleason & Cronquist, Man. Vase. Pis NE. U.S. & Canada: 469 (1963); O. Schwarz in Drudea 5: 60 (1965); Lauener in Notes Roy. Bot. Card. Edinburgh 27: 60 (1966); N. Robson in Davis, Fl. Turkey 2: 400, tt. 12 f. 4, 13 f. 23 (1967), in Tutin et al., Fl. Europaea 2: 269 (1968), in Rechinger f., Fl. Iranica 49 (Guttif.): 17 (1968), in Nasir & All, Fl. W. Pakistan 32 (Guttif.): 8, f. 2G-K (1973), in Cullen et al., Eur. gard.fl. 4: 60 (1995), in Wisskirchen & Haeupler, Standardliste Farn- u. Bliitenpfl. DeutschL: 270 (1998); Fosberg in Occas. Pap. Berenice Pauahi Bishop Mus. 24: 22 (1969); Jordanov & Koz. in Jordanov, Fl. R. P. Bulg. 4: 260, t. 49 ff. 3, 4 (1970); Stjep.-Vesel. in Josifovic, Fl. Srbije 3: 1 1 8 (1972); Anon., Iconogr. Cormoph. Sinicae 2: 879, f. 3488 (1972); Scoggan, FL Canada 3: 1097 (1978); Hagemann in Flora 183: 99, tt. l-8excl. 5B (1983); Ramos in Trab. Dept. Bot. Univ. Complut. Madrid 12: 49, t. 3 f. 2(l9S3),inActaBot.Malac. 11: 167,f.6a(1986), in Castroviejo et al., Fl. Iberica 3: 1 63, t. 45 ( 1 993); Valdes, Talavera & Galiano, FL Andalus. Occ. 1: 314 + map & fig. (1987); Li Xiwen in Fl. R.P. Sinicae 50(2): 67, t. 15 ff. 1-4 (1990); Biswas in Sharma & Sanjappa, Fl. India, 3: 73, f. 27 (1993); Benson & McDougall in Cunninghamia 4: 301 (1995); Azadi in Fl. Iran, 27: 11 + map (1999). Type: 'Habitat in Europae pratis' (probably Sweden), Herb. Linn. 943/34 (sphalm. 94) (LINNI-lectotype, Robson, 1968). Fig. 8 (for icones and maps see under subspecies). Perennial herb 0.2-0.6(-1) m tall, erect or rarely decumbent to procumbent from creeping and rooting base, budding on roots late in season,3 with stems numerous to few, much branched especially distally. Stems 2-lined, with few black glands on lines; internodes 5- 25 mm, shorter than leaves. Leaves sessile to 1 mm petiolate; lamina (5-)10-25(-30) x (l-)3-10(-15) mm, oblong or elliptic to linear or rarely orbicular or triangular-lanceolate, paler beneath, chartaceous; apex obtuse or apiculate to rarely acute, margin plane or ± recurved to revolute, base subcordate (or rarely cordate)-amplexicaul to rather narrowly cuneate; venation: c. 2 pairs of main laterals from lower quarter to eighth of midrib, tertiary reticulation lax or scarcely visible; laminar glands pale, scattered and sometimes black, few, punctiform; intramarginal glands black, spaced, interspersed with small dense pale ones. Inflorescence 3- to numerous-flowered, from 1-3 nodes, with flowering branches curved-ascending from up to 15 or sometimes more nodes below, the whole cylindric to broadly pyramidal or subcorymbiform; pedicels c. 0.5-2 mm; bracts and bracteoles up to 4(-7) mm long, narrowly lanceolate to linear, entire. Flowers 15-25(-35) mm in diam., stellate; buds narrowly ovoid, 'See Salisbury (1942: 106), Hagemann (1983: 106, f. 3 and refs.). N.K.B. ROBSON acute. Sepals 5, equal, 3-7 x 0.7-1.5 mm, narrowly oblong or lanceolate to linear, acute to finely acuminate with acumen some- times glandular, entire, erect in bud, recurved in fruit; veins 3(5), unbranched; laminar glands pale and often a few black, in 2(4) rows, striiform (basally) to punctiform; intramarginal glands few, black or absent. Petals 5, golden yellow, not tinged red in bud, (8-)12-15 x 5-6 mm, 3—4 x sepals, oblong to oblong-elliptic, asymmetric, distally ± crenate; laminar glands all pale to mostly black or rarely absent (subsp. chinense pro parte), linear or partly striiform to punctiform; intramarginal glands black, distal, in sinuses when present. Stamens 40-60, '3'-fascicled, longest 6-8 mm, c. 0.5-0.7 x petals; anther gland black. Ovary 3-locular, 3-5 x 1.3-1.8 mm, narrowly ovoid to ovoid-ellipsoid; styles 3, free, 4-6 mm, c. 1 .5-2 x ovary, broadly to rather narrowly spreading; stigmas narrow. Cap- sule 3-10 x 3-6 mm, 0.7-1.5 x sepals, narrowly to broadly ovoid or narrowly ovoid-conic; valves with dorsal vittae and lateral vittae or yellowish, striiform to punctiform vesicles. Seeds dark brown, c. 1 mm, cylindric, not carinate or appendiculate; testa finely linear- foveolate. 2n=32 (Noack, 1939, Mulligan, 1957etauct. plur.;n=16, Nielsen, 1924, Hoar & Haertl, 1932, Noack, 1939 et auct. plur.), 48 (Noack, 1939, Robson, 1981, Ciccarelli, Garbari & Martonfi, 2001), 2n=16 (Schwarz, 1965, as H. veronense; Strid & Franzen, 1981, as var. angustifolium', Papanicolaou, 1984, Ciccarelli, Garbari & Martonfi, 2001, as H. perforatuiri). Open woodland, meadows, grassland and steppes, riverbanks, stony and grassy slopes, roadsides, in dry or well-drained habitats; 10-c. 2750 m (China), -2780 m (Tajikistan), -3150 m (Afghanistan). General distribution of the species: Europe (except extreme north), north-west Africa, Canary Is., Madeira, Azores; Turkey, Cyprus, the Levant and western Saudi Arabia to north-west India (Uttar Pradesh), Transcaucasia, Turkmenistan to Altai, Angara-Sayan and NW Mon- golia; China (W. Xinjiang and from Gansu east to Hebei, south to Jiangxi and west to Yunnan). Introduced into many other parts of the world, viz. Canada, U.S.A., Mexico, Cuba, Haiti, Brazil (Parana), Uruguay, Argentina, Chile, Juan Fernandez, Sudan Rep. (Jebel Marra), S. Africa, Reunion, Australia, New Zealand, Japan. Hypericum perforatum is apparently an allotetraploid (2n=32) (Gustafsson, 1947; Robson, 1958«, 1975), which, for reasons of morphology and geography, would appear to have arisen from a cross between two diploid taxa (2n=16), viz. la. H. maculatum subsp. immaculatum (Balkans) and 6. H. attenuatum (western Sibe- ria to China). Morphologically it (or, rather, typical broad-leaved var. perforatum) is close to H. maculatum, but it differs in the 2-lined stem internodes, the more oblong leaves with lax reticulate venation and rather dense pale laminar gland dots, the lanceolate acute sepals, and the petals with black marginal glands, characters that all tend towards H. attenuatum. It differs from both presumed parents in its capsule valves with oblique vittae or vesicles, a glandular pattern that is unique in sect. Hypericum (of which it is the type species). The presence of linear laminar glands in the petals and pale laminar leaf glands suggests that the maculatum parent was var. immaculatum rather than var. maculatum, with its punctiform to striiform petal glands and usual lack of pale laminar leaf glands. If, as seems probable, the ancient hybridization occurred in Central Asia (the Altai region?), then (i) the current restricted distribution of var. immaculatum in the Balkans is a relict of a much wider one, and (ii) the extension of the current distribution of var. maculatum into Central Asia is relatively recent. Cytologically, H. perforatum behaves as a hybrid (Noack, 1939; Gustafsson, 1 946^7 ; Martonfi et al., 1996a). The pollen undergoes normal reduction division; but meiotic abnormalities (lagging chromosomes) lead to sterility that is usually about 30% but can be up to 70% (Nielsen, 1924; Hoar & Haertl, 1932). Only c. 3% of the embryo sacs are produced by normal meiosis (n=16), the remaining (unreduced) ones being produced parthenogenetically (n=32). The latter, however, are pseudogamous, i.e. they require pollination for seed development. Fertilization is therefore possible, though not necessary. Occasionally the n=32 embryo sacs are fertilized, result- ing in hexaploid (2n=48) plants. The records of H. perforatum with 2n= 1 6 chromosomes (see p. 63) would appear to be from dihaploids, i.e. from plants that have resulted from the development of reduced but unfertilized embryo sacs. Since in allotetraploids (i) each chro- mosome is duplicated and (ii) the more different the parents are, the less is the likelihood that there will be multiple chromosome asso- ciations, it seems reasonable to expect dihaploids to occur in H. perforatum. Two of these occurrences have been reported in the more reduced forms of the species (var. angustifolium and var. microphyllum), and two (Papanicolaou, 1984; Ciccarelli, Garbari & Martonfi, 200 1 ) were in H. perforatum without qualification. Gagnieu & Wilhelm's report ( 1 965) of n= 1 2 for var. angustifolium is no doubt either a miscount for n=16 or, just possibly, from fertilization by a triploid pollen grain. This cytological variation is accompanied by wide morphological variation, which is complicated in the wild by 'back-crosses' to at least two of the subspecies of H. maculatum (Robson, 1981: 168, f. 55) (see p. 104). The morphological variation in H. perforatum, though great, appears to be continuous and therefore theoretically indivisible. For reasons of practicality, however, it is convenient to recognize some previously described variants as subspecies4, a rank that reflects their geographical basis better than the hitherto more usual varieties. For one well-known taxon, however, even this is impossible. The southern European plants with narrow leaves and vesicular capsule vittae ('var. angustifolium DC.') cannot be separated from those with small (narrow to broad) leaves and similar capsule vittae ('var. microphyllum DC.'). Indeed, the latter are probably polyphyleti- cally derived from the former, and some may be merely the result of habitat-induced variation. Bearing in mind these reservations, the overall variation in H. perforatum can be described and classified in four intergrading subspecies. (a) From its presumed origin in western Siberia into northern and central Europe and eastward into NW Mongolia the initially sessile leaves become gradually shortly petiolate (although remaining sessile in northern Russia (always?) and sometimes in Scandinavia), and the base changes from rounded to broadly cuneate, the capsule vittae remaining continuous and almost always narrow. This is the type form (subsp. a. perforatum). In dry habitats the leaves become narrow with revolute margins, thus superficially resembling a form of subsp. veronense (see c. below); but the leaves are petiolate and the capsule glands are different. Such plants have frequently been recorded as 'var. angustifolium DC.', but the vesiculate capsule vittae differ from those of similar forms of subsp. veronense. Where the capsule is vesiculate (as it is, for example, in southern England on chalk, sand, wall-tops, etc.), such capsules are derived from continuous lateral vittae, as is shown by the occurrence of intermediate states. True 'var. angustifolium' would appear to have been derived from forms in which the lateral vittae have been inter- rupted before becoming enlarged and vesiculate. Another development of vesiculate capsules of this type in subsp. 4For H. perforatum var. latifolium Gaudin and synonyms, see under 1 7xb. H. x desetangsii nothosubsp. carinthiacum nothoforma perforatiforme. STUDIES IN HYPERICUM perforatum has occurred in the southern Balkans. In the mountains of Bulgaria and northern Greece there are erect to procumbent plants with small, obovate to orbicular leaves and capsules with lines of flattish vesicles ('van humile Stranski'). These are interpreted here as an extreme montane development of subsp. perforatum, there being continuous variation from (i) typical subsp. perforatum to (ii) a smaller-leaved form with rather dense inflorescences and small fruits, then to (iii) a similar, decumbent form with capsule valves having two lines of flattish 'vesicles', and finally to (iv) a dwarf form with small narrower leaves. (b) Both presumed parents have sessile leaves, with the base rounded in H. maculatum and subcordate to cuneate in H. attenuatum. Therefore the one variant of//, perforatum with sessile cordate- amplexicaul leaves would also appear to have a good claim to at least approach the primitive form. This is subsp. b. songaricum (var. songaricum (Ledeb. ex Rchb.) K. Koch), of which the tallest plants with the largest leaves and most luxuriant growth (approaching those of the presumed parental species) occur in eastern Kazakhstan and probably also in NW Xinjiang. The leaves become gradually smaller southward (Kyrgyzstan - '//. komarovW) and westward (W. Kazakhstan, S. Russia and the southern Ukraine - var. 'gracile ') and in dry regions develop a glaucous lower surface and recurved margins. (c) From the Caucasus region into eastern Turkey and northern Iran narrower-leaved plants become frequent ('var. collinum Woron.), and the capsules, which are still relatively large, have interrupted and often somewhat swollen oblique vittae. The leaves then become first narrower and then smaller (often with revolute margins) eastward into central Asia (Tadjikistan) and India (east along the Himalaya to Uttar Pradesh) and westward into the Mediterranean region, where the typical narrow-leaved and small-leaved plants of subsp. veronense (= respectively var. angustifolium DC. and var. microphyllum DC.) occur and ex- tend into Macaronesia, NW Africa, SW Saudi Arabia (Asir) and the Sudan Republic (Jebel Marra), although whether its occur- rence in Jebel Marra is natural is doubtful (Wickens, 191 6b: 76, 99, map 59). It has been introduced into various other countries and continents. Where its distribution meets that of subsp. perforatum in southern and central Europe, the intermediate forms that occur are presumably hybrid in origin, unless subsp. veronense has arisen polyphyletically in southern Europe, the Caucasus region and (possibly) Central Asia. In the trends in this subspecies the leaves first become narrow (vars collinum, angustifolium) and then small (var. microphyllum=subsp. veronense); but in relatively mesophytic areas these small leaves may become broad again (e.g. var. ellipticum Freyn non Durand & Pittier in mountain Turkmenistan). (d) In China, after a gap in distribution from NW Mongolia to Gansu and adjacent Qinghai (where some specimens closely approach subsp. perforatum in form), there is a southward trend to Yunnan and Guizhou in which the leaves and flowers become smaller, the latter being crowded in small (terminal and lateral) inflorescences at the end of elongated branches (subsp. d. chinense). This subspecies has been introduced into Japan ('//. foliosissimum Koidz.'). 5a. Hypericum perforatum subsp. perforatum Stjep.-Vesel. in Josifovic, Fl. Srbije 3: 118 (1972); Ramos in Castroviejo et al., Fl. Iberica 3: 165, t. 45 ff. f-j (1993). Fig. 8, Maps 6-8. H. offidnarum Crantz, Stirp. austr. 2: 66 (1763), 2nd ed.: 99 (1769); 89 Moench, Methodus: 129 ( 1 794), nom. illegit. Crantz cites the Sp. pi. phrase name (except 'Floribus trigynis' ) and number. Type as for //. perforatum L. H. vulgare Lam., Fl. France 3: 151 (1778); Rupr. in Mem. Acad. Imp. Sci. Saint-Petersbourg 7, Ser. 15, no. 2: 248 (1869), nom. illegit. Type as for H. perforatum L. H. officinale Gaterau, Descr. pi. Montauban: 135 (1789); Steud., Nomencl. Bot.: 421 (1821) qua syn.; Raynal in Taxon 17: 516 (1968), nom. illegit. Type as for H. perforatum L. H. perforatum var. [fi] sensu Choisy, Prodr. monogr.fam. Hyperic.: 50(1821). ? H. perforatum var. [£] albiflorum Choisy in DC., Prodr. 1: 550 (1824). Type: not found (1974). H. perforatum var. vulgare Schimp. & Spenn. in Spenn., Fl.friburg 3: 888 (1829); Neilr., Fl. Nied.-Oesterr. 2: 826 (1859); Hegi, ///. Fl Mitt.-Eur. 5(1): 528 (1925); Briquet, Prodr. fl. Corse 2(2): 150 (1936); Gu§ul & Nyar. in Sjavul., Fl. R. P. Roman. 4: 29 (1956); Grossg., FL Kavk. 2nd ed. 6: 1 75 ( 1 962). Type as for//, perforatum L. H. marylandicum Biroli ex Colla, Herb, pedem. 1: 467 ( 1 833), 7: 97, t. 50 f. 2 (1837). Type: no locality cited, Herb. Birol. (TO- holotype, photograph!). H. perforatum var. [y] latifolium sensu W. Koch in Rohling, Deutsch Fl. 3rd ed. 5: 349 ( 1 839), Syn.fl. germ. helv. 2nd ed. 1: 146 ( 1 843), Syn. deut. schweiz. Fl. 2nd ed. 1: 155 ( 1 846) pro parte omnes excl. typum. The type of Koch's var. latifolium is that of var. latifolium Gaudin (1829), which he cites; his specimens (L) are typical H. perforatum subsp. perforatum. See 5xbb. H. x desetangsii nothosubsp. carinthiacum nothoformaperforatiforme for Gaudin's plant. H. assurgens Peterm., Anal. Pfl.-Schliissel: 67 (1846), in synon.; Rouy in Rouy & Fouc., Fl. France 3: 337 (1896). Type: Germany, Leipzig, bei Roumaritz, Petermann s.n. (W!-holotype). The 'type specimen' appears to be pure H. perforatum, not H. humifusum x perforatum as reported by Rouy & Foucaud; and the name does not appear on Petermann's Fl. Lips.: 563 (1838) as cited by them. H. perforatum var. petiolatum Peterm., Anal. Pfl.-Schliissel: 67 (1846). Type: Germany, Leipzig, Konnevitz, Petermann s.n. (LZf) H. lineolatum Jord. in F.W. Schultz, Arch.fl. France Allem. 1: 343 (1854). Type: France, Ain, 'montagnes du Bugey a Junnimont', Jordan s.n. (LY-lectotype, selected here; BM!-isolectotype). ? H. deidesheimense Sch. Bip. ex Trev., Hyper, sp. animadv.: 10 (1861). Type: Germany, Rheinhessen-Pfalz, Deidesheim, Schultz- Bip. s.n. (P?-holotype). H. perforatum var. alpinum Parl., Fl. Ital. 5: 512 (1872); Fiori & Paol., Fl. Italia 1: 388 (1898); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 524 (1911). Type: Italy, Piemonte, sopra i Bagni di Valdieri al Vallasco nella regione superiore del Faggio, 1300 m, Parlatore s.n. (Fl-lectotype, Ciccarelli, Garbari & Robson, 2002); come pure nella regione superiore dell'Abeto del Monte Cramont', Parlatore s.n. (FI- syntype). Frohlich (1911: 524) compares (synonymizes?) this with his forma lineolatum of subsp. vulgare. H. perforatum var. anomalum Frid. in Lange, Haandb. Danske fl. 4th ed.: 700 (1887). Type: Denmark, Slesvig [Scederjylland], Haderslev, 1886 (fl), Haspelmath s.n. (C!-holotype). H. perforatum var. [a] typicum Beck, Fl. Nieder-Osterr. 2: 530 (1892); Fiori & Paol., Fl. Italia 1: 388 (1898). Type as for H. perforatum. H. perforatum var. vulgare subvar. lineolatum (Jord.) Rouy in Rouy & Fouc., Fl. France 3: 333 ( 1 896). Type as for H. lineolatum Jord. H. perforatum var. ellipticum Durand & Pittier in Bull. Soc. Roy. 90 N.K.B. ROBSON Fig. 8 H. perforatum. A. subsp. perforatum: (a) habit; (b) sepal; (c) petal, omitting laminar glands; (d) capsule. B. subsp. songaricum: (e) branch; (f) capsule. C: subsp. veronense: (g) branch; (h) sepal; (i) petal, omitting laminar glands; (j) capsule. D. subsp. chinense: (k) stem and branch; (1) leaves (depauperate); (m) capsule (a, e, g, k, 1 x 2/3; rest x 6). A. (a-c) Martinez s.n.; (d) Sennen s.n. B. (e, f) Becker s.n. C. (g-i) Riera et al. 17197; (j) Lacaita 28. D. (k) Wilson 2425; (1) Bullock s.n.; (m) Father Hugh s.n. STUDIES IN HYPERICUM Bot. Belgique 20: 80 (1881). Type: Switzerland, Vaud, bois pres de Rol\e,fide 'H. perforation [var.] a' sensu Rapin, Cat.fl. vaud., 2nd ed. : 123(1 862) (G?-holotype). Hegi ( 1925: 528) reduced this to a form of var. vulgare. H. vulgare Bubani, Fl. pyren. 3: 345 (1901), non Lam. (1778). Bubani ascribed this name to pre-Linnaean authors. H. perforatum var. normale Woron. in Kuzn., Busch & Fomin, Fl. Cauc. Crit. 3(9): 56 (1906). Type as for H. perforatum L. H. perforatum subsp. vulgare (Schimp. & Spenn.) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 522, f. 2 (1911), in Mitt. Naturwiss. Vereins Steiermark 51: 240 (1915); Hayek, Prodr. Fl. Pen. bale. 1: 533 (1925). H. perforatum subsp. vulgare forma brevisepalum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 524 ( 1 9 1 1 ). Type: Austria, Styria, Costing bei Graz, 1 1 July 1 909 (fl), Frohlich GZU16368 (GZUI-lectotype, selected here). H. perforatum subsp. vulgare forma lineolatum (Jord.) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 524(191 1), mMitt. Naturwiss. Vereins Steiermark 51: 240(1915). Type as for H. lineolatum Jord. Frohlich also describes this taxon in adnot. as 'subvar. lineolatum'' , a later homonym than that of Rouy(1896). H. perforatum subsp. vulgare forma lucidum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. KL, 1, 120( 1 ): 524 (191 1), in Mitt. Naturwiss. Vereins Steiermark 51: 240 (1915). Type: Austria, Styria, Stiftingtal bei Graz, 26 July 1910 (fl), Frohlich GZU16378 (GZUI-lectotype, selected here). H. perforatum var. humile Stranski (date?) [description not found]; Stoj. & Stef., Fl. Bulg. : 774 ( 1 948); Stoj., Stef. & Kitan., Fl. Bulg. 2: 720 (1967). Type: Bulgaria (SO). H. perforatum var. vulgare forma brevisepalum (A. Frohl.) Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925). 91 H. perforatum var. vulgare forma anomalum (Frid.) Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925). H. perforatum var. vulgare forma lineolatum (Jord.) Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925), sphalm. 'lanceolatum'. H. perforatum var. vulgare forma lucidum (A. Frohl.) Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925). H. perforatum subsp. vulgare var. lineolatum (Jord.) Hayek, Prodr. Fl. Pen. bale. 1:533(1925). H. perforatum var. ceretanicum Sennen, ined.? 'Type' : Spain, Lerida, La Cerdana, Targassonne, 1600 m, 15 September 1916 (fr), Sennen s.n. (BC, BM!). H. perforatum var. decompositum Nyar. in Bui. Grad. Bot. Univ. Cluj 19: 85 (1939); Gu§ul. & Nyar. in Savul., Fl. R. P. Roman. 4: 29 (1956). Type: Romania, Transsilvania, distr. Treiscaune, ad balneas §uga§, supra opp. Sf. Gheorghe, 7 September 1928 (fr), Nydrddy s.n. (CL-holotype, photocopy!). H. perforatum var. semihumifusum Nyar. in Bui. §ti. Acad. Republ. Populare romme, Sect. §ti. Biol. etc. 7: 226, t. 2 f. 3 (1955); Gu§ul. & Nyar. in Savul., Fl. R. P. Roman. 4: 29 (1956). Syntypes: Romania, Walachia, Pite§ti, Mt. Cozia, 1200-1400 m, 17 May, 20 June & 19 August 1950 (fl & fr?), Nydrddy s.n. (CL). H. perforatum subsp. perforatum var. pellucidum O. Schwarz in Drudea 5: 61 (1965), nom. illegit. (Art. 37.1). Type: none cited. H. perforatum var. perforatum - N. Robson in Wisskirchen & Haeupler, Standardliste Farn- u. Bliitenpfl. Deutschl. : 270 ( 1 998). Icones: Rchb., Ic. fl. germ. helv. 6: t. 343, f. 5177 (1844); Ross- Craig, Draw. Brit. PL 6: t. 7 (1952); Ramos in Castroviejo et al., Fl. Iberica3: t. 45 ff. f-j (1993). Leaves usually petiolate; lamina (5-)12-25(-30) x (2-)5-10 mm, broadly to narrowly oblong or ovate to rarely elliptic or orbicular or obovate (l:b=(l-)2-3(-5)), base rounded to broadly cuneate, not Map 6 5a. H. perforatum subsp. perforatum, Eurasia •, other records O. The north European plain, Bavaria and the upper Danube region under-represented. 92 glaucous beneath. Inflorescence not usually congested, with branches relatively short, straight. Petals with laminar glands all pale to mostly black. Capsule valves with lateral vittae linear, narrow or rarely swollen distally but not interrupted, the vesicles forming a regular row, not scattered irregularly. Distribution of the species excluding (i) the Mediterranean region, Macaronesia, Jebel Marra, SW Saudi Arabia (Asir), nearly all Asiatic Turkey, Transcaucasia and Iraq to NW India; (ii) Kazakhstan and adjacent southern Russia; (iii) China. Introduced into North America, Mexico, Cuba, Haiti, Brazil (Parana), Uruguay, Argen- tina, Chile, Juan Fernandez, Hawaiian Islands and possibly New Zealand and Australia (no confirmed records). SCOTLAND. Sutherland: Kildonan, 1882 (Anthony, 1976: 64). Easter Ross: Strathpeffer, 1967 (Duncan, 1980: 40). Inverness: Beauly, 1954 (McCallum Webster, 1978: 115). Moray: Forres, near Greshop House, 12 August 1961 (fl), McCallum Webster 5864 (K). S. Aberdeen: Ballater, mouth of R. Muick, 30 July 1955 (fl), Robson s.n. (BM). Forfar: Clova, Birkhill, n.d. (fl), Gardiner s.n. (BM). Mid Perth: Aberfeldy, July 1929 (fl), Meinertzhagen s.n. (BM). Stirling: Loch Lomond. Balmaha, 30 July 1962 (fl), Kenneth s.n. (BM). Mid Ebudes: Mull, Achnadrish, July 1968 (e. fl), Kingston s.n. (BM). Midlothian: Inveresk, 1851 (fl), Syme s.n. (BM). Berwick: Nenthorn parish, near Newton Don, 3 August 1960 (fl), Bangerter 948 (BM). Kirkcudbright: Tongland, July 1883 (fl), Coles s.n. (BM). ISLE OF MAN. Peel, 5 September 1944 (fl), Paton s.n. (BM). ENGLAND. Cumbria: Seascale, September 1903 (fl), N. Simpson 3025 (BM). Northumberland: Middleton Hall, Wooler Water, 4 July 1958 (bud), Bangerter 645 (BM). NE Yorks.: near Hill Ghyll, 4 August 1926 (fl), Foggitt s.n. (BM). Mid-W. Yorks.: Arncliffe village, 18 July 1959 (fl), Gerranslll (BM). E. Yorks.: Wauldley, 2 August 1951 (f[),Radfords.n. (BM). S. Lanes.: Liverpool, Oglet, 13 July 1972 (fl), Greenwood s.n. (LIV). Notts.: Budby, Fanny's Grove, 20 July 1963 (fl), Bowden & Hillman 37 (BM). S. Lines: Grantham, Sattersford, 1 9 July 1913 (fl), Fishers.n. (BM). Shropshire: Ferny Dingle, lOJuly 1961 (f\), Butler &Milward4\5(BM). W.Gloucs.:Tidenham, 1 6 July 1 909 (fl), Riddelsdell s.n. (BM). Bucks.: 1 .6 km SE of Ellesborough, Coombe Hill, 28 July 1954 (fl), Melderis 681 (BM). E. Norfolk: Lopham Middle Fen, 3 July 1973 (fl), Jones & Vickery 29 (BM). N. Essex: Berechurch, 2 July 1954 (fl), Jermyn 226 (BM). E. Kent: Isle of Sheppey, near Warden Point, 20 September 1953 (fr), Bangerter 205 (BM). Surrey: near Croydon, Riddlesdown, 17 July 1954 (fl), 7. &M. Cannon 2761 (BM). N. Hants.: near Alton, Selborne Hanger, 22 July 1974 (fl), Gerrans 1817 (BM). Dorset: Parkstone, 23 October 1 927 (fr), Salmon s.n. (BM). N. Somerset: Cle vedon, 1 1 July 1934 (fl), Waterfall s.n. (BM). S. Devon: Buckfastleigh, 4 July 1973 (fl), Chicken 524 (BM). W. Cornwall: N. of Camborne, near Godevry and Gwithian, 27 June 1953 (fl), Murphy s.n. (BM). WALES. Glamorgan: Parkmill, July 1903 (fl), Riddelsdell s.n. (BM). Radnor: near Llangunlo Stn, 5 August 1956 (fl), P.C. & J. Hall 52/56 (BM). Merioneth: Penmanpool, 7 July 1961 (fl), Raven [& Benoit] 16308 (BM). Caernarvon: Llandudno junction, Conway estuary opposite Conway, Au- gust 1 962 (fr), Brummin 62.224 (LIV). Denbigh: S. of Llangollen, Pengwern, 28 August 1877 (fl), Bailey s.n. (BM). IRELAND. E. Cork: near Fernroy, 1 849 (fl), Chandlee s.n. (BM). Mayo: Mulranny, 16 August 1947 (fl & e. fr), Ross-Craig & Sealy 1520 (K). Carlow: Staplestown, n.d. (fl), Collector! (BM). Wexford: W. of Bunclody, Drumderry quarry, 5 September 1962 (fl & fr), McCallum Webster ZQ48 (K). Wicklow: Ovoca, July 1 882 (fl), Fawcett s.n. (BM). Down: near Newcastle, Slidderyford, 6 September 1952 (fr), Turrill s.n. (K). Antrim: Logan Canal, 1.6-5 km from Belfast, n.d. (e. fr), A.G. More s.n. (BM). CHANNEL ISLES. Jersey: St. Helier, 14 July 1883 (fl), Fawcett s.n. (BM). THE NETHERLANDS. Gelderland: Bennekom, b/d Vijrus, 20 July 1927 (fl), Vermeulen s.n. (K). Brabant: Valkenswaard, 4 August 1 979 (fl), P. vanRoyen 11833(K,L*). BELGIUM. W. Vlanderen: near Proven, 1 July 1918 (fl), G.C. Brown s.n. (BM). Brabant: Opwijk, vallei van de Brabantse beek, 18 August 1981 (fl), Robrecht 1533 (BM, BR*). Hainault: Obourg, 8 August 1862 (fr), Martinis in van Huerck I 42 (BM, K). Namur: Wavreille, June 1914 (fl), G.A.B. s.n. (BM). N.K.B. ROBSON FRANCE. Ardennes: Rethel, 1 August 1897 (fr), Gerard in Soc. Rochel. 4049 (BM). Oise: Paris, foret de Chantilly, 15 June 1912 (fl), Ronniger s.n. ( W). Hts de Seine: Bois de Meudon pres Paris, October 1 875 (fl), Bonnet s.n. (K). Finisterre: Brittany, Haelgost, 22 July 1913 (fl), Perrycoste 3639 (BM). Loire atl.: Mauves, July 1 860 (fl), Gadeceau s.n. (BM). Maine-et-Loire: la route a Montfaucon s. Moine, 10 July 1888 (fl), Gadeceau s.n. (BM). Hte Vienne: Le Dorat, 21 July 1929 (fl), Jalicon s.n. (BM). Cher: Noirlac, 22 June 1 864 (fl), Deseglise s.n. (BM). Hte Saone: entre le Loritellet et le mont pres Conflans, 2 August 1 856 (fl), Perrier 211 (BM). Jura: Salins les Bains, 425 m, 15 August 1892 (fl), Deschamps (BM, BR*). Hte Savoie: Mont Saleve, 29 July 1922, Lacaita 29/22 (BM). Savoie: Belleville, 300 m, 2 September 1861 (fr), Perrier in Billot 3349bis (BM). Puy de Dome: Au- vergne, Mont Dore, August 1920 (fl), Norman s.n. (BM). ANDORRA. Vallee du Valira, July 1933 (fl), Font in Sennen s.n. (BM). SPAIN. La Coruna: near Coruna, August 1 888 (e. fr), Guardia s.n. (BM). Orense: c. 50 km E. of Orense, Monte Manzaneda, 1410 m, 22 August 1982 (fl), Goyder & Jury 606 (BM, RNG). Santander: Picos de Europa, between Fuente Be and Espinama, 877-1200 m, 5 August 1969 (e. fr), Grimes 319 (BM, RNG). Viscaya: Monasterio de Nuestra Senora de Guadalupe, Caceres, 24 June 1958 (fl), Guinea 2789 (RNG). Lerida(?): Cerdagne, Estavar, 1230 m, 8 September 1929 (fr), Sennen s.n. (BM). Madrid: Madrid, Real Casa de Campo, June 1930 (fl), Martinez s.n. (BM, MA*). Avila: Hoyacasero, Rio Alberche at Venta del Obispo, 1250 m, 14 July 1983 (fl), M. & S. Gardner 2002 (BM, RNG). PORTUGAL. Minho: [Braga] juxtaGuimaraeus [Guimaraes] in Durminia [Minho?], n.d. (fl), Baron Paiva s.n. (K). ITALY. Novara: Alpi Lepontine, Valle Formazzo, Altillone, 1300 m, 4 August 1912 (fl), Roggiani s.n. (BM). SWITZERLAND. Ticino: Monte Generoso, 6 July 1 856 (fl), Murray s.n. (BM). Valais: Brieg [Brig] to Viesch, 22 July 1881 (fl), Linton s.n. (BM). Vaud: near Vevey, 30 June 1856 (fl), Murray s.n. (BM). AUSTRIA. Niederosterreich: Weinverteil, Hagenbrunn, zwischen Kronawettberg und Tradenberg, 23 June 1993 (bud), Vitek s.n. (BM). Steiermark: ad pagum Costing prope urbem Graz, 460 m, September 1909 (fl & fr), Reiterm Hayek, Fl. Stir. Exsicc. 1016 (BM). Tirol: Seefeld, 1200m, 23 August 1936 (fl), Wyatt 77 (K). GERMANY. Baden-Wurttemburg: Schwarzwald, Heselbach, 4 August 1931 (fl), Williams s.n. (BM). Hessen: Milseburg, 13 September 1913 (fl), Arnold s.n. (FR). Thuringen: Strasse Wilhelmstal-Etterwinden, 7 August 1948 (fl), Launert 166 (BM). Schleswig-Holstein: Forst Neumunster, 17 July 1969 (fl), Holm-Nielsen et al. in Fl. Germ. Exsicc. 16 (BM, TAI). DENMARK. Jylland: Ajstrup, S. of Aarhus, 23 July 1965 (fl), Pedersen in Fl. Jutl. Exsicc 140 (BM, TAI). Sjaelland: Nordsjaelland, Tisvilde Hegn., 1913(fl),Da«/D.45b(BM). NORWAY. Vestfold: Jahlsberg, Oslo Fjord, Aasgaardstrand, 10 Septem- ber 1934 (fl), Williams s.n. (BM). SWEDEN. Blekinge: Kellovik, Injauley, 191- (fl), F.G. Johansson s.n. (BM). Malmohus: Malmo, July 1 886 (fl), G. Johansson s.n. (BM). Goteborg: Ringon, 23 July 1957 (1. fl), Blom s.n. (BM). Smaland: Madesjo, 29 July 1909 (fr), Medelius s.n. (BM). Gotland: Visby, July 1912 (fl), E. Th. Fries s.n. (BM). Sodermanland: Millsten, 8 August 1929 (fr), Asplund s.n. (BM). Uppsala: Soderby Karl, Brolunde, 12 July 1956 (fl), H. Smith 2037 (BM). Varmland?: par. Hilared, ad Tarsered, 12 July 1916 (fl), Ohlson s.n. (BM). FINLAND. Uusimaa: Varsinais-Suomi, Korppoo, Jurmo, Huwudskar, 23 July 1965, Kause & Seikkula s.n. (JE). ESTONIA. Tartu: Dorpat bei Ropkoi, 25 July 1860 (e. fr), Gruner s.n. (BM). BELORUS. Distr. Nowogrodek [Novogrudsk], ad Niarikow, 1893 (fl), Dybowski in Rehman & Wolszczak, Fl. polon. exsicc. 148 (BM). RUSSIA. St Petersburg?: Ingria, June-August 1860 (fl), Herb. Fl. Ingricae 134 (BM, K). The species is recorded by Gorschkova (1949: 249) from all regions, except Karelia-Lapland and the Arctic, east to Angara- Sayan (Chuna Angara watershed). The precise areas of overlap between the distributions of subsp. perforatum and subsp. songaricum are uncertain but appear to be in the central Ukraine and the southern Altai (see Maps 6 and 9). The species (no doubt subsp. perforatum) has also been recorded from the Mongolian side of the Altai mountains (Gubanov, 1996: 76). POLAND. Lublin: Lublin distr., 1 July 1 884, Krolestwa s.n. (JE). Bielsko- Biala: Wadowice, Wiepoz k. Andrychowa (pr. Andrychov), 1 8 July 1 938 (fl), hincucka in PI. Pol. exsicc. 328 (BM, JE, K). Jelenia Gora: Sudetia, 93 Hirschberg, Perle des Westens, 1 August 1958, Kohler & Zippold s.n. (JE). Walbrzych?: Reichenstein [Zloty Stok], Dorndorf, 20 August 1899 (fl), Troedel s.n. (FR); Schweidnitz [Swidnica], Koltschenberg, September 1882 (1. fl), Peck s.n. (FR). CZECH REPUBLIC. Bohemia: prope vicum Pruhonice haud procul Pragam, c. 300 m, 22 August 1958 (1. fl & fr), Nitka in Fl. Czech. Exsicc. 1 48 (BM). SLOVAKIA. BratislavskyKn: N. Vratislava, 1803, Collector? s.n. (JE). HUNGARY. [Province?]: Gau Mayn?, August 1873 (fl), Leichternissl s.n. (BM). ROMANIA. Transylvania: [Carus Severin] Banat, N. of Turnu Severin, 24 km above Baile Herculane up Cerna valley, 27 July 1971 (fl), Frazer Jenkins 3499 (BM). Moldavia: Mun. Baciiu, prope pagum Magum, c. 400 m, 25 August 1971 (fl), D. Mititelu, Baraba§r & L. Mititelu in Fl. Exsicc. Distr. Bacov. 351 (BM). Walachia: Oltenia, distr. Gorj, Cimpul Experimental Ciociriau, 29 June 1965 (fl), Cirtu & Zaharia in Fl. Olten. Exsicc. 960 (BM). BULGARIA. Tolbukhin: Suddobrudscha, Kap Kaliskra, 16 August 1958 (fr), Meyer 645 (JE). Varna: Suddobrudscha, Varna, Weg vom Strande zum Aladsha Monastir, 11 August 1958 (fl & fr), Meyer 414 (JE). Sliven: prope Sliven, in monte Sini Kamil, 20 June 1 907 (fl), Schneider It. bale. 583 (BM). Yambol?: Strandsha-Planina, zwischen Blgari und Kosti, c. 300 m, 14 October 1958 (fr), Meyer 2088 (JE). Stara Zagora: Stara Planina, Krstec, zwischen Stara Zagora und Trnovo [Veliko Turnovo], 28 October 1958 (fl), Meyer 2234 (JE). Smolyan?: in Rhodope Centralis ad Selca-Djorlen, 12 July 1924 (fl), Stefanoffs.n. (BM) ['van humile']. SLOVENIA. Carniola, September 1854 (fl), Ball s.n. (K). CROATIA. Dalmacija: Velebit Mts, 840 m, 2 July 1962 (fl), Mathew & Pycraft2l2(EM). SERBIA. Srbija: Vranja [Vranje], 1919 (fl), Tmdal Lucas 104 (BM) [hybrid?]; circa urbem Gevlegi supra pagum Sermenin, Mt Rozin, 17 July 1937 (fl), Grebenchikqffs.n. (K). Kosovo: Bertiscus (Alpes boreales albanicae) in fauce rivuli Susica prope oppidum Pec (Ipek), 750-950 m, (fl), K.H. Rechlnger & Scheffer Iter bale. 1933 782 (K). BOSNIA. Sarajevo, 750 m, 12 July 1960 (fl), McCallum Webster 4036 (K). MONTENEGRO. Distr. Vasojavidi, Korpice, 30 August 1903 (fl & e. fr), Hannibal in Baldacci Alb. X 44 (BM). Distr. Kuci, Poprat (Trijapsi), 4 July 1900 (fl), Baldacci Alb. VII 56 (BM, K). ALBANIA. Patok, Mamuras, 6 m, 15 May 1936 (fl), Pennington 206 (K). MAKEDONUA. Sar Planina, 900- 1 200 m, 9 September 1 968 (fl), Roper 9 ( B M ). Gosti ver to Navrovo, 3 km NE of top of pass, 1 1 00 m, 1 9 July 1 970 (fl), Edmondson 227 (BM). GREECE. Thraki: Thracia occid., in monte Karlik-Dagh prope Komotini (Giimiildschina), 1200m, 2 July 1936(fl),A".//. & F. RechingerlterGraecum IV 10478 (BM). Makhedonia: N. Khalkidiki, Oros Cholomon, c. 8 km WSW of Arnaea, 1100 m, 10 September 1981 (fl & fr), Husain, Jury & Rutherford 99 (BM, RNG*) ['van humile'}. TURKEY. Balikesir: Mt. Ida, prope Kareikos, 3 July 1 883 (ft),Ascherson in Sintenis 474 (BM). INTRODUCTIONS CANADA. Newfoundland:/?^ Scoggan ( 1 978: 1097). Prince Edward Island: Royalty Road, 9 August 1 888 (fl), Macoun s.n. (BM). Nova Scotia: Bridgetown, 5 July 1872 (fl), Fowler s.n. (P). New Brunswick: Rothesay, 1 September 1875 (fl), Mathew s.n. (P). Quebec: Quebec, foot of Heights of Abraham, 13 August 1959 (fr), Stearn 2403 (BM); Co. Sherbrooke, Jouvence, 17 September 1976, Bourassa 43 (H). Ontario: Stanford Township, 10 August 1943 (fl), K.O. Wood s.n. (BM); Ottawa (West), Nurminutly, 15 July 1927, Pesola s.n. (H). Br. Columbia: W. Vancouver, corner 17th Street and Duchess Ave., 9 August 1963 (f\),Bird 9061 (BM); Vancouver I., Cowichan Lake Road, 171 m, 28 June 1967, Turner et al. 552 (H). U.S.A. Maine: Marion Co., Marion, 28 August 1956 (fl), Rees 403 (BM). Vermont: Windham Co., Townshend, 10 July 1937 (fl), E. & H. Moldenke 9893 (BM). New Hampshire: Rockingham Co., just S. of Portsmouth on US Rt. 1, 22 August 1967 (fr), Miles 69436 (BM). Massachusetts: Hampshire Co., Northampton, off Daymen Rd., 19July 1979(fl&fr),AAtes87101 (BM). Rhode I.: Newport Co., Tiverton, Four Corners, 2 July 1 979 (fl), Miles 83 1 82 (H). New York: Tompkins Co., Connecticut Hill, 7 September 1983 (fl), Kearney 39 (BM). New Jersey: Somerset Co., Watchung, 4 July 1934 (fl), Moldenke 8066 (BM, TAI). Pennsylvania: Westmoreland Co., c. 2.5 km S. of Ligonier on PA hwy 71 1, 400 m, 1 July 1979 (fl), Utech 79-327 (BM, W). Maryland: Prince Georges Co., Beltsville, 8 July 1 950 (fl), Petrak 3 1 4 ( W). Map 7 5a. H. perforatum subsp. perforatum, North America (introduced) •, other records O. 94 N.K.B. ROBSON Map 8 5a. H. perforatum subsp. perforatum, South America (introduced) Virginia: Giles Co., just off Va. 700, entrance to Mountain Lake Biol. Stn., 1 146 m, 28 July 1968 (fl & fr), Culwell s.n. (BM). N. Carolina: Avery Co., E. side of Grandfather Mountain, below summit, 1560 m, 21 July 1968 (fl), Culwell s.n. (BM). Tennessee: McMinn Co., Etowah, 20 July 1969 (fr), Rogers 43984 (H). Arkansas: Pope Co., US 64 at Happy Bend Rd., 6.7 km E. of junction with Ark. 105 in Atkins, 120 m, 12 June 1968 (fl), Culwell s.n. (BM). Missouri: Marion Co., near Hannibal, 13 July 1918 (fl), Davis 8969 (H). Illinois: Cork Co., E. from Des Plaines R., S. of Irving Park Rd., 4 July 1956 (fl), Bennett s.n. (W). Indiana: Lake Co., Roby, 12 July 1906 (fl), Lansing 255 1 (W). Ohio: Trumbull Co., Vienna Twp., 25 July 1965 (fl & fr), Sturm 160 (W). Michigan: Barago Co., Ford Forestry Center, 13 July 1974 (fl), Bourdo 28729 (W). Wisconsin: Waukesha Co., Bishop's Woods, 22 June 1970 (fl), Klopatek s.n. (H). Colorado: Boulder Co., S. Boulder Creek 9.6 km S. of Boulder, 27 June 1967 (fl), Weber 13190 (H). Washington: Olympic Peninsula, Deer Park, 390 m, 17 July 1965 (fl), Fisk & Chapman 97 (H). Oregon: Jackson Co., P.O. Prospect, Rogue R. bottoms, Farewell Forestry Camp, 780 m, 28 July 1959 (1. fl), Demaree 41365 (BM). Califor- nia: Trinity Co., New River, Up Grade Mine, 10 August 1950, Wheeler 611 4 (BM, RSA*). Also recorded from Connecticut, Delaware, D.C., S. Carolina, W. Virginia, Kentucky, Texas, Oklahoma, Kansas, Nebraska, Iowa, Minne- sota, N. & S. Dakota, Wyoming, Montana and Idaho (Robson, ined.). MEXICO. Kbg., 1865 (fl), Bilimeck s.n. (MEXU). No precise locality, n.d. (fl), McMinn s.n. (P). CUBA. Matanzas: Matanzas, December 1822 (fl & fr), Poeppig s.n. (BM, P, W). Oriente: Sierra de Nipe, Woodfred, c. 500 m, 24 September 1922 (st), Ekman H.9313 (S). HAITI. Massif de la Selle, Petionville, c. 1400 m, 31 July 1924 (fr), Ekman H.1217 (S, US); above Port au Prince, 1600 m, 1947 (fl), Heldring- Talma s.n. (U). BRAZIL. Rio de Janeiro: Serra do Itataia, Retiro, 1 7 May 1 902 (fl & fr), Dusen 258 (S). URUGUAY. Montevideo: Sayago, 15 February 1936 (fr), RosengurttB 2049 (GH). ARGENTINA. Buenos Aires: Bella Vista, October 1915 (st), Hicken 444 (GH). Chubut: Lago Puelo, 10 February 1955 (fr), Burkhart 19915 (US). CHILE. Talca: Empedrado, 11 March 1972 (fr), Mahu 8710 (H). Conception: Saltos del Laja, 5 January 1971 (fl), Lourteig 2518 (S). Nuble: Termas-Chillan,8-10kmdelasTermas,2000m,7January 1971 (ft), Lourteig 2535 (P, S). Bio Bio: Parlahueque, December 1929 (fl), Piria 197 (GH). Malleco: Manzanar, 13 January 1986 (e. fr), Pedersen 14214 (BM). Cautin: Dpto Victoria, around Termas de Tolhuaca and 1 6 km towards Curacuatin, 950-1 180 m, 12 March 1939 (fr), Morrison & Wagenknecht 17488 (GH, K, MO). Valdivia: La Union Reserva Forestal, January 1 969 (fl), Mahu 9739 (H). Llanquihue: Alerce, 14 February 1936 (fr), Cabrera 3691 (F, S). JUAN FERNANDEZ. Masafuera: Papal, 23 January 1955 (fl), Kuschel 147 (S). HAWAIIAN ISLANDS. Hawaii: North Kona, road up Huaalalni, Kaupulehu, 1500 m, 9 September 1961 (fl & fr), Fosberg 42087 (BISH); Hualalai [Puu Hualalei], 1 July 1964 (fl), Carlson s.n. (BISH). Subsp. perforatum typically has broad, relatively large leaves and large flowers and fruits with narrow linear oblique vittae. In dry habitats (e.g. chalk soils, walls), however, narrow-leaved smaller- flowered plants superficially resembling subsp. veronense are often seen; but these have lateral capsule vittae typical of subsp. perforatum or with the distal ends forming a line of vesicles, not irregular like those of Mediterranean forms of subsp. veronense. Such plants occur in England (and probably elsewhere in NW Europe) as well as in the U.S.A. and Canada, the true subsp. veronense being absent from these areas. Where the distributions of subsp. perforatum and subsp. veronense overlap (in southern France, southern Germany, Austria and northern Italy), forms intermediate between the subspe- cies occur. These are similar to the northern dry-habitat forms in having narrow petiolate leaves, but the capsule glands are also intermediate in form, i.e. ± irregularly vesiculate but with several dorsal linear vittae. Examples of such intermediate specimens from Austria (Styria) were cited by Frohlich (1911) as subsp. angustifolium, e.g. Foiling bei Maria Trost, 23 July 1910 (fr), Frohlich GZU 16326 (GZU). There is evidence that such popul- ations show cytological irregularities (Garbari, in litt.) STUDIES IN HYPERICUM Map 9 5b. H. perforatum subsp. songaricum •. It seems likely that subsp. perforatum was introduced into North America independently in the east and the west. A low, erect to procumbent southern Balkan plant with small, rather broad leaves (1 : b= 1-2.5) and capsule valves bearing lines of vesicles has been named H. perforatum var. humile Stranski, but no valid description has been traced. On account of intermediate forms that link it with typical subsp. perforatum, it apparently belongs to that subspecies. Specimens have been seen from Albania, Bulgaria and Greece. 5b. Hypericum perforatum subsp. songaricum (Ledeb. ex Rchb.) N. Robson, stat. nov. Fig. 8, Map 9. H. perforatum var. [f3] latiglandulosum Choisy in DC., Prodr. 1: 550 (1824). Type: Ukraine, Tanais [R. Don], 1819 (fl), Goldbach s.n. (G-DC!-holotype). H. songaricum Ledeb. ex Rchb., Icon. hot. pi. crit. 3: 72, t. 283 f. 446 (1825); Spreng., Syst. veg. 4(2): 297 (1827). Type: Kazakhstan, 'ad lacum Noor-Saisan dictum', fl., Ledebours.n. (LE-holotype). H. perforatum var. songaricum (Ledeb. ex Rchb.) K. Koch in Linnaea 15: 714 (1841); Woron. in Kuzn., Woron. & Fomin, Fl. Cauc. crit. 3(9): 56 (1906). H. perforatum var. gracile Gruner in Bull. Soc. Imp. Naturalistes Moscou41(3): 132 (.869). Type: Ukraine, Zaporozh'ye, 'Gubern. Catherinoslav, in demissis ad Borysthenem infra urbem Alexandrowsk' [Zaporozh'ye], 20 June 1865 (fl), Gruner s.n. (BM!-holotype?). H. komarovii Gorschk. in Bot. turn. SSSR 24: 428, f. 1 (1939). Type: Kirghizstan [Kirghizia], in monte Arslanbob supra Namas-tasch, 23 August 1925 (fl), Massagetov 335 (LE-holotype). Icon: Rchb., Icon. hot. pi. crit. 3: t. 283 f. 446 (1824). Leaves sessile (at least on main stems); lamina 6-15 x 2-10 mm, oblong to oblong-ovate (l:b=2.5-4), base ± shallowly cordate- amplexicaul, often glaucous beneath. Inflorescence not congested, with branches relatively short, straight. Petals with laminar glands all pale. Capsule valves with lateral vittae linear, narrow or slightly swollen but rarely interrupted. Northern Kazakhstan and adjacent Kyrgyzstan and China (NW Xinjiang), south-eastern Russia and the Ukraine (including Krym). CHINA. Xinjiang: Altay Xian, 1 100 m, 18 August 1956 (fl & fr), Ching 2434 (CDBI, IBSC); Huocheng [Suiding] Xian, 1500 m, 22 June 1965 (fl), Cheo et al. 650862 (IBSC, KUN, NAS). KAZAKHSTAN. Almatinskaya Obi.: Songaria, in dumetis montium Alatau, n.d. (fl), Schrenk s.n. (K, LE*); Semiryechensk prov., Vyernyi distr., Zaylinskiy (Trans-Ilian) Alatau, 14 July 1907 (fl), Sokalsky 118 (BM). Zapadnyy-Kazakhstan Obi.: Chingirlauskiy rayon, Kara-Alag, 16 July 1935 (e. fr), Akastovy(l) s.n. (A*, BM, LE). KYRGYZSTAN. *Osh: in monte Arslanbob [Pik Skobeleva] supra Namas-tasch, 23 August 1925 (fl), Massagetov 335 (LE). RUSSIA. Astrakhanskaya Obi.: Sarepta, 16 June 1879 (fl), Becker s.n. (BM). Stavropolskiy Kray: S. of Karachayevsk, near Gora Dembay Ul'gen, 11 July 1971 (fl), Fraser Jenkins 3219 (BM). UKRAINE. Orlovskaya Obi.: Prov. Orel, circut. Felitz, inter Polna et pag. Morskaja, 20 June 1868 (bud), Gruner s.n. (BM). Dnipropetrovskaya Obi.: Gubern. Catherinoslav [Dnipropetrovs'k], in demissis ad Borysthenum [Dnieper] infra urbem Alexandrowsk [Zaporozhye], 20 June 1865 (fl), Gruner s.n. (BM). Krym: prope Karasubazar [Belogorsk, Bilohirs'k], 23 June 1 900 (fl), Callier 564 (BM, FR, JE, K); in aprices Tauriae montosae, n.d. (fl), Pallas s.n. (BM). Some Crimean specimens lack the cordate leaf base of this subspe- cies but are otherwise clearly a reduced form of it. The following specimens are intermediate between subsp. songaricum and subsp. perforatum in having the leaves on the main stem sessile but those on branches petiolulate: UKRAINE. Cherkasky Obi.: Uman, Farco, June 1867 (fl), Golde s.n. (BM, K). ? Obi.: Valka, zwischen Veselaya und Zingalni (?), 5 May 1 865 (fl), 96 N.K.B. ROBSON Grune r s.n. (BM); an Ackerrandran (?) bei Condo-dorf, 1 8 July 1 859 (fl), Gruner s.n. (BM). The following specimen is intermediate between subsp. songaricum and subsp. veronense in having narrow leaves with those on the main stem cordate at the base. CHINA. Xinjiang: [Kashi Xian] Yarkand Expedition, 1 870 (fl), Henderson s.n. (K). The Fraser-Jenkins specimen from the Russian Caucasus listed above also shows some characters intermediate between those of subspecies songaricum and veronense. 5c. Hypericum perforatum subsp. veronense (Schrank) H. Lindb. in Ofvers. Forh. Finska Vetensk.-Soc. 48: 73 (1906); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 530, f. 5 (191 1), in Mitt. Naturwiss. Vereins Steiermark 51: 241 (1915); Hayek, Prodr. Fl. Pen. bale. 1: 533 (1925); Stjep.-Vesel. in Josifovic, Fl. Srbije 3: 1 19 (1972); Robson in Bull. Soc. Ech. Vase. Eur. Bass. Medit. 27: 20 (1998), Ciccarelli, Garbari & Robson, (2002). Type: as for H. veronense Schrank. Note: Lindberg's '//. perforatum L. *Veronense (Schrank)' is to be regarded as a new combination at subspecies rank. He indicated new combinations at varietal rank in this paper by an initial small letter. Fig. 8, Maps 10-12. 1 Hypericum perforatum [var.] angustifolium Borkh. in Neues Mag. Bot. 1: 26 (1794); N. Robson in Wisskirchen & Haeupler, Standardliste Farn- u. Blutenpfl. Deutschl.: 270 (1998). Type: Germany, not stated. The varietal epithet angustifolium has previ- ously always been attributed to De Candolle, but Borkhausen's H. perforatum angustifolium has priority. By the date of his publica- tion, a trinomial is assumed to be a variety. Borkhausen's type, however, is unknown (Stafleu & Mennega, 1993: 340); and so whether his plant belongs to the narrow-leaved Mediterranean form with vesicular capsule glands (var. angustifolium DC.) or a narrow-leaved form of var. perforatum (with vittate, not vesicular capsule valves) cannot be ascertained. In either case De Candolle's var. angustifolium is a later homonym. However, as the variation between narrow-leaved plants with large and small leaves in the Mediterranean region is continuous, allowing the recognition of no more than one taxon, De Candolle's epithet microphyllum is available for it as a variety. H. veronense Schrank in Neues Bot. Taschenb. Anfdnger Wiss. Apothekerkunst 22: 95 ( 1 8 1 1 ); Link, Enum. hort. berol. alt. 2: 276 ( 1 82 1 ); Jord. in Schultz & Billot, Annots Fl. France & Allemagne: 49 ( 1 855); Engl. in Verh. hot. Vereins Brandenburg 12: 49 ( 1 870); O. Schwarz in Drudea 5: 61 (1965). Type: Italy, Verona, an den ehemaligen Sitzen des Amphiteaters zu Verona, October 1808 (fr), Schrank (not found). Neither Professor Garbari (Univ. of Pisa; in litt.) nor I have been able to locate type material of this 'species', which, according to Prof. Garbari, does not grow now at the type locality. (Schrank himself refers to strenuous efforts to extirpate it, even at that time.) There is no doubt, however, that Schrank's plants belonged to the narrow-leaved Mediterranean subspecies, even though the plants that he grew from their seeds were somewhat larger with leaves (some or all?) shortly petiolate. In the absence of type material, a neotype is therefore required: Italy, Verona, Anfiteatro di Verona, August 1874, Cesati s.n. (RO!-neotype, Ciccarelli, Garbari & Robson, 2002, ined.). H. perforatum var. [5] angustifolium DC. in Lam. & DC., Fl. France, 3rd ed. 5: 630 ( 1 8 1 5); Gaudin, Fl. helv. 4: 628 ( 1 829); W. Koch, Syn.fl. germ, helv.: 134 (1835), in Rohling, Deutschl. FL, 3rd ed. 5: 349 (1839); Beck, Fl. Nieder-Osterreich: 530 (1892); Rouy in Rouy & Fouc., Fl. France 3: 333 (1896); Zelen., Prodr. fl. Taur.: 233 (1906); Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925); Briquet, Prodr. fl. Corse 2(2): 151 (1936); Gus,ul. & Nyar. in Savul., Fl. R. P. Roman. 4: 29, t. 2 f. 2 ( 1 956); N. Robson in Tutin et al., Fl. Europaea 2: 269 (1968), in Cullen et al., Eur. gard.fl. 4: 60 (1995); Jordanov & Koz. in Jordanov, Fl. R. P. Bulg. 4: 261 (1970). Type: France, Pyrenees Orientales, Roussillon, Prades, 1814 (fl), Coder s.n. (G-DC-holotype, microfiche!). H. perforatum var. [E] microphyllum DC. in Lam. & DC., Fl. France, 3rd ed. 5: 630 (1815); Choisy, Prodr. monogr. fam. Hyperic. : 5 1 ( 1 82 1 ). in DC., Prodr. 1: 550 ( 1 824); Guss., Fl. sicul. syn. 2(1): 378 (1843); Rouy in Rouy & Fouc., Fl. France 3: 333 (1896); Fiori, Nuov. Fl. Italia 1: 522 (1924); Briquet, Prodr. fl. Corse 2(2): 151 ( 1 936); Gu§ul.& Nyar. in Savul., Fl. R. P. Roman. 4: 30 (1956); N. Robson in Tutin et al., Fl. Europaea 2: 269 ( 1 968), in Cullen et al., Eur. gard. fl. 4: 60 ( 1 995), in Wisskirchen & Haeupler, Standardliste Farn- u. Blutenpfl. Deutschl.: 270 (1998); Jordanov & Koz. in Jordanov, Fl. R. P. Bulg. 4: 261, t. 49 f. 3 (1970). Type: France, Lozere, aux environs de Mende, 1810 (fl), Prost s.n. (G-DC-holotype, microfiche!). H. perforatum var. [y] sensu Choisy, Prodr. monogr. fam. Hyperic. : 50(1821). H. perforatum var. [y] elatum Choisy in DC., Prodr. 1: 550 (1824), in Zoll., Syst. Verz. 1-2: 151 (1854). Type: Italy, Florence, 10 August 1808 (fl), De Candolle s.n. (G-DC!-lectotype, selected here). H. stenophyllum Opiz, Naturalientausch no. 9: 158(1 825). Syntypes: Czech Republic, 'in Boheim', Opiz s.n. (PR-syntype); Austria, 'in Oberosterreich', Brittinger s.n. (PR-syntype). A specimen from the Czech Republic, Hodkovicka [Prague], 25 July 1850 (fr), Opiz (PR!), labelled 'Hypericum stenophyllum Opiz' by Opiz himself, is subsp. veronense and, in the apparent absence of the syntypes, can be taken as a representative specimen of his species. H. perforatum var. nanum Gaudin, Fl. helv. 4: 627 (1829). Type: not stated. 'An H. perforatum (3 microphyllum Dec. Fl. Fr. Suppl. p. 630?' (LAU-holotype). H. perforatum var. veronense (Schrank) Ces. in Cattaneo, Not. Nat. Civ. Lombardia: 291 (1844); Beck, Fl. Nieder-Osterreich 2: 530 (1892); R. Keller ex Albov in Trudy Tiflissk. Bot. Sada 1: 4 (1895); Beguinot in Fiori, Beguinot & Pampini, Flora Italica Exsiccata, no. 817 (1904); Woron. in Kuzn., Busch & Fomin, Fl. Cauc. Crit. 3(9): 56 (1906); Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925); Grossg., Fl. Kavk., 2nd ed. 6: 54, 175 (1962). Soldano (1991: 248) claimed that Cesati made a comb. nov. at the rank of subspecies; but there would seem to be no justification for inter- preting 'Hypericum perforatum L. (3 veronense Schrnk.' as other than a variety. He certainly did not make the combination H. perforatum subsp. angustifolium (DC.) Cesati attributed to him by Gamisans & Jeanmonod (1993: 1 83). See Robson & Lambinon (2000). Beguinot's combination was made independently. H. perforatum var. stenophyllum (Opiz) Wimm. & Grab., Fl. Siles. 2(2): 82 (1829); Neilr., Fl. Nieder-Oesterr.: 826 (1859). H. schlosseri Heuff. in Flora 36: 626 ( 1 853); Schlosser & Vokut., Fl. croat.: 381 (1869); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 537 (1911) in adnot. Type: Croatia, in rupibus Zagoniae, Schlosser s.n. (W?). H. microphyllum Jord. in F. Schulz, Arch. fl. France Allem.: 341 (1854). Type: France, Rhone, environs de Lyon, Jordan s.n. (LY- holotype; BM!-isotype). H. chrysostictum Webb, Fragm. fl. aethiop.-aegypt.: 54 (1854). Type: [Saudi Arabia?; not recorded from Ethiopia] 'Occurrit in herb. Figariano sine loci natalis indicatione.' (FI!-holotype). STUDIES IN HYPERICUM 97 •~ • ••• r-t »% V» > '• •^TWW11^'.*. ; *v* •«* *•«%•££ ^_v%. :^ r* *: r Wrfflfcr* Map 10 5c. H. perforatum subsp. veronense, Mediterranean and S. Europe •, other records O , introductions A. Reported from south Germany. H. aetnaeum Tornab., Fl. sicul.: 171 (1887), Fl aetnea 1: 213 (1889). Type: Sicily, Mt. Etna, 'in elatis arenosis et ad nemora', Nicolosi s.n. (CAT-holotype). H. noeanum Boiss., Fl. orient., Suppl.: 130 (1888). Type: Bulgaria, Rumelia, 'Hab. in monte Haemo minore Thraciae' [Iztochni Rodopi], Noe s.n. (G!-holotype). H. perforatum var. [y] angustifolium subvar. lineolatum Rouy in Rouy & Fouc., Fl. France 3: 333 (1896). Type: France, Cher, Mehun, Sables de Beauvoir, 3 July 1 840 (fl), Deseglise in Billot exsicc. no. 3349 (P?-holotype; BM!, JE!-isotypes). H. perforatum var. [8] mediterraneanum Rouy in Rouy & Fouc., Fl. France 3: 333 (1896). Type: France?, not stated. H. perforatum var. typicum forma microphyllum (DC.) Fiori in Fiori & Paol., Fl. Italia 1: 388 (1898). H. perforatum var. ellipticum Freyn in Bull. Herb. Boissier II, 3: 1064 (1903), non Durand & Pittier (1881). Type: Turkmenistan, Aschabad, in monte Ak-tepe, 17 June 1900 (1. fl), Sintenis 564a (G-lectotype, selected here; BM !, K!-isolectotypes); prope Suluklu ad fines Persiae, 4 September 1900 (1. fl), Sintenis 564b (G- syntype; JE!). H. perforatum var. moesicum Velen. in Sitzungsber. Ko'nigl. Bohm. Ges. Wiss. Prag 1903(28): 2 (1904), op. cit. 1910(8): 3 (191 1); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 538 (1911) in synon. Type: Bulgaria, in Sredna Gora ad Adzar [Svezhen], 1902 (fl), Stfibrny s.n. (W?). H. perforatum var. collinum Woron. in Kuzn., Busch & Fomin, Fl. Cauc. Crit. 3(9): 55 (1906); Grossg., FL Kavk. 2nded. 6: 55, 175 (1982). Type: Georgia, Sukhumi, no sklonam' gori Cherniyav- skago, June 1904, Woronow s.n. (LE-lectotype, selected here). H. perforatum var. corioides Vokut., nomen in scheda (WU); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 538 (1911), in synon. H. perforatum subsp. angustifolium (DC.) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 534, f. 6 (1911), in Mitt. Naturwiss. Vereins Steiermark 51: 241 (1915); Hayek, Prodr. Fl. Pen. bale. 1: 533 (1925); Stjep.-Vesel. in Fl. Srbije 3: 1 1 9 (1972); A. Ramos in Castroviejo et al., Fl. Iberica 3: 165, t. 45 ff. a-e (1989). H. perforatum var. floribun dum Sennen, ined.? Type': Spain, Bar- celona, S. Celloni, 20 July 1922 (fl), Sennen 4864 (BC, BM!). H. nachitschevanicum Grossh. in Izv. Azerbajdzansk. Fit., no. 1: 46 (1941), Fl. Kavk., 2nd ed. 6: 171, karta 195 (1962). Type: Azerbaijan, Nakhitchevan, inter pp. Urmys et Pazmara, 21 Au- gust 1933 (fl & fr), Karjagin & Isaev s.n. (BAK-holotype; LE!-isotype; BM!, K!-photographs). H. perforatum var. longicapsulum Jordanov & Koz. in Jordanov, Fl. R. P. Bulg. 4: 261, 709, t. 49 f. 4 (1970) ' longicapsula' . Type: Bulgaria, Mt. Vitosa [Cherni Vrukh], in pratis prope refugium 'Pogledec', loco dicto 'Karkama', 1600 m, 18 September 1967 (fr), N. Vihodcevsky (SO-holotype). Icones: Valdes, Talavera & Galiano, Fl. Andalucia Occid. 1: 315 (1987); Castroviejo et al., Fl. Iberica 3: t. 45, ff. a-e (1993). Leaves usually sessile (at least on main stems); lamina 9-20 x 1-4 mm, usually linear (l:b=5-9) but sometimes narrowly triangular- lanceolate or linear-oblong (l:b=2.5^), or occasionally broadly ovate to elliptic or obovate, but then small (c. 5-10 x 4-5 mm), base cuneate, paler but not glaucous beneath. Inflorescence occasionally congested, with branches relatively short, straight or curved-ascend- ing. Petals with laminar glands all pale or rarely mostly black. Capsule valves with lateral vittae swollen at base (but not forming a regular distal row) or ± interrupted to punctiform and wholly swollen (vesicular). Southern Russia, the Ukraine (Krym) and the Caucasus region east to Kazakhstan and N. India (Uttar Pradesh) and west to the Mediter- 98 N.K.B. ROBSON ranean region and southern Europe (N. to central France, southern Germany and southern Poland) and extending to Macaronesia, north-west Africa, Sudan Republic (Jebel Marra) and Saudi Arabia (Asir). Introduced into Hawaii, New Zealand, Australia and South Africa. Records of var. angustifolium from Canada and the U.S.A. would appear to refer to the narrow-leaved form of subsp. perforatum found in dry habitats in England (see p. 94). The record of H. perforatum from Karpathos (Turland, Chilton & Press, 1993: 94) is an error for H. tetrapterum. RUSSIA. Ciscaucasia: Stavropolskiy Kray, N. of Pyatigorsk, Mt Mashuk, 18 November 1975 (fl & fr). O. & J. Degener 33,789 (NY); Karachayevo- Cherkesskaya ASSR, sudlich Teberda, c. 1350 m, 11 August 1957 (fr), Meyer, Lippold & Kohler251 (JE); Severo-Osetinskaya ASSR, Vladikavkas [Ordzhonikidze], August 1981 (fl & fr), A. & V. Brotherus 192 (BM, H). UKRAINE. Krym: Yalta, Massandra, 22 June 1969 (fl), Rintamn s.n. (H). GEORGIA. Abkhazskaya ASSR: Pizunda [Pitsunda], 19 July 1966(fr), F.K. & J. Meyer 8560 (JE). Gruziya: prope Tiflis [Tbilisi], June 1981 (fl), A.M. & V.F. Brotherus 192a (BM, H); Batumi, ostlich Machindzauri, 25 July 1978 (fr), H. & R. Manitz s.n. (JE). AZERBAYDZHAN. Nachrespublica, inter Urmys et Paxmara, 21 Au- gust 1933 (fr), Karjagin & Isaev s.n. (BM and K photographs, LE); Chizy distr. (olim Kuba), prope p. Alty-agatsh, 21 June 1935 (fl), Gurvitsh s.n. (K); Nakhichevan - see type of H. nachitschevanicum Grossh. in synonymy. ARMENIA. In monies 'Bazumski khrebet' in vicinitate oppidi Kirovakan (Karalkisa), 1600-1800 m, 22 July 1975 (fl), Vasdk s.n. (W); Sevanskiy raion, Goranrotuv n.-o. Sevan, 6 July 1926 (fr), Savur s.n. (LE); Kafanskiy raion, gor. Kadzharan, Tandzasar, 1850-2150 m, 16 September 1966 (fr), Manakian s.n. (BM). TURKEY. Kars: Ardahan, Giircayir village, 31 July 1976 (fl), Sanda 1196 (BM). Bitlis: Corgih to Bitlis, 2250 m, 31 July 1976, Tong 207 (E). Hakkari: Cilo Dag, in Diz deresi above Kursi, 1890 m, 7 August 1954 (fr), Davis & Polunin D. 23962 (BM, E, K). Coruh: Borsa to Hopa, 300 m, 22 June 1957 (fl), Davis & Hedge D. 29882 (BM, E, K). Erzurum: Palandokea Dag, 25 km from Cat to Erzurum, 2350 m, 27 July 1966 (fl), Davis 47385 (E). Trabzon: Hohe sudlich Trabzon, 250 m, 26 June 1931 Go'rz 654 (BM). Amasya: Erbaa to Kozlu, 550 m, 5 June 1 967 (fl), Tobey 2 1 90 (E). Malatya: 5 km NW Darende, 1200 m, 8 September 1977 (FL), Sorger 77-105-22 (WU). Hatay: Hassa, Tiiyek village, 9 July 1974 (fl), E. Sezik 680 (BM). Nevs.eb.ir: Goreme, c. 10 km W. of Urgup, c. 1000 m, 3 August 1956 (fl), NcNeill 399 (E, K). Ic.el: Bulgar Dagh, 'Gullek Boghas', 1 140 m, 23 August 1853 (fl & fr), Kotschy 282a (G). Kastamonu: Daday to Azdavay, 30 km, 1 100 m, 30 July 1962 (fl & fr), Coode & Yaltirik D. 38658 (E, K). Ankara: Elma Dagh bei Ankara, c. 1600m, 25 June 1932 (fl),Kotte 3 17 (K). Antalya: Manavgat, 4 June 1973 (fl), Himmetoglu H17 (BM, BAS*). Bolu: Ala Dag distr., Kartal Kaya (Mt.), 2000 m, 11 August 1960 (fl), Khan, Prance & Ratcliffe 490 (E). Eski§ehir: Eskishehir, 500 m, 27 May 1971 (fl), Price 638 (K). Isparta: Egridir distr., Barla Dag foothills, 1200 m, 1 August 1960 (fl), Khan, Prance & Ratcliffe 4219 (E, K). Istanbul: Alaton, 28 July 1967 (fl), Baytop & Atila 1 1585 (E, ISTE*). Kiitahya: Gediz, 850 m, 7 July 1962 (fl), Davis & Coode D. 36969 (E, K). Burdur: Yesjlova to Denizli, c. 6.5 km, by Salda Golu, 1100 m, 1 June 1962 (fl), Dudley D. 35303 (E, K). Kirklareli: Demirkoy, 1 9 July 1959 (fl), Baytop 5488 (BM). Izmir: Sel?uk, Ephesus, 3 1 May 1972(fl),£. <£G.Sez//t260(BM,HUB*).Mugla:Marmaris,c.4kmE. of Marmaris, s.l., 18 May 1983 (f\),Murto 1376 (H). GREECE. Thasos: Glifada, 18 May 1986 (fl), Raitraluna? s.n. (C). Thraki: Nom. Xanthis, NE of village Dimarion, c. 26 km N. of Xanthi, 550 m, 28 July 1977 (fr), Strid & Georgiadhou 13485 (C). Makedhonia (E): [Nom. Kavala] c. 4 km W. of Kavalla, 1 1-12 July 1970 (fr), K.H. Rechinger 38260 (W). Makedhonia (W): Nom. & Ep. Grevenon, Mt. Smolikas, 2-3 km S. of Samarina towards Armata, 1275- 1350m, 27 August \915(f\),Hartwig 6 Seberg 4814 (C). Epirus: Distr. Janina, ad Paleochori Syrakou, 2 August 1895 (e. fr), Baldacci III 304 (BM). Ionian Is.: Korfu, S. of Korfu town, Kanoni. 20 May 1978 (bud), A. Hansen 441 (C). Thessalia: prov. Trikala, distr. Kalambaka, Meteora, between Ajia Triada and St. Roussanis, 600 m, 17 June 1977 (fl), Steam A.64 (BM). Sterea Ellas: Gravia, 480 m, 20 June 1937 (fl), Balls & Gourlay B. 3352 (BM). Peloponnisos: Prov. Achaia, Distr. Kalavrita, Mt. Helmos, lower SE slopes, 3 km W. of village Feneos, 1000- 1200 m, 30 July 1980 (fr), Baden & Franze'n 684 (C). Evvoia: Mt. Dirphys, c. 1930 (fl), Guiol s.n. (BM). Sporadhes: Rhodes, S. of Kremasti, 9 May 1966 (fl), A. Hansen s.n. (C). KRITI. Distr. Kissamos, Enneachoria, inter Myli et Elos, c. 100 m, 30 May 1942 (fl), K.H. Rechinger Iter Graecum VI 13437 (BM, K). ALBANIA. Nordalbanische Alpen (Prokletija), Tethi, Shtegu i Dhenve, c. 1400-1600 m, 26 September 1961 (fl & fr), Meyer 6427 (BM, JE); Korea, Mali i Moraves, bei Drenova, 1 100-1200 m, 12 September 1962 (fr), Meyer 6141 (BM, JE); Viossa-Tal, an der Landstrasse nordlich Perati, c. 300 m, 10 July 1959 (fr), Meyer 3835 (JE). MAKEDONUA. Gorge of Crni Drim near Lukovo, N. of Struga, 650 m, 12 June 1971 (fl), Chater 432 (BM). MONTENEGRO. Distr. Vasojavici, Murino, 1 September 1903 (fl & e. fr), Baldacci Iter Alb. X 46 (BM); 1 0 km S. of Titograd [Podgorida], 50 m, 20 June 1965 (fl & e. fr), Wrigley 65/1360 (K). BOSNIA & HERZOGOVINA. Velez planina, 1 500 m, 6 July 1912 (fl), Sagorski s.n. (BM); Jajce, 1925 (fl), Leathes s.n. (BM); Sarajevo, 750 m, 12 July 1960 (fl), McCallum Webster 4036 (K). SERBIA. Kosovo, c. 10 km S. of Pristina towards UroSevac, 600 m, 9 September 1972 (fr), Kukkonen 8565 (H); Savac-Pomoravlje, Damin, 13 September 1 979 (fl), Cvetkovic 1 5 (BM) ; Vranja, 1 9 1 9 (fl & fr), Tin dall Lucas 104 (BM). CROATIA. Velebit Mts, pass above Karlobag, 690 m, 17 July 1962 (fl), Mathew & Pycraft 339 (BM); Insel Krk, Omisaly, 17 July 1931 (fl & fr), HuteH s.n. (BM). SLOVENIA. Istrien, zwischen Tulisevica und Draga di Lovrana, 23 June 1935 (fl), Ronniger s.n. (W); within 9.5 km of Bled, Mt. Triglav, 300-1200 m, 1923 (fl), Leathes s.n. (BM). BULGARIA. Mikhailovgrad: Predbalkan, Ostabhang des Midshur [Midzor], 4 September 1958 (fr), Meyer 1171 (JE). Sofiya: Mt. Vitosha, c. 1400 m, 26 July 1952, Vihodcevsky s.n. (JE). Blagoevgrad: Struma-Tal, bei Marikostinovo, 23 September 1958 (fr), Meyer 1500 (JE). Plovdiv: in valle Akdere [Tundzha] prope Kalofer, 1 1 August 1893 (fr), Wagner 26 (BM, W*). ROMANIA. Transylvania: Cluj, Turda Gorge, 23 July 1963 (fl), Dandy 1425 (BM). Dobrogea: Constanja, Tekirg Cirol [Techirghiol], n.d. (fl), Cretzoiu s.n. (K). HUNGARY. Pest: Pass between Doroy and Pilisovorosvar on Budapest road, 7 July 1962 (fl), Curie 44 (BM). SLOVAKIA. Vrchol Sitna, c. 1000 m, 1957, Hosticka & Zeleny (PRC*), only locality (fide Zeleny, Jasickova & Zahradnikova, 1982: 310). CZECH REPUBLIC. Possibly occurs (fide Celakovsky, 1875: 521). POLAND. WaHirzych: Silesia, Schweidnitz [Swidnica], Koltschenberg, September 1882 (1. fr), Peck s.n. (FR). GERMANY. Thiiringen: Gera, Jena, Lichtenhain am Bahndamm, 20 June 1950 (fl), Launert 269 (BM) (verg. ad subsp. perforatum?). Baden- Wurttemburg: Istein in Baden, July 1919 (fl & fr), Meebold s.n. (K). AUSTRIA. Salzburg: Leopoldskroner-Moor, 490 m, June 1 877 (fl?), Eysn s.n. (JE). Steiermark: bei Kelsdorf (Graz), 2 August 1910 (fl & fr), Frohlich GZU 1 6323 (GZU). Karnten: Carinthia, fluv. Gail, July 1 864? (fl), Pichlers.n. (JE, K). Tirol (E.): prope Lienz, July 1 863 (fl & fr),Ausserdorfer s.n. (BM, K). SWITZERLAND. Zurich: Albis, 3 August 1854 (fl), Hort s.n. (BM). Vaud: Des Pierrettes, pres Lausanne, 7 September 1877, Ferrat s.n. (JE). Geneve: Compesieres, August 1855 (fl), Chavin & Perrier55 (BM). Ticino: [Monte] Generoso, Bela Vista, 10 August 1899, Gyoperzer s.n. (JE). ITALY. Piemonte: Liguria, Portofmo Peninsula, 10 September 1963 (fr), Robson 1813 (BM). Lombardia: Brescia, Lago d'Iseo between Velio and Toline, 195 m, 2 June 1983 (fl), Hind 165 (BM, RNG*). Venezia: Verona, Torri del Benaco, 100-200 m, 15 July 1904 (fl), Rigo in Fl. Ital. Exsicc. 817 (BM). Campania: Ravello, above Sta Cattarina, 2 October 1 879 (st), Lacaita s.n.(BM).Puglia:Brindisi,8kmW.oftown,31 May 1 960 (fl), Brummitt 60. E. 802 (K, LIV*). Calabria: Aspromonte, c. 50 km SW of Reggio di Calabria, below Bova, c. 750 m, 10 June 1979 (fl), Davis & Sutton D. 64779 (BM). SICILY. Palermo: c. 5 km W. of Cefalu, 30 m, 25 May 1979 (fl), Davis & Sutton D. 63726 (BM). Monreale: Weg nach San Martino, 27 May 1927 (fl), Ronniger s.n. (W). Messina: Taormina, 28 May 1964 (fl), Brummitt 4567 (K). Pantellaria: P. Spadillo, 27 May 1960 (fl), Davies s.n. (K). MALTA. Malta: Kordin (Haslam, Sell & Wolseley, 1977); occurrence doubtful or plant extinct (Lanfranco, 1982). SARDINIA. Sassari: near Stintino, La Salina, 5 June 1971 (fl), Dunford & Moves 143 (BM). STUDIES IN HYPERICUM 99 CORSICA. Haute-Corse: Bastia, oberhalb der Stadt, 1 June 1914 (fl), Ronniger s.n. (W); Cirque de Bonifato, pres de la Maison forestiere, c. 530 m, 8 July 1 999 (fl), Lambinon 99/Co.276 (BM, LG*). Corse-du-Sud: Zouya to Porto Vecchio, 21 April 1933 (fl), A.C. & E. Godman 127 (BM). FRANCE. Vendee: Mortagne, 24 June 1869 (fl), Genevier s.n. (BM). Deux Sevres: Puy St. Bonnet, 26 June 1861 (fl), Genevier s.n. (BM). Charante-mar.: Autran, July 1884, Giraudias s.n. (JE). Cher: Allogny, 12 July 1878 (fl & fr), A.Z.P. s.n. (BM). Saone-et-Loire: Rougers, 4 August 1891 (fl & fr), Gillot s.n. (BM). Hte-Saone: Goren, pres Conflans, 14 September 1858 (fr), Perrier s.n. (BM). Rhone: Arnas, August 1897 (fr), Gandoger s.n. (H). Isere: Viray, vallee du Vezy, August 1910 (fl & e. fr), Giraudias s.n. (BM). Puy-de-D6me: Volvic, 750 m, 18 September 1893 (fr), Heribaud in Soc. Rochel. 3438 (BM). Cantal: Anglards de Salers, July 1928 (fl), Chanar s.n. (BM). Hte Savoie: Passy, coteaux au Reyard, 1 1 July 18 — (fl), Depierre s.n. (BM). Savoie: Verel-Pragondran, pres de Chambe'ry, 19 August 1852 (fl), Chabert in Billot 1847 (BM, JE). Htes-Alpes: environs de Gap, July 1 854 (fl), Blanc in Billot 1 846 (BM). Alpes-mar.: Cipieres village, 700 m, 29 June 1992 (fl), Hepper943\ (BM, K*). Drome: Donzere, barrage between Canal and R. Rhone, 17 June 1969 (fl), Verdcourt 2682 (K). Var: c. 2 km NW of Cotignac, 350 m, 5 August 1981 (fl), Longton 4443 (BM). Vaucluse: Entrechaux a Faucon, 23 June 1981 (fl), Bamps 6960 (BM, BR*). Bouches-du-Rhone: Camargue, Canal de Fumemort, 2 km from Tour du Valat, 10 June 1968 (fl), Kendrick & Moyes 206 (BM). Card: Saint-Christol- les-Ales (S. d'Ales), c. 170 m, 13 July 2000 (fr), Lambinon O/F/246 (BM, LG*). Ardeche: St. Jean de Pourcharesse, 600 m, 3 September 1984 (fl & e. fr), Dechamps 10013 (BM, BR*). Aveyron: Cherondels, 15 July 1905 (fl), Carbonel s.n. (BM). Lot: Causse d'Alvignac, July 1868 (fr), Mah— s.n. (BM). Landes: near Dax, June 1924 (fl), Heard s.n. (BM). Aude: Moussan, 28 May 1903 (fl), Septimin-Donat s.n. (BM). Pyrenees-or.: Lorede, 21 June 1906(fl), Conills.n. (FR). BALEARIC ISLANDS. Mallorca: Alcudia, Mai Pas beach, 9 August 1967 (fr), J. & M. Cannon 3366 (BM). Menorca: San Luis, E. of Punta Prima, c. 10 m, 14 April 1967 (o. fr), Bowden & Sims 1180 (BM). Ibiza: San Juan Bautista, between Puerto de San Miguel and San Miguel, c. 30 m, 16 January 1971 (st), Ferguson 2113 (BM). SPAIN. Gerona: N. of Puerto de la Salva, c. 20 m, 17 June 1959 (fl), Chamberlain 8 1 5 (BM). Barcelona: San Celoni, 20 July 1 922 (fl), Secondaire in Sennen, Pis d'Espagne 4864 (BM, RNG). Huesca: Sierra de Guara, between Casa de Fabara and Fabara Gorge, 15 August 1970 (fr), I. & E Richardson 21 (RNG). Viscaya: Ordufia, emirna del Balneariode la Nueva, 10 July 1941 (fl), Guinea 1555 (RNG). Leon: near Cistierna, 6 July 1927(fl), Wilmott s.n. (BM). Madrid: Guadalupe, 14 June 1958 (fl), Guinea s.n. (RNG). Ciudad Real: 3 km SW of Alhambra, Sierra de Alhambra, 950 m, 29 June 1974 (1. fl), Leadley & Petty 172 (BM, RNG). Albacete: Almansa, W. end of Sierra de El Mugr6n, 1 150 m, 22 June 1979 (fl), Reading Univ. Bot. Dept. Exped. 1 50 (BM, RNG). Valencia: Utiel, Utiel to Pantano de General- issimo, 21 km N. of town, 641 m, 20 June 1979 (bud), Reading Univ. Bot. Dept. Exped. 55 (BM, RNG). Almeria: Laujar de Andarax, Nacimiento, c. 900 m, 8 July 1978 (fl), Roivainen s.n. (H*, RNG). Jaen: Sierra de Cazorla, near Cueva de la Magdalena, 900 m, 7 July 1951 (fl), Heywood 1058 (BM, RNG). Cordoba: Sierra Morena, Cjo N. S. of Fuenta Sagra, 7 km W. of Cordoba, 1 0 June 1 927 (fl), Wilmott s.n. (BM). Granada: Puerto de la Ragua, crossroads with road to Alpujana, 1400 m, 19 June 1988 (fl), Valdes et ai. 1011 (RNG). Malaga: Marbella, 500 m, N. de Puerto Cabopino, 30 m, 10 August 1984 (fl), Michel 7029 (BM, BR*). Sevilla: entre Cantillana y Alcolea del Rio, Km 9.0-9. 1,11 May 1986 (bud), Lopez & Molina CL 7917 86 (RNG, SEV*). Huelva: c. 5 km S. of Aracena on road N433, 600 m, 3 July 1974 (fl & fr), Leadley & Petty 205 (BM, RNG). GIBRALTAR. Guadacarte, 23 May 1927 (fl & fr), Croffen s.n. (BM). PORTUGAL. Minho: Caldas do Geres, June 1 887 (fl), Murray s.n. (BM). Tras-os-Montes: 6.4 km N. of Braganca, 450 m, 9 August 1957 (fl), Epsom College 325 (BM). Beira littoral: Coimbra, prope Carreira do Tiro, 50 m, 8 June 1936 (fi),Rothmaler s.n. (JE). Estremadura: Torres Vedras,arredoresde Runa, July 1 88 1 (fr), Barros & Cunha s.n. (BM). Ribatejo: Serra Monte Junto, 2 km from Cereal, above Sao Salvador, 300 m, 6 July 1 974 (fl), Leadley & Petty 259 (BM, RNG). Baixo Alentejo: Sines, proximo do Monte de Chamine, 13 June 1981 (fl), Barbosa, Gomes & Morena 13889 (RNG). AZORES. Graciosa: Monte d'Ajuda, 30 m, 7 July 1971 (fl), Gonfalves 3003 (BM). MADEIRA. Monte, 550 m, 19 October 1984 (fl & fr), Short 13 (BM). Ponta do Clerigo, 480 m, 1 1 June 1985 (fl), Press 896 (BM). CANARY ISLANDS. Santa Cruz: Calderata, 1 June 1913 (fl), Sprague Map 11 5c. H. perforatum subsp. veronense, Orient and Central Asia ' 100 N.K.B. ROBSON & Hutchinson 189 (K). Tenerife: Teneriffe, 1845 (fr), Bourgeau 246 (BM). Las Palmas: near S. Cruz, June 1892 (fl), Murray s.n. (BM). MOROCCO. Tanger: Tangier, July, Schousboe s.n. (JE). Rif: Targuist, Targuist to Al Hoceima, 3 km W. of Beni-Hadifa, 960 m, 29 June 1993 (fl), Jury \ 1278 (BM, RNG). Centre nord: Zerhouan, ruins of Roman city of Volubilis, 390 m, 4 June 1994 (fl), Jury 14865 (BM, RNG). Moyen Atlas: Taza, c. 42 km from Taza to Sidi Abdallah, around Jbel Tazzeka, 1200 m, 15 June 1992 (fl), Optima her V 996 (RNG). Haut Atlas: Demnat, Djebel Tihlatin, tribu de Ait Maala [Mahala], 25 July 1879 (fr), Ibrahim s.n. (BM). ALGERIA. Oran: Tlemcen, 15 June 1917 (fl), Faure s.n. (BM). Alger: Gorge deChiffa(Blida-Medea), 500m. 1 1 June 1971 (fl), Davis 5345 1 (BM). Kabylie: between Tizi Ouzou and Makouda, Oued Sebaou, 50-100 m, 17 June 1975 (fl). Davis 59146 (BM). TUNISIA. NW mountains (regions Kroumirie, Mogods, Dorsale Tunisienne),./?^ Pottier-Alapetite (1979: 508). SUDAN REPUBLIC. Darfur: Jebel Marra, Wadi Saria, 2300 m, 8 March 1983 (fr), S. Mielie 958 (BM); Jebel Marra, S. of Jebel Abau, c. 2600 m, 7 December 1954 (1. fl), Jackson 3335 (K). SAUDI ARABIA. Asir: Taif-Al Baha-Jabal Ibrahim road, 1 5 km N. of Al Mindak, 2040 m, 17 March 1987 (fl & e. fr), Collenette 6079 (BM, E*, K). ISRAEL. Upper Galilee: near Dafneh [Dafna?], 27 May 1945 (fl), Norris s.n. (BM). LEBANON. Coast: environs de Brumana. July to September 1979?, Cramer s.n. (G); Baynu, 1 3 June 1 943 (fl), Davis 64 1 2 (BM, K). Lebanon: Cedars, 20 July 1865 (fl), G. Post s.n. (K); Zahleh [Zahle], 7 May 1875 (fl), Post 809 (K). SYRIA. North: Monts Nusairy, Ain Halakim, 750-900 m, June 1 9 1 0 (fl), Haradjian 3537 (K); 20 km S. of Urdu [Ordu], 800-900 m, 8 June 1938 (fl), Dinsmore 20360 (K). CYPRUS. Kyrenia: environs du Monastere Armenien, 600-900 m, 26- 30 May 1912 (fl), Haradjian 348 (K). Paphos: Lisso [Lyso], c. 600 m, 10-16 June 1913 (fl), Haradjian 851 (K). Troodos: distr. Limassol, between Vasa and Agios Amvrosios, 21 May 1941 (fl), Davis 3538 (BM, K). IRAQ. Amadiyah: Rowandiz [Rowanduz] Gorge, 24 June 1934 (e. fr), Field & Lazar 856 (BASBG, K, W); 35 km NE of Zakho, Daimka village, 820 m, 5 July 1976 (fl), Al-Kaisi & Hamid 45365 (K). Rowanduz: Bekarl, 2200 m, 2 August 1956 (fl), Haley 17 (BM); Erbil distr., Monies Qandil, Pushtashan, c. 1 100 m, 28 July-1 August 1957 (fl), K.H. Rechinger 1 1035 (W). Sulinianiyah: Sulimanieh, in mont., June 1867 (fl), Haussknecht s.n. (BM, K); Sulaimaniya liwa, A'hmidau swamp, 28 July 1964 (fl), Makki 551 (W). IRAN. Azarbaijan: Tabriz to Marand, 1500m, 27 July 1971 (\.ft),Terme EV1N 34352E (BM, EVIN*). Kordestan: 64 km SW of Mahabad, 1500 m, 29 July 1962 (fl), Furse 3494 (K, W). Lorestan: Bicheh, 1 100 m, 29 May 1937 (fl), Koie 1506 (C*, W). Kohkiluyeh va Buyer Ahmadi: Tale' Khosravi to Sissakht, 29 July 1949 (fl), Beh houdi 975E (W). Zanjan: pass between Zandjan and Rudbar, 36 km from Zandjan, 2 100m, 9 July 1972(fl), Goldblatt 776 (BM). Hamadan: Hamadan, 2100 m, 7 September 1929 (fl), Rogers 9507 (BM, K). Gilan: Astara to Resht, 16 km S. of Astara, 30 m (by Caspian shore), 7 June 1962 (fl), Furse 2846 (K, W). Mazandaran: Elburz Mts, above Allamul, between Chalus and Karadz, c. 1980 m, 23 June 1966 (fl), Archibald 2403 (E, K). Tehran: prope urbem Teheran, 1 843 (fl), Kotschy 756 (BM, K); Haft Hartz Valley, N. of Teheran, 2100 m, June 1934 (fl), Trott 181 (K). Khorasan: in montibus inter Budjnurd [Bujnurd] et Morawe Tappeh, c. 1 300 m, 25-27 July 1 937 (fl & fr), K.H.&F. Rechinger 1 875 (W). AFGHANISTAN. Farah?: Khash-Dt., 2400 m, 8 August 1937 (fr), Koelz 1 29 1 3 (W). Fariab: c. 6.5 km N. of Darrah Zang village, c. 1 320 m, 30 June 1971 (fl), Grey-Wilson & Hewer 1312 (K). Jozjan: Tuzkar, 1200 m, 15 September 1939 (fr), Koelz 13977 (W). Baghlan: Salang Pass, N. side, 1800 m, 24 June 1966 (fl), Furse 8056 (K). Parwan: montee sud vers Col de Salang, 2500 m, August 1964 (fr), Haysu 487 (W). Kabul: Paghman, 25.6 km W. of Kabul, 2400-2700 m, 23 June 1939 (fl), Chaworth-Musters s.n. (BM). Logar: Ghorband, 1700 m, 14 August 1938 (fr), Koie 2789 (W). Konar: 17.6 km NW of Kamdesh towards Barg-i-Matal, c. 1567 m, 3 July 1969 (fl), Hewer 1 378 (K). Nangarhar: Mama Kheyl, 1 200 m, 23 May 1937 (ft), Koelz 11576(W). PAKISTAN. Kurram: Kurram Valley, Ziran, Imam Bagh, 30 August 1 972 (fr), Kazmi 3749 (H). Dir: Lowari, 6 July 1 968 (fl), Nasir 5 108 (RAW). Swat: Khawazakiela to Shangla, c. 1300 m, 2 June 1965 (fl), Lamond 1691 (E,KUH).Hazara: Jabri, 1 0 km from Shogran, 15 April 197% (f\),Muq. Shah & Oil. Khan 2403 (BM, ISL*). Chitral: Kesu, 1290 m, 24 May 1958 (e. fr), Bowes Lyo/i 714 (BM, RAW, W).Gilgit: Buni, 2100m, 1 876? (fr), Giles 392 Map 12 5c. H. perforatum subsp. veronense, Australia and New Zealand (introduced) A. Also in W. Australia (see records). STUDIES IN HYPERICUM (K). Baltistan: Balti, 2400 m, June 1915, Schlich 56 (BM). INDIA. Kashmir: Manasbal, 1 560 m, 5 June 1940 (fl), Ludlow & Sherriff 8120(BM). Himachal Pradesh: Pangi, Chenab R., N. bank between Darwas and Ishtwari, c. 2400 m, 12 June 1981 (fl), Sayers 3713 (BM). Uttar Pradesh: Tihri-Garhwal, Ganges valley near Jangla, 2400-2700 m, 12 July 1883(st),Dwtf»>977(BM). TURKMENISTAN. Aschabad: ad montem Aktepe [Kopet-Dag], 17 June l900(fl),Swtenis564a(BM, K); Kisil Arwat, Karakala,m. Sundsodagh, 31 May 1901 (fl), Sintenis 1862(JE). UZBEKISTAN. Taschkent, bei Tschimgan, westlich Auslaufer des Tian Shan, 1700 m, 14 July 1975 (fl), Breckle 3632 (K); Denauskiy raion, Akhunbabeva, 1 July 1938 (fl), Parabskaya s.n. (LE). TAJIKISTAN. Pamiro-Alai, Zalekhsi, N. slope of Aatni, R. Gulibysta 4- 5 km N. of Stalinabada [Dushanbe], 7 June 1942 (fl), Grigoriev 225 (LE); Kondor Gorge, 30 km N. of Dushanbe, W. side of R. Varzob, 1100m, 13 July 1975 (fl & e. fr), Pankhurst & James 93 (BM). KYRGYZSTAN. Tian-schan occid., in valle fl. Tales, prope pagum Pokrovka, 1 4 June 1 9 1 9 (fl & fr), Kushelev in HTU 077243 (TAI); Ferganskiy khrebet, Arslanbobskiy nut farm, R. Yasky-chu, 11 October 1933 (fr), Sokolov s.n. (LE). KAZAKHSTAN. Semirychensk prov., Vyernyy [Almaty ] distr., Zayliyskiy (Trans-Ilian) Alatau, 14 July 1907 (fl), Sokalsky 118 (BM); Ketmenskiy khrebet, Ushchelye, R. Ketmen, 20 September 1931 (fr), Rodin 1508 (LE). INTRODUCTIONS SOUTH AFRICA. Cape Province: Stellenbosch, College of Agricul- ture, 23 November 1948 (fl), Hulme s.n. (BM, K); Cape Town, slopes of Papegaaiberg, c. 105 m, 15 January 1969 (1. fl), Taylor 7376 (K, PRE*). REUNION. See Robson & Stevens, Flare des Mascareignes 49. Guttiferes: 6 (1980). These populations probably belong to subsp. veronense, but the specimens have not been re-studied. AUSTRALIA. New South Wales: Faulconbridge, 435 m, 17 November 1982 (fl), Coveny 11405 (K, NSW*). A.C.T.: Canberra, Jarrabomberra Avenue, just S. of junction with Narrabundah Lane, 580 m, 5 December 1 980 (fl), Canning 5031 (BM, CBG*). Victoria: Far NE, Corryong, near Anzac Drive, 25 November 1972 (fl), R.V. Smith 72/57 (K, MEL*). South Aus- tralia: Mt Lofty Ranges, Blackwood, 251 m, [comm. April 1929] (1. fl), Pritchard s.n. (BM, K, GH). Western Australia: approx. 8 km NE of York on York-Meckering road, 7 November 1979 (bud), Dixon s.n. (K). NEW ZEALAND. North Island: South Auckland, Taumarunui, 20 April 1928 (fl), Allan s.n. (K); Wellington, Masterton, 10 January 1929 (fl), Allan s.n. (K). South Island: Nelson, near Nelson City, 23 January 1929 (fr), Allan s.n. (K); Otago, Oamaru, 9 January 1929 (fl), Allan s.n. (K). HAWAIIAN ISLANDS. Hawai'i: North Kona, Huallai, 1961, Fosberg 42087 (BISH*) (fide Wagner, Herbst & Sohmer, 1999: 526). Hypericum perforatum var. collinum Woron. (from Georgia) and records of H. veronense and H. perforatum var. veronense from the Crimea (Vul'f, 1953: 110) and the Caucasus region both apparently refer to a narrow-leaved form of subsp. veronense with interrupted rather than vesicular lateral capsule vittae. 5d. Hypericum perforatum subsp. chinense N. Robson, subsp. nov.5 A subsp. perforatum inflorescentiis vel inflorescentiis partialibus vel inflorescentiis totis et partialbus congestis, ramulis pro ratione longis curvato-ascendentibus, petalis glandulis in lamina insertis nigris vel carentibus, differt. Type: China, Sichuan, Wa-shan, 2100-2400 m, July 1908 (fl), Wilson 2425 (BM!-holotype; A-isotype). Fig. 8, Map 13. H. perforatum var. microphyllum H. Leveille in Bull. Soc. Bot. France 54: 595 (1908); Lauener in Notes Roy. Bot. Card. Edin- burgh 27: 5 (1966) qua syn.; non DC. (1815). Type: China, Guizhou, Kouy-yang, pied de la montagne, 28 June 1897 (fl), 101 Bodinier 1665 (E-holotype; BM!-isotype). H. foliosissimum Koidz. in Acta Phytotax. Geobot. 3: 215 (1934); Nemoto, Fl. Japan, Suppl.: 483 (1936). Type: Japan, Honshu, Prov. Sima, Kamomura, Arasima, 27 July 1934, T. Ito s.n. (? - holotype). See note on next entry. H. perforatum var. angustifolium sensu Ohwi, Fl. Japan: 781(1 936); Y. Kimura in Bot. Mag. (Tokyo) 52: 406, f. 9 (1938), in Nakai & Honda, Nova fl. jap. 10: 133, ff. 47, 48 (1951); Momiyama in Satakeetal., Wild Fls Japan 2: 118, t. 1 14 f. 3 (1982). Momiyama states that the Japanese plant was originally introduced from Europe; but examination of Japanese specimens indicates that they match subsp. chinense rather than the south European subsp. veronense. Icon: Li Xiwen in Fl. R. P. Sin. 50(2): 68, t. 15, ff. 1^ (1990). Leaves petiolate; lamina (7-) 10-25 x 3-7(-15) mm, ± narrowly elliptic to narrowly oblong or linear (l:b=2.5-3.5), base rather narrowly cuneate to subcordate-amplexicaul, ?glaucous beneath. Inflorescence and/or partial inflorescences congested, branches rela- tively long, curved-ascending. Petals with laminar glands black, linear to punctiform, or often absent. Capsule valves with lateral vittae linear, narrow or somewhat interrupted or shortened but not swollen. China (eastern Qinghai and Gansu east to Shandong and south to Yunnan and Guizhou). The northern populations tend to have larger and relatively broader leaves. CHINA. Hebei :/? Li Xiwen (1990). Shanxi: fide Li Xiwen (1990). Shaanxi: Zhouzhi Xian, Chenhe Xiang, 1400 m, 14 August 1958 (fl & fr), Zhang 44 (IBSC, KUN, NAS); Chungmanshan, Hsingpaochuan, 20 August 1934 (fl & fr), Pai 1071 (PE); Tai-pei Shan, 1910 (fr), Purdom s.n. (K). Gansu: Yanchang Xian, Gongjiagou, 2150 m, 1 July 1989 (fl & fr). Yuan 1 008 ( A. BM, CAS, PE*); Tianshui, 1 350 m, 20 July 1 95 1 (fl & fr), Zhang 46 (KUN, PE). Henan: Lushi Xian, 1660 m, 2 October 1958 (fr), Fu 1640 (IBSC); Xichuan Xian, Lugou Sandaohe, 220 m, October 1956 (fr), Henan For. Dept. 1445 (PE). Shandong: Fei Hsien, Mong Shan, 12 July 1936 (fl), Cheo & Yen 38 (BM, W). Jiangsu. fide Li Xiwen (1990). Jiangxi: Kew Kiang [Jiujiang], 1873 (fl), Shearer s.n. (BM). Hunan: Wugang Xian, Yunshan, Qishucao, 1 150 m, 17 October 1963 (fr), Liu & He 16288 (IBSC); Changsha, Xiaonamen, 9 May 1964 (fl), Chen & Yue 1498 (NAS). Hubei: Wuchang, Luojia Shan, May 1932 (fl), Chung 9034 (A, NY, W); Badong Xian, 15 July 1957 (fl), Fu & Zhang 776 (IBSC, KUN); Shennongjia For. Distr., along Qiadonggou Canyon, 1900 m, 27 August 1980 (fl & fr), Sino- Amer. Exped. 396 (A, NY). Sichuan: Wenchuan Xian, 23 July 1928 (fl), Fang 1492 (GH, K, NY, PE, US); Fengjie Xian, Zhuyuan, 600 m, 24 July 1958 (fl & fr), Chang 25617 (IBSC, KUN. PE); Dujiandyan Municipality, Longxi For. Farm, 800 m, 2 September 1988 (fr), Bouffordel al. 14433 (A, BM, CAS, NY, PE*). Guizhou: Pin-fa, Kweiting, 400-550 m, 27 June 1930 (fl), Tsiang 5341 (IBSC, NAS, NY, PE); Yinjiang Xian, Huguosi, W. side of Fanjing Shan, 22 September 1986 (fl), Sino-Amer. Guizhou Exped. 1453 (BM, NY). Yunnan: vicinity of Yunnan-sen [Kunming], 1906 (fl), Maire 1505 (K); no precise locality, July 1936 (fl & fr), McLaren U.109 (A); [also Cavalerie 3125 (K)]. INTRODUCTIONS JAPAN. Hokkaido: Furano city, Yamabe, Tokyo University Forest, 13- 1 5 July 1 979 (fl), Sohma & Takahashi 478 ( H ). Key to hybrids of Hypericum perforatum This key should be used bearing in mind (a) the variability of the parental taxa, (b) the continuity of variation between parents and hybrids of H. x desetangsii nothosubsp. desetangsii and (c) that subsidiary lines present in younger stems may disappear in old ones 'Some specimens of subsp. chinense have been labelled 'subsp. confertiflorum (Debeaux) N. Robson'. but the type of H. perforatum var. confertiflorum has proved to belong to 6. H. attenuatum. 102 N.K.B. ROBSON Map 13 5d. H. perforatum subsp. chinense •, introduced O. 1 Stem internodes 2-lined (from leaf bases) or with weaker and usually incomplete subsidiary lines (from between leaves); leaves relatively laxly reticulate-veined 2 Stem intemodes ± completely 4-lined; leaves relatively densely reticulate-veined 5 2(1) Stem internodes 2-lined only; sepals usually narrow, more rarely broad, always acute with a hair point; capsule valves with not very numerous vittae to punctiform vesicles 3 Stem internodes also with weakly developed subsidiary lines; sepals narrow or quite broad, acute; capsule valves with relatively numer- ous narrow longitudinal vittae 4 3(2) Sepals entire, lanceolate; capsule with lateral vittae linear and shortened; leaves spreading, venation sometimes densely reticulate 5xx. x medium Sepals eroded-denticulate to glandular-ciliate, elliptic to lanceolate; capsule with lateral vittae linear or absent, not shortened; leaves often deflexed, venation laxly reticulate ... 5xbb. x desetangsii nsubsp. carinthiacum nf. perforatiforme 4(2) Sepals narrowly oblong to linear-lanceolate, finely acuminate or apiculate, eroded-denticulate to entire; leaves usually rather densely pale-gland-dotted 5xa. x desetangsii nsubsp. desetangsii Sepals broadly elliptic to linear-lanceolate, acute to acuminate, ± eroded-denticulate to subfimbriate or entire; leaves sparsely pale- gland-dotted ...5xbb. x desetangsii nsubsp. carinthiacum nf. perforatiforme 5(1) Leaves usually rather densely pale-gland-dotted, venation very laxly to rather densely translucent-reticulate; sepal apex usually very acute or eroded-denticulate with fine hair-point; sepals and petals with sparse dark glands 5xa. x desetangsii nsubsp. desetangsii densely translucent-reticulate; sepal apex obtuse or subacute, entire or finely denticulate, without distinct hair-point; sepals and petals without or with relatively numerous dark glands 6 6(5) Petal laminar glands all or some black; leaf apex obtuse, not acute, base cuneate to rounded ... 5xba. x desetangsii nsubsp. carinthiacum nf. maculatiforme Petal laminar glands all pale; leaf apex obtuse or sometimes some- times acute, base cuneate to cordate-amplexicaul 5xc. x desetangsii nsubsp. balcanicum 5x. Hypericum x desetangsii Lamotte in Bull. Soc. Bot. France 21: 121 (1874); Bonnet in Bull. Soc. Bot. France 25: 277 (1878); Schinz in Bull. Herb. Boissier II, 3: 11, 13 (1902), in Vierteljahrsschr Naturf. Ges. Zurich 49: 241 (1905); Tourlet in Bull. Soc. Bot. France 50: 307 (1903); Thellung in Allg. Bot. Z. 1912: 19 (1912); Salmon in J. Bot. (Lond.) 51: 317, t. 528 (1913); R. Keller in Engl. & Prantl, Nat. Pflanzenfam., 2nd ed. 21: 179 (1925); N. Robson in Bot. Soc. Brit. Isles Proc. 3: 99 (1958), mBull. Br. Mus. (Nat. Hist.), Bot. 8: 168, f. 55 (1981), in Wisskirchen & Haeupler, Standardliste Farn- u. Bliitenpfl. Deutschl: 268 (1998); A. Frohl. in Preslia 32: 97-99, f. 2 (1960); O. Schwarz in Drudea 5: 64 (1965). Type: France, Aube, Chapelle-St.-Luc pres Troyes, 11 July 1841 (1. fl), Des Etangs s.n. (CLFMectotype, selected here); Garenne de Villechetif pres Troyes, 5 August 1840 (e. fr), Des Etangs s.n. (K!-syntype). But see note under Ix. H. x laschii (p. 76) regarding the identity of these Des Etangs specimens. Fig. 9, Maps 14, 15. H. maculatum Crantz x H. perforatum L.; A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 557 (1911). Leaves without or almost without pale gland dots, venation rather Stems almost completely 4-lined to 2-lined. Leaves broadly ovate or STUDIES IN HYPERICUM 103 n Fig. 9 H. x desetangsii: A. nsubsp. desetangsii: (1): (a) branch; (b) stem section; (c) leaf; (d) sepal; (e) petal. B. nsubsp. desetangsii: (2): (f) branch; (g) stem section; (h) leaf; (i) sepal; (j) petal; (k) capsule. C. nsubsp. carinthiacum nforma perforatiforme: (1) stem section; (m) leaf; (n) sepal; (o) petal. D. nsubsp. carinthiacum nforma maculatiforme: (p) stem section; (q) leaf; (r) sepal; (s) petal. E. nsubsp. balcanicum: (t) stem section; (u) leaf; (v) sepal; (w) petal (a, f x 2/3; b, c, g, h, 1, m, p, q, t, u x 2; rest x 4). A. Anon. s.n. (France, Puy de Dome). B. Salmon s.n. (England, Sussex). C. Kerner 3649. D. Lillefosse s.n. E. Rechinger 782. 104 N.K.B. ROBSON broadly elliptic or ovate-oblong to linear; tertiary reticulate venation dense to lax; pale laminar glands dense (especially in upper leaves) to very few. Sepals broadly elliptic to linear-lanceolate, obtuse to acuminate, entire or ± eroded-denticulate to subfimbriate. Petals with black laminar glands rather numerous to few, mostly linear to mostly punctiform, or absent; black marginal glands present. Cap- sule with vittae all linear and vertical or some lateral, oblique and irregular to elongate-vesicular. Intermediate between the parents, but back-crossing to both at the tetraploid level of H. maculatum (subsp. obtusiusculum) has re- sulted in a complete series of intermediate forms of//, x desetangsii nothosubsp. desetangsii (cf. Crackles, 1990). At the diploid level (subsp. maculatum), two distinct forms of nothosubsp. carinthiacum occur, one triploid (2n=24) (H. maculatum (n=8) x H. perforatum (n= 1 6) or, probably rarely but conceivably, H. perforatum (n= 1 6, i.e. reduced) x H. maculatum (n=8)), the other pentaploid (2n=40) (H. perforatum (n=32, i.e. unreduced) x H. maculatum (n=8)). In the cross with subsp. maculatum (nothosubsp. carinthiacum), these forms are at least sometimes distinguishable (respectively nothoforma maculatiforme and nothoforma perforatiforme)', but in the cross with the presumed diploid subsp. immaculatum (nothosubsp. balcanicum), no distinct morphological forms are apparent and no chromosome counts have yet been made. As the specimens with this alleged parentage are intermediate between the parents, however, they are likely to be triploid. See Noack (1939) for a discussion of the cytology, but not the nomenclature. Although records are scattered, distribution of the hybrids would appear to be confined to the areas of distribution overlap of the parents; but vegetative propagation and (in nothosubsp. desetangsii) partial fertility of the hybrids have sometimes resulted in their occurrence in the absence of one or both parents. 5xa. Hypericum x desetangsii Lamotte nothosubsp. desetangsii - N. Robson in Wisskirchen & Haeupler, Standardliste Farn- u. Blutenpfl. Deutschl.: 268 (1998). Map 14. 5. H. perforatum L. x Ic. H. maculatum subsp. obtusiusculum (Tourlet) Hayek H. perforato-quadrangulum Lasch in Linnaea 4: 415 (1829). H. linnaeanum Callay & Gren. in Gren., Fl. jurass. 1: 154 (1863), nomen. ?//. x mixtum Du Moulin in Oesterr. hot. Z. 17: 390 ( 1 867); A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. I, 120(1): 568 (1911), in Mitt. Naturwiss. Vereins Steiermark 51: 227 (1915). Type: Germany, Bavaria, bei Neuburg, Bertolzheim, Du Moulin s.n. (W?- holotype). It is not certain from the descrip- tion whether this is an earlier synonym of H. x desetangsii or a later synonym of//, maculatum Crantz (cf. Martonfi, Repc^ak & Mihokova, 1996ft). Frohlich (1911: 568) was unable to find the type specimen in W; so, unless or until it is discovered and proves to be //. x desetangsii, it seems best to maintain the well known name. H. desetangsii var. genuinum Bonnet in Bull. Soc. Bot. France 25: 277 (1878); Burnat, Fl. Alpes marit. 2: 27 (1896); Schinz in Bull. Herb. Boissier II, 1: 14, 21 (1902), in Vierteljahrsschr. Naturf. Ges. Zurich 49: 24 1 ( 1 905). Type as for H. x desetangsii Lamotte. H. acutum subsp. desetangsii (Lamotte) Rouy in Rouy & Fouc., Fl. France 3: 336 (1896), comb, illegit. (Art. 52). H. acutum subsp. desetangsii var. genuinum (Bonnet) Rouy in Rouy & Fouc., Fl. France 3: 336 (1896), comb, illegit. (Art. 52). H. quadrangulum subsp. desetangsii (Lamotte) Tourlet in Bull. Soc. Bot. France 50: 307 (1903); A. Frohl. in Oesterr. Bot. Z. 63: 18 (1913), in Mitt. Naturwiss. Vereines Steiermark. 51: 220 (1915); Hegi, ///. Fl. Mitt.-Eur. 5(1): 518 (1925). H. maculatum subsp. erosum (Schinz) A. Frohl. x H. perforatum L. sensu A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl. 120(1): 559(1911). H. maculatum subsp. obtusiusculum (Tourlet) Hayek x H. perforatum L., A. Frohl. in Oesterr. Bot. Z. 63: 15 (1913). H. x desetangsii nothovar. desetangsii qua nothomorph sensu N. Robson in Tutin et al., Fl. Europaea 2: 269 ( 1 968), in Cullen et al., Eur.gard.fl. 4:60(1995). H. erosum (Schinz) O. Schwarz in Drudea 5: 63 (1965) pro parte quoad typum. Type as for H. quadrangulum var. erosum Schinz. Icon: Salmon in / Bot. (Lond.) 51: t. 528 (1913). Stems usually partially 4-lined. Leaves ovate or broadly elliptic to narrowly oblong or linear; tertiary reticulate venation dense to lax; pale laminar glands usually present, sometimes dense, especially in upper leaves. Sepals narrowly oblong to lanceolate or linear-lanceo- late, eroded-apiculate to acuminate and entire or denticulate. Petals with black laminar glands usually few, linear to elongate-puncti- form, and some marginal. Capsule with vittae all linear and vertical or some lateral, oblique and linear to elongate-vesicular. 2n=32, 40 &48 (Robson, 1956, 1975, 1 981; Robson & Adams, 1968;Martonfi et al., 1996a). For record of 2n=24, see p. 105. Habitat range of the parents excluding extremes of humidity. Distribution of H. maculatum subsp. obtusiusculum, i.e. NW Eu- rope south to central France and Switzerland and east to western Germany and Austria. SCOTLAND. S. Aberdeen: Drum, bank of R. Dee, 15 August 1956 (fr), Robson s.n. (BM). W. Perth: Perth, R. Tay, Moncrieffe L, 17 July 1938 (fl), Mackechnie & Wallace s.n. (BM, K). Argyll: Dunoon, 23 July 1 888 (fl), King s.n. (BM). Ayr: West Kilbride, 24 July 1973 (fl), B. Simpson s.n. (BM). Renfrew: Mr Hutcheson's School playing fields, 1 2 September 1 987, Dickson s.n. (GL). Lanark: Rotten Calder at Redlees Quarry, 1 3 September 1 987 (fl), Dickson s.n. (GL). Berwick: Earlston, 1926 (e. fr), Hayward s.n. (BM). ENGLAND. Cumberland: between Little Skelton and Ellaby, 19 June 1969 (fl), Halliday s.n. (LANC). Durham: near Nunstanton Carr, 82 m, 18 July 1975 (fl), Graham 8714 (BM). NW Yorks.: Swaledale, Langton, 23 August 1967 (1. fl), Gillingham s.n. (BM). SE Yorks.: Harswell old station, 20 July 1988 (fl), Duncan s.n. (BM). S. Lanes.: Wigan, Gathurst, 1947 (fl), Edmondson LIV 1975.560.7 (LIV). Cheshire: Castle Mill, 1 August 1867 (fl), Bailey s.n. (BM). Shropshire: Craig Llwyn, 28 September 1971 (fr), Benoit s.n. (BM). Oxford: Hailey, Synge Wood, 1 3 September 1 946 (1. fl), N. Simpson 46302 (BM). Bucks.: between Cadmore End and Fingest, 11 August 1946 (fl), Sandwith 3139 (BM). S. Essex: Woodhatch nearTheydon Boys, 19 July \9\4(fl),C.E. Britton 1247(K). W.Kent: S. of Ightham Court, 8 September 1957 (e. fr), McClintock s.n. (BM). Surrey: Burgh Heath, Perrott's Farm, 15 August 1929 (1. fl), Wallace s.n. (BM). E. Sussex: Lewes, Seaford Road, 11 July 1936 (fl), N. Simpson 36868 (BM). Dorset: near Wimborne, 22 August 1928 (fl), L.B. Hall in N. Simpson 45808 (BM). S. Devon: A 373 between Honiton and Awliscombe, 19 July 1989 (fl), Margetts s.n. (BM). WALES. Glamorgan: near Lisvane, Cefn On Halt, 3 July 1968 (fl), Pinkard s.n. (H). Radnor: near Builth Beacon, 1 8 August 1 940 (fl), Pugsley s.n. (BM). Merioneth: near Barmouth Junction (Morfa Mawddach) station, 5 October 1971 (fr), Benoit s.n. (BM). Caernarvon: Bangor, Talybont, Penbryn, 27 August 1972 (fl), Roberts (BM). Denbigh: Llangollen, 15 August 1953 (fl), Herb. Liverpool Mus. 1973.305.356 (LIV). IRELAND. Recorded from West Kerry only (Scannell & Synnott, 1 987), but likely to be more widespread. FRANCE. Aisne: Villers-Cotterets, 17 August 1879 (1. fl & e. fr), Bonnet in Soc. Dauph. 2409 (BM, FR). Oise: Aulmont pres Senlis, 25 September 1835 (fr), Gay s.n. (K). Seine-et-Marne: Nemours, 17 July 1892 (fl), Jeanpert 63 1 (BM). Aube: Villechetif [loc. class.], 29 July 1989, Guyot s.n. (H); see p. 102. Cote-d'Or: Magny-la-Ville, June 1887 (fl), Wilmotts.n. (K). STUDIES IN HYPERICUM Map 14 5xa. H. x desetangsii nsubsp. desetangsii •. Recorded in Ireland only from the south-west (Co. Kerry), but possibly more widespread. Saone-et-Loire: Curgy, pres Autun, 6 August 1889 (fl), Gillot 435 (BM). Allier: Sussat, la Veauce, July 1873 (fl), Deseglise s.n. (BM). Hte-Vienne: le Dorat, 5 July 1929 (fl), Elias in Heribaut- Joseph 441 (BM). Hte-Savoie: Constans, 2 August 1896 (fl), Boreau s.n. (BM). Savoie: Foret d'Apremont [pres Chambery], 30 July 1906 (1. fl), Chabert s.n. (BM). BELGIUM. Namur: entre Doisches et Petit-Doisches centre la frontiere francaise, au SW du Baquet, 16 July 1977 (fl), Duvigneaud 77B642 (H). GERMANY. Rheinland Pfalz: Zweibrucken, n.d.. Herb. Ekart/Irmisch s.n. (JE). Baden- Wurttemburg: [Schwabischen Alpen] Donstetten, August 1875 (fl & fr), Kemmler s.n. (FR). SWITZERLAND. Valais: Champery, 4 September 1 88 1 (fl), Lacaita s.n. (BM). Zurich: Affoltern, July 1902 (fl), Schinz s.n. (FR). St. Gallen: zwischen Litzibach und Vilters, c. 500 m, 8 July 1951 Biiel s.n. (BM, ZT*). AUSTRIA. Salzburg: bei Hellbrun, 26 August 1913 (fl), Frohlich s.n. (BM). Steiermark: ad vicum Maria-Trost prope urbem Graz, inter parentes, c. 430 m, July 1911 (fl), Frohlich in Hayek, Fl. stir. exs. 1 199 (BM, GZU, H). The above description covers the range of variation in this hybrid, which becomes increasingly difficult to recognize as it approaches each parent in form. The typical (f,) form has incomplete subsidiary stem lines, rather broadly oblong leaves with intermediately dense venation and laminar glands, oblong sepals with eroded-apiculate apex, and capsule vittae (both dorsal and lateral) linear. The chromo- some counts of 48 and 40 were, presumably, the results respectively of the fertilization of an unreduced embryo sac and a back-cross from that to another unreduced embryo sac. 5xb. Hypericum x desetangsii nothosubsp. carinthiacum (A. Frohl.) N. Robson [in B.S.B.I. News, no. 49: 50 (1988) in adnot.] in Wisskirchen & Haeupler, Standardliste Farn- u. Blutenpfl. Deutschl.: 268 (1998); Lihova, Martonfi & Martonfiova in Caryologia 53: 127 (2000). Type as for H. x carinthiacum A. Frohl. Fig. 9, Map 15. 5. H. perforatum L. x Ib. H. maculatum Crantz subsp. maculatum H. maculatum subsp. typicum A. Frohl. x perforatum L., A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 557 (1911); in Oesterr. Bot. Z. 63: 14 (1913). H. x carinthiacum A. Frohl. in Mitt. Naturwiss. Vereins Steiermark 51: 228 (19 15). Frohlich (19 13) listed a series of localities in Carin- thia where this hybrid was found, but did not designate a type either there or in his 1915 paper. See 5xba. H. x desetangsii nothosubsp. carinthiacum nothoforma maculatiforme for lectotype. H. x desetangsii nothomorph. carinthiacum (A. Frohl.) N. Robson in Repert. Spec. Nov. Regni Veg. 74: 23 (1967), in Tutin et al., Fl. Europaea 2: 269 (1968); in Watsonia 9: 203 (1972). Stems almost completely 4-lined to 2-lined. Leaves broadly ovate or broadly elliptic to ovate-oblong; tertiary venation dense to lax; pale laminar glands very few. Sepals broadly elliptic to linear-lanceolate, acute to acuminate, entire or ± eroded-denticulate to (glandular-) subfimbriate. Petals with black laminar glands rather numerous to few, mostly linear to mostly punctiform, or absent, black marginal glands present. Capsule with vittae all linear, dorsal and lateral, oblique, the latter regular or sometimes irregular. 2n=24, 40 [48, 56 experimentally] (see below). Habitat range of the parents? The above description covers both triploid and pentaploid hybrids with both varieties of subsp. maculatum. As regards these hybrids, Noack (1939) found the triploid to be intermediate between the parents, but could not distinguish the pentaploid from pure H. perforatum. Martonfi et al. (19966), on the other hand, were able to distinguish the pentaploid, and thus indirectly confirmed Frohlich's (1913, 1915, 1960) recognition of two distinct forms of this hybrid. Seeds obtained from a wild-collected pentaploid plant yielded 5 x 2n=40, 5 x 2n=48 and 3 x 2n=56 plants (Lihova, Martonfi & Martonfiova, 2000). 106 N.K.B. ROBSON 5xba. Hypericum x desetangsii nothosubsp. carinthiacum nothoforma maculatiforme (A. Frohl.) N. Robson in Wisskirchen & Haeupler, Slandardliste Farn-u. Blutenpfl. Deutschl. : 269 (1998). Type as for H. x carinthiacum forma maculatiforme A. Frohl. Map 15. H. quadrangulum var. erosum Schinz in Bull. Herb. Boissier II, 3: 21, 22 (1902). Type: Switzerland, Zurich, Dreilander Stein an der Hohe Rone, 31 July 1902 (fl), Schinz s.n. (ZMectotype, selected here - see also Martonfi et al., 1 999: 340). Frohlich (1911) raised Schinz's taxon to subspecific rank in H. maculatum and later (1913, 1915) reduced the combination to a synonym of H. maculatum subsp. obtusiusculum (Tourlet) Hayek. All the speci- mens cited by Schinz, however, have proved to be hybrids belonging to the above nothoforma. H. quadrangulum subsp. erosum (Schinz) Schinz in Vierteljahrsschr. Naturf. Ges. Zurich 49: 240, 241 (1905). Type as for H. quadrangulum var. erosum Schinz. H. maculatum Crantz x perforatum L. forma sub-maculatum in Oesterr. Bot. Z. 63: 14 (1913). H. x carinthiacum forma maculatiforme A. Frohl. in Mitt. Naturwiss. Vereins Steiermark 51: 229 (1915); Gu§ul. & Nyar. in Savul., Fl R. P. Roman. 4: 46 (1956). Type: Austria, Carinthia, Nordhange der Golitza sudlich Rosenbach, c. 800 m, 16 August 1911 (fl), Frohlich GZU 16301 (GZUMectotype, selected here). H. perforatum subsp. latifolium var. obscurum O. Schwarz in Drudea 5: 61 (1965), nom. illegit. (Art. 37). Stems with subsidiary lines almost complete though much less prominent than principal lines, i.e 4-lined. Leaves ovate to ± broadly elliptic or elliptic-oblong, with tertiary reticulation dense, quite visible. Sepals ovate to lanceolate or linear-oblong, rounded or obtuse to finely apiculate or acute, entire or finely denticulate, usually black-glandular-punctate. 2n=24 (Noack, 1939). SCOTLAND. Moray: Boat-of-Garten, station yard, 4 August 1978 (fl), McCallum Webster 19460 (BM). Argyll: head of Loch Feochan, 1 1 July 1994 (ft),B.H. Thompson s.n. (BM); Kintyre, Skipness, 3 1 July 1 %99(f\),Somerville s.n. (BM). Lanark: Bothwell Castle, 26 July 1987 (fl), Dickson s.n. (GL). NORWAY. Hordaland: Hardanger, Strandebarm, 10 July 1940 (fl), Lillefosse s.n. (BM, K, O*). FRANCE. Loire: St. Miedard, July 1895 (fl), Anthdme s.n. (H). Hte- Savoie: near Chambery, above St. Jean Arvey, c. 600 m, 1 8 July 1954 (fl), De Wit 5368 (WAG). GERMANY. Hessen: Grossherz, Vilbeler Wald, 12 July 1973 (fl), Diirer s.n. (FR); Kr. Alsfeld, Schnellhausen, Hahnberg, 8 July 1950 (fl), Hupke s.n. (JE). Thiiringen: Erfurt, Chaussee, July 1931, Reinecke s.n. (JE). SWITZERLAND. Jura: La Dole, 1 864 (fl), Favrat & Barbey 77 pp. (K). Zug: Geissboden, c. 950 m, 19 July 1902 (fl), Schinz s.n. (Z). Schwyz: Aufsteig zum Rikliklosterli von Goldau aus, 8 September 1902 (fr), Schinz & Brumes s.n. (Z). Zurich: Limmatal bei Herdern-Altstetten, c. 400 m, 3 August 1925, W. Koch s.n. (H). AUSTRIA. Steiermark: bei Strassgang bis Graz, n.d. (fl & fr), Frohlich s.n. (BM, GZU); bei M.[aria] Trost, 10 August 1911 (fl), Frohlich s.n. (BM, GZU). Niederosterreich: Berchtoldsdorf, July 1919 (fl), Fens! s.n. (K). ROMANIA. Transylvania: in monte Transsilv., July, Schur s.n. (JE). On the basis of specimens seen, nothoforma maculatiforme seems to be rarer than nothoforma perforatiforme. 5xbb. Hypericum x desetangsii nothosubsp. carinthiacum nothoforma perforatiforme (A. Frohl.) N. Robson in Wisskirchen & Haeupler, Standardliste Farn- u. Blutenpfl. Deutschl.: 269 (1998). Map 15. Hypericum perforatum [var.] P latifolium Gaudin, Fl helv. 4: 627 (1829); W. Koch in Rohling, Deutschl. Fl. 3rd ed. 5: 349 (1839), Syn. fl. germ, helv., 2nd ed.: 146 (1843), Syn. deut. schweiz. Fl. Map 15 5xb. H. x desetangsii nsubsp. carinthiacum. a. nforma maculatiforme •; b. nforma perforatiforme 0 . 5xc. nsubsp. balcanicum A. STUDIES IN HYPERICUM 2nd ed: 155 (1846), pro parte omnes quoad typum; Neilr., Fl. Nieder-Oesterr.: 826 (1859); Rouy in Rouy & Fouc., Fl. France 3: 333 (1896); Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925); Gusul. & Nyar. in Savul., Fl. R. P. Roman. 4: 29 (1956); Jordanov & Koz. in Jordanov, FL R. P. Bulg. 4: 261 (1970); N. Robson in Cullen et al., Eur. gard. fl. 4: 60 (1995), in Wisskirchen & Haeupler, Standardliste Farn-u. Blutenpfl. Deutschl.: 270 (1998). Type: Switzerland, no precise locality, Gaudin s.n. (LAU-holotype). The specimens labelled var. or subsp. latifolium that I have seen all have partly developed subsidiary stem lines and otherwise also agree with Frohlich's description of H. carinthiacum f. perforatiforme and specimens identified by him as such. Other than Koch himself, all the above authors cited this taxon as H. perforatum var. latifolium Koch; but Koch cited Gaudin, to whom this name should be credited. I have not seen Gaudin's type; but never having seen an undoubted H. perforatum with broad sepals, I assume that it belongs here. H. perforatum var. platycalyx Celak., Prodr.fl. Bohmen: 520 ( 1 875). Type: Czech Rep., Stfedo&sky, Jungbundzlau [Mlada Boleslav], Celakovsky (PR?). Celakovsky cited ^latifolium Koch' after his 'y platycalyx '; the type must therefore be that of var. latifolium Koch, i.e. var. latifolium Gaudin (see above). Celakovsky's 'type' is not in PR (Dr B. Skocclopolova, in litt. 2001). The only Hippelli specimens from Jungbundzlau in that herbarium are of subsp. veronense, whilst the only one labelled 'var. platycalyx' (15 July 1910) by Celakovsky is subsp. perforatum. H. perforatum [var.] P latifolium subvar. lineolatum Rouy in Rouy & Fouc., Fl. France 3: 333 (1896). Type as for 'H. lineolatum Jord. pro parte'. Rouy 'split' Jordan's type of//, lineolatum between his subvar. lineolatum of var. vulgare and subvar. lineolatum of var. latifolium. As Jordan's specimen is the type of var. vulgare, the subvar. of var. latifolium becomes typeless. H. perforatum subsp. latifolium (Gaudin) A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 525, f. 3a (1911), in Mitt. Naturwiss. Vereins Steiermark 51: 240 (1915); Hegi, ///. Fl. Mitt.-Eur. 5(1): 528 (1925); Hayek, Prodr. Fl. Pen. bale. 1: 533 (1925); Stjep.-Vesel. in Josifovc, Fl. Srbije 3: 119 (1972). H. perforatum subsp. latifolium forma dentatum A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 527, f. 3b, c (1911). Type: Austria, Steiermark, Plesch bei Grat- wein, 5 September 1909 (fl & fr), Frohlich GZU 16341 (GZUMectotype, selected here). H. perforatum subsp. latifolium forma fimbriatum A. Frohl. in Sitzungsber. K. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 529, f. 4(1911). Type: Austria, Steiermark, Miihlbachkogel bei Gratwein, c. 1000 m, 18 July 1909 (fl), Frohlich GZU 16345 (GZU!- lectotype, selected here). H. maculatum Crantz x perforatum L. forma sub-perforatum A. Frohl. in Oesterr. Bot. Z. 63: 14 (1913). H. x carinthiacum formaperforatiforme A. Frohl. in Mitt. Naturwiss. Vereins Steiermark 51: 229 (1915); Gu§ul. & Nyar. in Savul., Fl. R. P. Roman. 4: 46 (1956). Type: Austria, Karnten, Drauwiesen bei M.[aria] Elend, 16 July 1911 (fl), Frohlich GZU 16300 (GZUI-lectotype, selected here). H. maculatum subsp. desetangsiiforme A. Frohl. in Mitteil. Naturw. Ver. Steierm. 51: 219 (1915); Hegi, ///. Fl Mitteleuropa 5(1): 518 (1925). Type: Austria, Steiermark, bei Graz, Judendorf, July 1913 (fl), Frohlich GZU 16155 (GZUl-lectotype, selected here). H. maculatum subsp. desetangsiiforme var. aporosum A. Frohl. in Mitteil Naturw. Ver. Steierm. 51: 220 (1915). Type: Austria, Steiermark, bei Maria Trost, July 1913 (fl), Frohlich GZU 16171 (GZU!- lectotype, selected here). 107 //. x desetangsii nothomorph perforatiforme (A. Frohl.) N. Robson in Bull. Br. Mus. (Nat. Hist.), Bot. 8: 168 (1981); Martonfi, Repcak & Mihokova (19966). H. carpaticum Martonfi in Folia Geobot. 36: 374 (2001). Type: Slovakia (E.), [Volovske vrchy Mts] village of Pracovce, path near dextral feeder of Hrelikov potok brook, 27 July 1995 (fl), Martonfi 2012 (BRA-holotype). Martonfi (2001: 378) describes this pentaploid population as 'most likely of hybridogenous origin, with H. perforatum and H. maculatum [i.e. subsp. maculatum] as parents.' He claims, however, that it 'was stabi- lized and sufficiently reproductively isolated from its parents due to inherited apomixis' to allow it to be treated as a species. On the basis of his description and illustrations, however, it seems to be no more distinct from the rest of nothoforma perforatiforme than are parts of that taxon (e.g. 'subsp. desetangsiiforme'). It seems most appropriate, therefore, to include it in nothoforma perforatiforme. Stems with incomplete and scarcely prominent subsidiary lines or 2- lined. Leaves elliptic to rather narrowly oblong, with tertiary reticulation denser than in H. perforatum but obscure. Sepals elliptic to lanceolate and acute to finely acuminate or oblanceolate and apiculate, sometimes eroded-denticulate, usually black-glandular- striate to -punctate, those on main stems sometimes becoming markedly depressed ('desetangsiiforme'). 2n=40 (Noack, 1939, as H. perforatum x quadrangulum; Martonfi et al., I996a, b; Lihova, Martonfi & Martonfiova, 2000 [also 2n=48, 56 experimentally]. SWEDEN. Goteborg: Ringon, 23 July 1957 (1. fl), Blom s.n. (BM, GB*). FINLAND. Uusimaa: Nylandia, par. Kyrkslatt, Tero in pago Osterby, 8 August 1907 (fl), H. Lindberg PI. Fin. Exsicc. 806 (K). FRANCE. Hautes Pyrenees: between Gedre and Gavarnie, 900 m, 21 July 1922 (fl), Hutchinson, Matthews & Riley 14 (K). GERMANY. Mecklenburg-Pommern: Dalwitz, 1 864 (fl), Kokeils.n. (K) CZECH REPUBLIC. Bohemia, Jihocesky Kraj: *in silvis Novosedlecky revir prope opp. Pisek, August 1941, M. Deyl PR 164950 (PR). Moravia, Severomoravsky Kraj: Hohenstadt [Zabreh], bei Unterheinzendorf, July 1933 (fl), Hruby s.n. (H). SLOVAKIA. Kosicky Kraj: Prakovce, 3 August 1993 (fl), Martonfi 1452 (BM); *Zylinsky Kraj: Kysuce, near Nova Bystrica, c. 600 m, 7 July 1959, Futdk s.n. (SAV). AUSTRIA. Steiermark: Muhlbachkogel, bei Gratwein, 25 July 1910 (fl), Frohlich GZU 1 6338 (GZU). Karnten: [Cult, ex] Karawanken, nordlich der Goliza, n.d. (fl), Frohlich GZU 215 (GZU). Niederosterreich: in hort. bot. Vindobon., n.d. (fl), Kerne r in Fl. Exs. Austro-Hung. 3649 (BM, K). SWITZERLAND. ? : Pont d'Of, 16 July 1856 (fl), Depierre s.n. (BM). ITALY. Botzen [Bolzano]: Ritten, n.d. (fl), Hausmann s.n. (BM, IBF*). HUNGARY. *Comit. Vas: Totszgyorgy', 12 July 1882, Borbds BP 261 470 (BP). ROMANIA. Transylvania: Adamsdorf [Adamos], bei Brunes, 15 June 1873(fl), Schur 12138(BM). BOSNIA. Igman planina, am Veliko Palje, 14 June 1908 (fl), Ualy s.n. (K). Specimens apparently correctly identified as H. perforatum var. latifolium K. Koch are indistinguishable from those keying out to H. x carinthiacum var. perforatiforme; and the distribution of the H. perforatum variety does not exceed that of //. maculatum subsp. maculatum. I have therefore concluded that it, too, is a manifestation of this hybrid, not a form of H. perforatum itself. In addition, the voucher specimen for the count of 2n=40 by Martonfi et al. (see p. 105) agrees with the description and authentic specimens of H. maculatum subsp. desetangsiiforme A. Frohl., which I have there- fore also included in the synonymy of nothoforma perforatiforme. 5xc. Hypericum x desetangsii nothosubsp. balcanicum N. Robson, nothosubsp. nov. Type: Greece, E. Makedhonia, Nom. Serron, 108 N.K.B. ROBSON Mt Vrondous, near the EOS ski centre, 1700-1 800 m, 26 July 1981 (fl), Farsakoglou, Franzen & Strid 19521 (C!-holotype). Map 15. H. maculatum Crantz subsp. immaculatum (Murb.) A. Frohl. x perforation L. A nothosubsp. carinthiacum foliis sessilibus plerumque latioribus apice interdum acutis basin interdum cordatis, glandulis laminaribus petalorum pellucidis baud nigris, differt. Leaves sessile; lamina broadly ovate to elliptic-oblong, apex rounded to acute, base cuneate to cordate-amplexicaul. Petals without black laminar glands. SERBIA. Kosovo: Bertiscus, riv. Susica prope Pre (Ipek) [Pec], c. 800 m, 12 July 1933 (fl). Rechingerf. & Scheffer782 (EM). BULGARIA. Samokov: Vakarel, c. 750 m, 16 July 1945, Kitanov 370 (JE). Predbalkan: Ostabhang des Midshur [Midzor], 4 September 1958 (fr), Meyer 1171 (JE). Dobruja: Siiddobrudsha, Varna, Weg vom Strand zum Aladsha Monastir, 1 1 August 1958 (fl & e. fr), Meyer 414 (JE). GREECE. Makedhonia (E.): Norn. Kavalas, Mt Pangaion, summit area, Tsekour Madra, 1550-1750 m, 26 July 1977 (fl & e. fr), Papanicolaou 411 (C). These specimens are intermediate in form between H. perforatum and H. maculatum subsp. immaculatum and would therefore appear to be hybrids. Nevertheless, the Bulgarian specimens (with the possible exception of Kitanov 370, which I have not seen recently) and one of the Greek ones have apparently ripe or immature fruit, which, if they are triploids, would be unlikely. The type and the Kosovo specimen, however, are convincingly intermediate and would seem to be true hybrids. 5xx. Hypericum x medium Peterm., Fl. lips, excurs.: 563 (1838); Rchb., lcon.fl. germ. helv. 6: f. 5178 (1842-1844); Rouy in Rouy & Fouc., Fl. France 3: 337 (1896); N. Robson in Stace, Hybridization and the flora of the Br. Isles: 166 (1975). Syntypes: Germany, Leipzig, ad pagum Leutsch, July-August (fl), Petermann s.n. (LZ). 5. H. perforatum L. x 4. H. tetrapterum Fr. //. x sparsiflorum Schur in Verh. Naturf. Vereins Briinn 15(2): 158 (1877). Type: Moravia, bei Bisterz auf der Mnischihora, im Punkwathale na'chst Briinn, July-August 1 87-, Schur s.n. (BRNU? or LW?). H. perforato-acutum sec. O. Kuntze & Fockefide Rouy & Foucaud, Fl. France 3: 337 (1896). H. acutum x perforatum; A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 120(1): 592 (1911). H. x dubioides Druce in Bot. Soc. Exch. Cl. Br. Isles 8(5): 868 ( 1 929), nomen. Only the formula H. perforatum x quadrangulum was cited. Icon: Rchb., lcon.fl. germ. helv. 6: f. 5178 (1842-1844). Intermediate between the parents Stems without or with partial subsidiary lines. Leaves oblong to elliptic-lanceolate with dense to lax, rather obscure reticulate venation; pale laminar glands rela- tively few, varying in size. Sepals oblong to lanceolate, acuminate, without black glands. Petals without or with linear to striiform black laminar glands. Capsule with shortened lateral vittae. This hybrid is rare, on account of the differing habitats of its parents, but could occur anywhere where their distributions overlap. Frohlich (1911: 593) cites three German localities, but I have seen only the two French specimens, the third (Romanian) one having been determined by Dr Martonfi. FRANCE. Jura: Beauvoisin pres Chaussin, 6 July 1853 (fl), Michalet s.n. (K); Chaussin, September 1873 (fr), Michalet s.n. (CLF). GERMANY. Leipzig. See type. CZECH REP. Moravia. See type of H. x sparsiflorum Schur. ROMANIA. Transylvania: *Comit. Hunyad., Pareng mts, in Bordi, 21 June 1906, Jdvorka BP 1 1535 (BP). 5xxx. (Hypericum perforatum L. x H. maculatum Crantz) x H. tetrapterum Fr. 5x. H. x desetangsii Lamotte x 3. H. tetrapterum Fr. (H. maculatum Crantz x H. perforatum L.) x H. acutum Moench sensu A. Frohl. in Sitzungsber. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl. 120( 1 ): 59 1 , f. 1 3 ( 1 9 1 1 ). Approaching H. tetrapterum in habit. Stems with wider subsidiary lines than H. perforatum x maculatum. Leaves broadly ovate, almost semi-amplexicaul, with laxer reticulation venation and quite densely and finely pale-gland-dotted. Inflorescence more compactly flowered. Flowers smaller than in the hybrid parent, c. 20 mm in diam. Sepals relatively broadly ovate, acuminate; laminar glands all or mostly pale, striiform to punctiform. Petals darker yellow than in H. tetrapterum, like those of the hybrid parent, with pale glands only. Stamens c. 2/3 x petals. Ovary broadly ovate (no fruit seen). Switzerland (Schwyz, St. Gallen). SWITZERLAND. Schwyz: Roblosen, near Einsiedeln, 900 m. Thellung s.n. (Z*). St. Gallen: Fiirstenland, in den Senke ostlich von Griindenwald bei Winkeln, 640 m, 2 August 1946 (fl), Koch 46/421 I (H). Like H. x medium, this hybrid has parents with widely differing habitat requirements and must therefore be rare. The above data have been extracted mainly from Frohlich's account (191 1: 591). Hypericum assurgens Peterm., Anal. Pflanzenschl. hot. Excurs. Leipzig: 67 (1846), which Petermann himself published as a syno- nym of his H. perforatum var. petiolatum (described on the same page), was claimed by Otto Kuntze (Flora 63: 293, 1880) to be the unlikely hybrid H. humifusum x perforatum. On the basis of the Petermann specimen named H. assurgens in W!, I regard this plant as a rather long-petioled form of H. perforatum subsp. perforatum (see p. 89) . 6. Hypericum attenuatum Fisch. ex Choisy, Prodr. monogr. Hyperic. : 47, t. 6 ( 1 82 1 ), in DC., Prodr. 1 : 548 ( 1 824); Turcz., Fl. Baical-Dahur. 1: 251 (1842); Ledeb., Fl. ross. 1: 448 (1842); Maxim., Primit. fl. amur.: 65 (1859), in Bull. Acad. Imp. Sci. Saint-Peter -sbourg 27: 433 (1882), Mel. Biol. Bull. Acad. Imp. Sci. Saint-Peter sbourg 11: 166 (1882); Regel, Tent.fl. ussuri: 33 (1861); Hance in J. Bot. 12: 259 (1874), op. cit. 16: 104 (1878); Franchet, PL david. 1: 56 (1884); Forbes & Hemsley in J. Linn. Soc. Bot. 23: 72 (1886); Diels in Bot. Jahrb. Syst. 29: 476 (1901); Freyn in Oesterr. Bot. Z. 52: 17 (1902); Komarov, Fl. Manshur. 3: 43 (1905); H. Leveille in Bull. Soc. Bot. France 54: 594 (1908); Nakai, Fl. kor. 1: 95 (1909), 2: 453 (1911); Komarov & Klob.-Alis., Opred r. Dal'nevost. kr. 2: 747, t. 232 f. 2 (1932); Kitagawa, Lineament. Fl. Mandshur. : 3 1 7 ( 1 939); Gorschkova in Shishkin & Bobrov, Fl. URSS 15: 236 (1949); Komarov, Fl. Manchzh. 5: 48 (1950); Y. Kimura in Nakai & Honda, Novafl. jap. 10: 213 (1951), in J. jap. Bot. 34: 323 (1959); Kitamura & Murati in Acta Phytotax. Geobot. 20: 197 (1962); Noda, Fl. N. E. Prov. (Manch.) China: 794 (1971); Anon., Iconogr. Cormoph. Sinicae 2: 875, f. 3480 ( 1 972); Wu C.-J. in Fl. Intramongol. 4: 9 1 , t. 42 ff. 1-3 (1979); Lee, T. B. ///. Fl. Korea: 545 cum f. (1989); STUDIES IN HYPERICUM 109 Fig. 10 A. H. elegans: (a) habit; (b) sepal; (c) petal; (d) capsule. B. H. attenuatum: (e) habit; (f) sepal; (g) petal; (h) capsule (a, e x 2/3; c, g, h x 4; rest x 8). A. (a-c) Blocky s.n.; (d) Launert s.n. B. (e) Kang 57; (f, g) Campbell s.n.; (h) Kang 68. 110 N.K.B. ROBSON Map 16 6. H. attenuatum Li Xiwen in Fl. R. P. Sinicae 50(2): 69, t. 15 ff. 5-8 (1990); N. Robson in Cullen et al., Ear. gard. fl. 4: 60 (1995); non H. attenuatum Link (1822) (= H. canariense L.). Type: Russia, Irkutsk, Dauria prope Macarova in valle granitica et alibi, n.d. (fl), Fischer s.n. (G-DC-holotype, microfiche!; P!-isotype). Fig. 10, Map 16. H. perforatum var. confertiflorum Debeaux in Actes Soc. Linn. Bordeaux 31: 130 (1876), Florule du Tche-fou: 35 (1877) ['confertiflora']; Maxim, in Bull. Acad. Imp. Sci. Saint- Petersbourg 27: 432 (1882), Mel. Biol. Bull. Acad. Imp. Sci. Saint-Petersbourg 11: 166 ( 1 882). Type: China, Shandong, Tche- fou [Chefoo=Yantai], 12 July 1860 (fl & e. fr), Debeaux Exp. de Chine 60. (Pl-lectotype, selected here). Although this name was adopted by me as the provisional basionym of the Chinese subspecies of//, perforatum, and I determined some specimens of it as H. perforatum subsp. confertiflorum, a syntype borrowed from P clearly belongs to H. attenuatum, a species for which the epithet confertiflorum is appropriate. ?//. kamtschaticum var. senanense sensu Y.N. Lee, FL kor. 3rd ed.: 232, f. 685(1998). Icones: Wu C.-J. in Fl. Intramongolica 4: 92, t. 42 ff. 1-3 (1979); Li Xiwen in Fl. R. P. Sinicae 50(2): 68, t.15 ff. 5-8 (1990). Perennial herb 0. 1-0.45(-0.7) m tall, erect from rootstock or creeping rhizomatous base, with stems numerous to few, caespitose, much branched. Stems 2-lined, with black punctiform and striiform glands on lines and often sparsely (reddish or black) elsewhere; internodes (10-) 15— 40 mm, usually exceeding leaves. Leaves sessile; lamina (8-) 15-31 (-38) x (3-)5-12(-15) mm, el- liptic or elliptic-oblong to oblanceolate or rarely ovate, paler beneath, chartaceous; apex obtuse to rounded, margin entire, plane, base subcordate to cuneate; venation: 2 pairs of main laterals from lower sixth of midrib, tertiary reticulation dense but often rather obscure and apparently lax; laminar glands puncti- form, small, pale and black, few or scattered, mainly distal; intramarginal glands black, spaced. Inflorescence many- to few- (rarely 1-) flowered from 1-4 nodes, sometimes with flowering branches from up to 4 nodes below, the whole cylindric to py- ramidal; pedicels (1.5-)3-4 mm; bracts and bracteoles 3.5-6.5 mm long, oblong-elliptic, entire. Flowers 13-20(-25) mm in diam., stellate; buds ovoid, subacute to acute. Sepals 5, unequal to subequal, (3.5-)5-10 x 1—4 mm, erect in bud and fruit, triangular- ovate to lanceolate, acute to subacuminate, entire; veins 5-7, distally branched; laminar glands pale, striiform to punctiform and black, punctiform, scattered, rather sparse or rarely absent; intramarginal and marginal glands black, punctiform, sparse, distal. Petals 5, golden? yellow, tinged red in bud, 8-12 x 4-7 mm, 2.4-3 x sepals, oblong-obovate, asymmetric, entire, laminar glands black, punctiform to striiform, scattered, marginal glands black, distally dense. Stamens c. 90, '3'-fascicled, longest 8-10 mm, c. 0.7-0.8 x petals; anther gland black. Ovary 3-locular, 2.5- 3.5 x 2-3 mm, narrowly ovoid; styles 3, free, 4-4.5 mm, 1.3-1.6 x ovary, widely spreading; stigmas narrowly capitate. Capsule (4-)6-10 x 2-6 mm, 2-3 x sepals, broadly ovoid or oblong-ovoid to narrowly conic; valves densely longitudinally vittate, occasion- ally with a few black longitudinal glandular streaks. Seeds dark? brown, c. 0.7 mm, cylindric, slightly carinate and apiculate, not appendiculate; testa finely linear- foveolate. 2n=16 (Probatova & Sokolovskaya, 1986). Fields, pastures, steppes, grassy and dry stony slopes, pebble shores, forest margins and clearings; China: s.l. (Liaoning) - 2000 m (Guiz- hou); Russia. Russia (E. Siberia: Angara-Sayan, Dauria, Lena-Kolyma; Far East: STUDIES IN HYPERICUM Zeya-Bureya, Uda, Ussuri, Sakhalin?6), Mongolia, China (Nei Mon- gol, Heilongjiang and Jilin south to Guizhou, N. Guangxi and Guangdong), Korea. RUSSIA. Angara-Sayan: Inter Irkutsk et Jenisseisk, ad fl. Angara (Tunguska super.) pro pagum Padun, 1 867 (fl), Czekanowski s.n. (BM); Prov. Irkutsk, distr. Balagansk, prope Bashejewskoje, 11 July 1904 (fl), Malzew 1 9 1 9 (BD, H). Dauria: Chita distr., Priiskovaya stn, valley of R. Nercha, NE slope of Mt Krasnyy mys, 28 June 1 952 (fl), Sergievskaya & Obolentsev s.n. (H); Nertschintsk, 1899 (fl), Karo 143 (BD, BM, H, JE, K). Lena-Kolyma: see type. Zeya-Bureya: Zejskaja Pristan am Zeaflusse, July 1 899 (fl), Karo 359 (BM, H, JE, K); Blagowjestschensk in Amurgebeite, 1 898 (fl), Karo 224 (BM, JE). Uda: present (fide Gorschkova, 1949: 237). Ussuri: Prov. Amurensis, fluvium Amur circa stationem Radde, 25 June 1 895 (fl), Komarov Fl. Man. 1092 (BM, K). Sakhalin? (see footnote 6). MONGOLIA. Khalka Gol, 26 July 1 902 (fl & e. fr), Campbell s.n. (BM); Ourato, Oulachan occid., August 1882? (fl), David 1944 (K). CHINA. Nei Mongol: Solon Qi, Honggolj, 17 August 1956 (fr), Liou T.N. et al. 8250 (NAS, PE); Ergun Qi, 720-850 m, 9 July 1951 (fl & fr), WangZ. et al. 1 1 97 (PE). Heilongjiang: Ningan Xian, Jingpo Hu, 20 July 1 975 (fl), Fu P.Y.et al. 3327A (IFP); Yilan Xian, Weiken R., 7 August 1959 (fl), Chang, Yu Liang et al. 1971 (IBSC). Jilin: Jilin Shi, Longtan Shan, 8 September 1956 (fr), Chang Y.L etal.911 (KUN,PE); Jiutai Xian, Tumenling Village, 300m, 28 August 1950 (fr), Fu P.Y. et al. 2027 (IBSC, PE); Mukden to Kirin, Tang- ho-ko, Sungari R. to Hui-fa R., 1886 (fl), James s.n. (K). Liaoning: Chikuanshan, Tiehling, 250 m, 18 July 1930 (fl), Kung H.W. 648 (PE); near Mukden, Lao-yeh Ling and other hills, May-August 1886 (fl), James s.n. (K). Hebei: Neiqiu Xian, Xiaolingdi Village, 1600-1700 m, 21 July 1950 (fl & fr), Liu K 13150 (NAS); Hsiaowutai Shan, 1500 m, 29 August 1913 (fr), Meyer 1376 (GH, K). Shanxi: Wutai Xian, Gengzhen Xiang, 2000m, 19 July 1959 (fl & fr), Kuan & Chen 2433 (PE); Xiangning Xian, Guanwang Temple, 17 July 1960 (fl), Liu X.Y. 20863 (HNWP); Wenshui, S. of Hechiata, 14 August 1925(fl),A>rtg68(BMp,GH).Shaanxi:Schansi, 1500m, 18 August 1935 (fl), Licent 125 19 (GH); Shang Xian, 1060 m, 6 August 1952 (fl & fr), Wang T.P. 16043 (PE); Mt Lao-y-san, Sciuz-scien, June 1897 (fr), Fr Hugh s.n. (BM). Gansu.fide Li Xiwen (1990). Shandong: Fei Xian, Meng Shan, 350 m, 12 July 1936 (fl & fr), Cheo & Yen 38 (GH); ibid., 20 July 1936 (fl & fr), Cheo & Yen 122 (BM); Wulian Xian, Wulian Shan, 20 July 1959 (fr), Cheo IK et al. 5128 (NAS). Henan: Lushi Xian, Laochun Shan, 555 m, 4 August 1935 (fl), Liou KM. 4892 (PE); Xinyang Xian, Youhe Gongshe, 500 m, 1 8 July 1965 (fl), Chen & Ming 14 (PE). Jiangsu: Wuxi, Hui Shan, 1 1 July 1956 (fl), Ting & Wang 925 (NAS); Zhenze, Dong Shan, Huachuan Ling, 29 July 1958 (fr), Wu W.X. 3782 (IBSC). Anhui: Wuhu, 20 June 1941 (fl), Migo s.n. (NAS); Chu Xian, Langyashan, 2 August 1951 (fr), Huadong Working Stn 3148 (PE). Zhejiang: Jinhua Xian, 24 July 1927 (fl), Keng Y. L. 873 (NAS, PE). Fujian: Wuyi Shan, peak of Huanggang Shan, 2130 m, 20 July 1980 (fl), Wuyishan Exped. 80-0147 (IBSC). Jiangxi: Suichuan Xian, 1100 m, 22 September 1963 (fr), Yue J.S. et al. 4159 (IBSC); Julian Shan, n.d. (fl & fr), Julianshan Forestry Farm 781088 (IBSC). Guangdong, Guangxi: fide Li Xiwen (1990). Hunan: Sangzhi Xian, Bamaoxi Gongshe, 1500 m, 3 July 1975 (fl), Li & Shao 750126 (IBSC); Fenghuang Xian, Liantouyang Xiang, 600 m, 17 September 1988 (fl & fr), Wuling Exped. 1209 (IBSC). Hubei: Ichang and immediate neighbourhood, 1885-1888 (fr), Henry 4157 (BM, GH, K); Wuchang, Luojishan, July 1932 (fr), Chung H. H. 9078 (A). Sichuan: Sintsin Hsien [Xinjin Xian], 600 m, 23 July 1938 (fr), Wang T.P. 8309 (PE). Guizhou: Hsing Shan, 2000 m, 1 1 October 1931 (fr), Steward et al. 536 (PE). KOREA (N). Namp'o-Si: in herbidis Chinnampo [Namp'o], August 1906 (e. fr), Faurie 61S (EM). Hypericum attenuatuni is most closely related to 1 . H. maculatum and 1 1 . H. scouleri (respectively from western Eurasia and western North America), differing from both inter alia by the constantly 2- lined stem, from H. maculatum by the acute sepals, and from H. scouleri by the gland-dotted petals. It would seem to be one diploid parent (with H. maculatum subsp. maculatum) of the amphidiploid H. perforatum. Ill Kitamura & Murati ( 1 962) sank the Japanese H. tosaense Makino (1903), with its f. insulare Y. Kimura (1951) and var. atumense (Y. Kimura)Y. Kimura (1959), and H. momoseanum Makino (1931) into H. attenuatum, but these taxa appear to be distinct from that species (see key, p. 67). See also Y. Kimura (1959), who included them all in an '//. attenuatum Superspecies'. 7. Hypericum elegans Stephan ex Willd., Sp. PL 3: 1469 (1802); M. Bieb., Fl. taur.-caucas. 2: 230 (1808), 3: 519 (1819); Poiret, Encycl. Suppl. 3: 701 (1814); Choisy, Prodr. monogr. Hyperic.: 55 (1821) 'non Steph.' (which is said to = perforatum), in DC., Prodr. 1: 551 (1824); Ledeb., FL altaic. 3: 368 (1831), FL ross. 1:450(1842); Spach, ///s/. not. veg.,Phan.S: 389 (1836), in Ann. Sci. Nat. Bot. 5: 357 (1836); Boissier, FL orient. 1: 805 (1867); Kerner in Oesterr. Bot. Z. 24: 165 (1874); Krylov, FL Altaya 1: 190 (1901); Woronow in Kuzn., Busch & Fomin, Fl. Cauc. Crit. 3, No. 9: 45 (1906); Hayek, Prodr. Fl. Pen. Bale. 1: 537 (1925); Stefanoff in God. Agr.-les. Fak. Univ. Sofia 10: t. 4 f. 24 (1932), 11: 175 (1933), 12: 87 (1934), in Pflanzenareale 4: karte 7 (1933); Krylov, FL Zap. Sib. 8: 1907 (1935); Fedtchenko & Shishkin, Fl. Yugo-Vost. Evrop. casti SSSR 5: 711 (1938); Stoyanov & Stefanov, Fl. bulg.: 774 (1948); Gorschkova in Shishkin & Bobrov, FL URSS 15: 243 (1949); E. Wulf, FL Kryma 2(3): 109 (1953); Karnaukh in Klokov & Visjulina, Fl. URSR 7: 307 (1955); Kornas in Szafera & PawBowski, FL Polska 7: 28 (1955); Rzazade in Karjagin et al. (Eds), FL Azerbaid_ana 6: 257 (1955); Go§ul. & Nyar. in Savul., Fl. R. P. Romano 4: 33, t. 3 f. 1 (1956); Janchen in Hofler & Knoll, Cat. Fl. Austr. 1: 257 (1957); Zeleny in Sbornik Klubu Pffrod. v Brne 32: 69 (1960); Grossg., FL KavL, 2nd ed. 6: 175, karta 191 (1962); Vasyleva in Pavlov, FL Kazakhstana 6: 163 (1963); Robson in Davis, FL Turkey 2:401, f. 12.25 (1967), in Tutin et al. (Eds), FL Europaea 2: 269 (1968), in Huxley & Griffiths (Eds), R.H.S. Diet. Gdng 2: 629 (1992), in Cullen et al. (Eds), Eur. Card. FL 4: 60 (1995); Jordanov & Koz. in Jordanov, FL R. P. Bulg. 4: 261, t. 49 f. 1 (1970); Stjep.-Vesel. in Josifovic, FL Srbje 3: 119, t. 23 ff. 5, 5a ( 1 972); Hegi, ///. FL Mitt. -Eur. , 2nd ed. 5( 1 ): 522, ff. 1 998 (map), 2007 (1975); Meusel, Vergl. Chor. Zentraleurop. FL 2: 288, Karte 285c (1978); Zeleny, Jasicova & Zahradnikova in Futak & Bertova, FL Slovenska 3: 3 1 1 , t. 38 f. 2, map 57 ( 1 982); Hagemann in Flora 173: 109, tt. 9-11, 39 (1983); Dostal, Nova Kvetena CSSR 1: 584 (1989). Type: Russia, 'Habitat in Sibiria', n.d. (fl), Stephan in Herb. Willd. 14473 (B-WILLD-holotype, micro.!). Fig. 10, Map 17. H. kohlianum Spreng., Fl. hal. tent, nov.: 214, t. 9 (1806); Baumg., Enum. stirp. Transsilv. 2: 392 (1816); Schur, Enum.pl. Transsilv.: 132(1 866). Type: Germany, Halle, 'in vineis ad Bennstadt rarius', July (fl), Sprengel s.n. (JE!-holotype). H. pulchrum Pallas ex M. Bieb., Fl. taur.-cauc. 2: 230 (1808), in synon. H. hyssopifolium [var.] ysensu Choisy, Prodr. monogr. Hyperic.: 56 (1821). Choisy cited 'Bieb. cauc. 2. p. 231'. 'Type' as for var. pauciglandulosum Choisy (1824). H. hyssopifolium var. pauciglandulosum Choisy in DC., Prodr. 1: 552 (1824). Type: Ukraine?, Tanais [R. Don], 1819, Goldbach s.n. (G-DC!-holotype). H. elegans var. pectinatum Celak. in Oesterr. Bot. Z. 24: 140 ( 1 874). Type: Russia, Ciscaucasus, 'in pratis ad rivum Kuma prope pagum Maslow, July 1843 (fl), Hohenackers.n. (W-holotype; K!- isotype). 6Maximowicz (1882) attributed H. attenuatum sensu F. Schmidt, Fl. Sachal. no. 87, to H. erectum, but Gorschkova (1949) included Sakhalin in the distribution of H. attenuatum. Vorobiev et al. (1974) did not include this species in their key to vascular plants of Sakhalin and the Kurile Islands. Maximowicz would seem to have been correct. 112 N.K.B. ROBSON H. elegans var. stepposum Savul. & Rayss, Mat.fl. Basarab. 3: 175 (1934); Gu§ul. & Nyar. in Savul., Fl. R. P. Roman. 4: 34, t. 5 f.l (1956); Jordanov & Koz, in Jordanov, Fl. R. P. Bulg. 4: 262 (1970). Type: Romania, Reg. Constanta, intra Valea Nucaliler §i Mahmudia (r. Tulcea), Savulescu ? s.n. (BUCA). H. elegans var. genuinum Gu§ul. in Savul., Fl. R. P. Roman. 4: 34 (roman.), 385 (latin) (1956). Type as for H. elegans Stephan ex Willd. H. elegans var. elegans Jordanov & Koz. in Jordanov, Fl. R. P. Bulg. 4:262(1970). H. elegans var. pedunculatum Jordanov & Koz. in Jordanov, Fl. R. P. Bulg. 4: 262, 709 (1970) ('pedunculata') in clav., nom. invalid, sine descr. lat. (Art. 36). Icones: Rchb., Iconogr. hot. pi. crit. 3: 68, t. 280 f. 443 (1825); Icon, fl. germ. helv. 6: t. 350 (1842-1844); Fedchenko & Shishkin, Fl. Yugo-Vost. Evrop. casti SSSR 3: 712, f. 487 (1938). Perennial herb 0. 1 8-0.45(0.55) m tall, erect or occasionally shortly ascending and rooting from rootstock, with stems solitary or few, usually branched at most nodes. Stems incompletely 2-lined (nodes terete near base), with ± dense black punctiform glands on or near lines and occasionally elsewhere; internodes 8-40 mm, shorter than to usually exceeding leaves. Leaves sessile; lamina 1 5-25(-33) x 3- 9(-12) mm, oblong-lanceolate or ovate-lanceolate to triangular-lanceolate, paler beneath, chartaceous; apex subobtuse, margin entire, revolute, base subcordate, subamplexicaul; venation: 3 pairs of main laterals from near base of midrib, tertiary reticulation obscure; laminar glands all pale, dense, or with a few black, scat- tered, punctiform; intramarginal glands black, rather dense. Inflorescence (l-)5-c. 65-flowered, from \-b nodes, with flowering branches from up to 5 nodes below, the whole corymbiform to ± narrowly pyramidal or narrowly cylindric; pedicels 1-2 mm; bracts and bracteoles 3.5-5 mm long, lanceolate, sparsely black-glandular- denticulate. Flowers ( 1 5-)20-25 mm in diam., stellate; buds narrowly ovoid, (sub)acute. Sepals 5, free, 4-6 x 1-2 mm, erect in bud and fruit, lanceolate to ovate-lanceolate, acute, with ± irregular promi- nent glands to long-glandular-ciliate; veins 3(5), unbranched; laminar glands pale, linear to distally punctiform; marginal glands black, ± elongate, sessile or on cilia. Petals golden? yellow, not tinged red in bud, (8-) 10- 12 x (3-)5-6 mm, 2.4-4 x sepals, oblong-obovate, asymmetric, entire to subentire, laminar glands pale, ± punctiform, distal, marginal glands black, immersed or rarely ± prominent, distally dense. Stamens 40-50, '3 '-fascicled, longest 7-10 mm, 0.8- 0.9 x petals; anther gland black. Ovary 3-locular, 2-2. 5 (-3) x 1-1 .5 mm, cylindric-ovoid; styles 3, free, 4-6 mm, 1.7-3 x ovary, widely spreading; stigmas narrowly capitate. Capsule (6)-7-8 x 4-5 mm, c. 1.5-2 x sepals, cylindric-ovoid to conic; valves rather densely longitudinally and obliquely vittate. Seeds yellowish-brown, 0.8-1 mm, cylindric, not carinate; testa finely linear-foveolate. 2n=32 (n=16, Chattaway, 1926). Well-drained habitats: meadows, stony and scrub steppes and stony slopes on chalk and limestone, Betula and Pinus-Betula woods on sand, shores of rivers and lakes; 0-500 m. Russia (E. Siberia: Angara-Sayan; W. Siberia: Ob (S.), Upper Tobolsk, Irtysh, Altai; European Russia: Volga-Kama, Transvolga, Lower Don, Ciscaucasia, Daghestan, Volga-Don, Upper Volga), Ukraine (Upper Dnieper [near Kremenets], Middle Dnieper, Upper Dniester, Black Sea, Crimea) [apparently absent from Transcaucasia, despite area records in Gorschkova (1949)]; Germany (Saxony, Rhineland), Czech Republic, Slovakia, Poland, Hungary, Romania, Moldova, Serbia, Bulgaria, Turkey (European) (?) RUSSIA. Angara-Sayan, Ob, Upper Tobolsk, Irtysh: all fide Gorschkova (1949). Altai: Chemal distr., Gora Biryutka, 29 June 1927 (fl), Shishkin s.n. (H). Volga-Kama, Transvolga: both fide Gorschkova (1949). Lower Don: Novotscherkassk, 9 June 1911 (fl), Jakuschev s.n. (BM, FR). Ciscaucasia: ad rivum Kuma prope pagum Maslow, July 1843 (fl), Hohenacker s.n. (BM, K); am Terek bei Galjugajer, August (fl & fr). Herb. Steven s.n. (H). Daghestan : fide Grossgeim (1962: map 191). Volga-Don: Prov. Kursk, prope urbem Koroczka, 1 July 1900 (fl), Schirjewsky 1604 (H). Upper Volga: fide Gorschkova (1949). UKRAINE. Upper Dnieper: fide Gorschkova (1949). Middle Dnieper: Poltava, n.d (fl), Rogouris s.n. (H). Upper Dniester: Probabin prope Horodenka [Gorodenka], n.d. (fl), Blacky s.n. (BM). Black Sea: Crimea, Simferopol, prope pagum Massanka, 5 July 1900 (fl), Collier 901 (JE, K); Tarkhankutskiy peninsula, balka Onsina near Otlesh, 4 June 1 935 (fr), Dzens- Litovskaya s.n. (LE). AZERBAIJAN?. E. Transcaucasia, fide Gorschkova (1949), but not Grossgeim (1962: map 191). GEORGIA?. W. Transcaucasia, fide Gorschkova (1949), but not Grossgeim (1962: map 191). GERMANY. Upper Saxony: Erfurt, Schwellenburg, 23 July 1852 (fl), Launert s.n. (BM); Erfurt, Miihlhausen, Weinberg, 1951, Meyer s.n. (JE); Halle, Sachsenburg, 1906(fl), Kappels.n. (BM); Halle, Rossleben, Bottendorf, \93\,Rothmalers.n. (JE). Rheinhessen-Pfalz: AlzeyerHiigelland, Wissberg bei Sprendlingen, W-Seite des Gipfelplateaus, 260 m, 9 July 1978 (fl), Kalheber 78-477 (H). CZECH REPUBLIC. Moravia: Jimoravsky Kraj, Hodonin distr., inter pagos Cejc et Cejkovice, c. 220 m, 13 July 1971 (1. fl & fr), Vicherek 1539 (H); Severomoravsky Kraj, Vyskov, prope Vetrmky, June 1936 (fl), Weber s.n. (H). SLOVAKIA. Two records - see Zeleny et al. (1982: map 57). POLAND. Warbrzych: Dolny Slaj>ka, 35-40 km W. of Nysa (fide Kornas, 1955: 28). HUNGARY. Pest: Comit. Pest, in monte Csiki-hagy prope Torok-Babut, 3 June 1926 (fl), Degen s.n. (BM). ROMANIA. TVansylvania [Alba]: Langenthal [Valea Lunga], 18 June 1871 (fl), Earth 515 (BD, BM, H, JE); distr. Alba, inter pagos Ciumbrud et Bagau, c. 300-350 m, 1 4 June 1 930 (fl), Nydrddy s.n. (H). Moldavia: distr. Ia§i, in foenatis montis 'CiocSiau' prope pagum Rediu-Tatar, c. 150m, 21 June 1938 (fl), Rdvdrut in Fl. Rom. Exsicc. 2003a (BD). Wallachia: Dobrogea, Radu- Negru (by wellau Constanta o Burarqu), 1 1 July 193 1 (fl), Hayren s.n. (H). MOLDOVA. Bessarabia: a Aktherman en Bessarabie, 1822, Tardent s.n. (G-DC). SERBIA. Voj\odma:fide Stjep.-Vesel. (1972: 120). BULGARIA. Varna: Baltschik, Schiblak, oberhalb des Strandes, 1 5 June 1961 (fl), Bisse & Schneider 166 (JE). Plovdiv: M. Rhodope centr. supra Stanimak [Azenovgrad], 10 June 1 892, Wagner in Degen s.n. (JE); Rhodope in collibus ad Alvan Dere, June 1901 (fl), Stribmy s.n. (BM). TURKEY. Istanbul: Biiyiik Han, c. 100 m, Davidov s.n. See Davis, Fl. Turkey 2: 401 (1967). 7x. Hypericum elegans Stephan ex Willd. x H. perforatum L.? Raciborski s.n. (June 1904), from Ukraine, Podolia, colles Miodobory, Okno [Okna?] (BM) seems to be this hybrid, which has not hitherto been recorded. It is morphologically intermedi- ate between the two suggested parents. 8. Hypericum tosaense Makino in Bot. Mag. (Tokyo) 17:79(1903); H. Leveille in Bull. Soc. hot. France 53: 502 (1906); Makino & Nemoto, Fl. Japan: 545 (1925), 2nd ed.: 753 (1931); Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 221, ff. 73.4, 74 (1951), Enum. Sperm, jap. 3: 1 84 ( 1 954); Ohwi, Fl. Japan (Engl. transl.): 632 (1965); Horikawa, Atlas of jap. fl. 2: 681 (1976); Momiyama in Satake et al. (Eds), Wildfls Japan 2: 118, t. 114 f. 2 (1982). Type: Japan, Shikoku, Prov. Tosa [Kochi], Ikku-mura, 27 Sep- tember 1892 (fl & fr), T. Makino s.n. (MAK!-holotype). Map 18; also Kimura (1959: 324, f.l) and Horikawa (see above). H. tosaense f. tosaense Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 222(1951). STUDIES IN HYPERICUM 113 114 N.K.B. ROBSON A* A* A A A A Q Map 18 a. 8. H. tosaense •; b. 9. H. iwatelittorale •; c. 10. H. momoseanum O; d. 1 1 . H. yezoense A, other records A. H. tosaense f. insulare Y. Kimura in Nakai & Honda, Novafl. jap. 10: 222 (1951), Enum. Sperm, jap. 3: 184 (1954); Momiyama in Satake et al. (Eds), WildFls of Japan 2: 118 (1982). Type: Japan, Shikoku, [Kagawa] Ins. Syodo [Shodo], Ikedamura, 30 August 1910 (fl), R. Hirama s.n. (TI!-holotype). H. attenuatum pro parte sensu Kitamura & Murati in Acta Phytotax. Geobot.2Q: 197(1962). Perennial herb (0.15-)0.25-0.57(-0.75) m tall, erect from creeping and rooting base, with stems solitary, branched above. Stems 2- lined, with black punctiform glands on and near lines; internodes 10-20 mm, mostly exceeding leaves. Leaves sessile; lamina 5-20 x 2-8 mm, elliptic or ovate-oblong to narrowly oblong or obovate- oblong, paler beneath, chartaceous; apex obtuse to rounded, margin entire, base rounded to broadly cuneate; venation: 2(3) pairs of main laterals from lower quarter of midrib; laminar glands pale, puncti- form, dense; intramarginal glands black, dense. Inflorescence 3-c. 30-flowered from up to 3 nodes, with flowering branches from up to 4 nodes below, corymbiform to cylindric and then dense; pedicels 2- 3 mm; bracts and bracteoles small, linear-lanceolate to linear, entire. Flowers 15-18 mm in diam, stellate; buds ellipsoid, obtuse. Sepals 5, unequal, 2.5-3 x 1-1.5 mm, triangular-ovate to lanceolate, gradually to abruptly acuminate, entire; veins 5, outer pair(s) branched; laminar glands pale, punctiform to linear-striiform; intramarginal glands black, punctiform, spaced. Petals 5, bright yellow, veined red in bud, 9-15 x c. 5-8 mm, 3.5-5 x sepals, oblong- elliptic, somewhat asymmetric, distally ± markedly crenate; laminar glands pale, linear to punctiform, scattered; marginal glands black, distally dense. Stamens 60-70, '3 '-fascicled, longest 7-11 mm, 0.65-0.8 x petals; anther gland black. Ovary 3-locular, 2.5-^.8 x c. 1.5 mm, ovoid; styles 3, free, 3.5^.5(-5) mm, 1-1.6 x ovary, spreading-outcurved; stigmas narrow Capsule 7-9 x 3.5-5 mm, c. 3 x sepals, ovoid-conic to ovoid; valves longitudinally vittate. Seeds dark brown, c. 1 mm long, cylindric; testa linear- foveolate. 2n=16 (Kogi, 1984). 'Sunny places on hills'. Japan (Shikoku (Tosa area) and Shodo I.). JAPAN. Shikoko: Kagawa: Ins. Syoda, Ikeda-mura, 30 August 1910 (fl), R. Hirama s.n. (TI). Kochi: Prov. Tosa, Ikku-mura, 27 September 1892 (fl & fr), T. Makino s.n. (MAK). Hypericum tosaense occurs in two restricted area of southern Japan (south coast of Shikoku and Shodo Island between Shikoku and Honshu) and would appear to be a relict species linking the mainland H. attenuatum with the equally relict 9. H. iwatelittorale from northern Honshu, both morphologically and geographically. Forma insulare (from Shodo I.) differs from the type form in having narrower leaves, more flowers per inflorescence and longer styles (2 x rather than 1.2 x ovary), characters that, in this group, do not seem to warrant separate taxonomic recognition at more than the forma level, if at all. 9. H. iwatelittorale H. Koidz. in J. PL Iwateken II, 2: 89 (1937) ('iwate-littorale'). Type: Japan, Honshu, Prov. Rikutiu [Iwate], Shimohei-gun, Todogasaki, 12 August 1934 (fl & fr), Y. Fukuda 569 in Herb. H. Koidz. 95611 (TNS!-holotype); loc. cit., 12 August 1934 (fl & fr), Y. Fukuda 569 in Herb. H. Koidz. 95612- 95613 (TNS!-isotypes). Map 18. Perennial herb 0.25-0.32 m tall, erect, with stems solitary or few and caespitose, branched above. Stems 2-lined, with black puncti- form glands on lines; internodes exceeding to equalling leaves. Leaves sessile (main stem) to c. 0.5 mm petiolate (laterals); lamina (8-) 1 2- 1 4 x 5-8 mm, ovate to elliptic-oblong (main stem) or elliptic to oblanceolate (laterals), paler beneath, chartaceous; apex rounded STUDIES IN HYPERICUM 115 to rounded-obtuse, margin entire, base cuneate to rounded- amplexicaul (upper); venation: 3-(4-) pairs of laterals from lower quarter to fifth of midrib; laminar glands pale, punctiform, dense; intramarginal glands all black or some reddish, dense to spaced. Inflorescence (l)3-c. 30-flowered from up to 3 nodes, with flower- ing branches from up to 4 nodes below, corymbiform to cylindric and then dense; pedicels 0.8-1 mm; bracts small, lanceolate, entire. Flowers 9-10 mm in diam., stellate?; buds ellipsoid, subacute. Sepals 5, subequal, 3-4 x 1-2 mm, ovate-oblong to lanceolate, acute to subacuminate, entire; veins 5, outer pair(s) branched; laminar glands pale, linear-striiform to punctiform; intramarginal glands black, few or absent. Petals 5, bright yellow, not tinged red in bud, 6-9 x c. 2-3 mm, obovate-oblong, distally subcrenate; laminar glands pale, linear to punctiform, and sometimes 1-2 black, distal, striiform-punctiform; marginal glands black, very few or absent. Stamens c. 50?, '3'-fascicled, longest c. 7 mm, c. 0.8 x petals; anther gland black. Ovary 3-locular; styles 3, free, 3.5-4 mm, ? x ovary, spreading-outcurved; stigmas narrow. Capsule 5-8 x 4-5 mm, c. 2 x sepals, ovoid-conic to ovoid; valves longitudinally vittate. Seeds dark brown, 1.2 mm long, cylindric; testa linear-foveolate. 2n=? Coastal. Japan (Honshu-Iwate). JAPAN. Honshu. Iwate: Shimohei-gun, Todogasaki, 12 August 1934 (fl & fr), Y. Fukuda 569 in Herb. H. Koidzumi 95611 (TNS). Hypericum iwatelittorale was included doubtfully by Kimura ( 1 95 1 ) in H. pseudopetiolatum (subsect. 2. Erecta), but the gland-dotted raised stem lines reveal that its true position is in subsect. 1. Hypericum. It is morphologically very close to H. tosaense but considerably distanced geographically. Taken together, the small morphological differences (see key, p. 68) and the wide spatial separation warrant its recognition as a species. 10. Hypericum momoseanum Makino in/. Jap. Bot. 7: 12 (1931); Nemoto, Fl. Japan, Suppl.: 485 (1936); Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 218, f. 73.3 (1951), in J. Jap. Bot. 34: 323, f.l (1959); Momiyama in Satake et al. (Eds), Wild Fls of Japan 2: 118 (1982). Type: Japan, Honshu, Prov. Sinano [Nagano], Mt. Hachibuse, 18 July 1930 (fl), Momose s.n. (TI!- holotype). Map 18. H. momoseanum var. momoseanum Y. Kimura in Nakai & Honda, Novafl. jap. 10:219(1951). H. momoseanum var. atumense Y. Kimura in Nakai & Honda, Nova fl.jap. 10: 219 (1951); in Satake etal. (Eds), Wild fls Japan*. 118 (1982). Type: Japan, Honshu [Aichi], Prov. Mikawa, Atumi-gun, Fukue, Kantekiyama, 3 August 1939 (fl), Yamamoto s.n. (TI!- holotype). H. tosaense var. atumense (Y. Kimura) Y. Kimura in J. Jap. Bot. 34: 326(1959). H. attenuatum pro parte sensu Kitamura & Murati in Acta Phytotax. Geobot. 20: 197 (1962). H. yezoense var. momoseanum (Makino) Ohwi in Fl. Japan (Engl. transl.): 632 (1965). The original (Japanese) edition (Ohwi, 1953: 782) mentions H. momoseanum under H. yezoense but makes no formal combination. Perennial herb 0.25-0.65 m tall, ascending from woody base, with stems single, unbranched or with short sterile branches throughout. Stems 2-lined, with black punctiform glands on and near lines; internodes 1 5-30 mm, equalling leaves. Leaves sessile, amplexicaul; lamina 15-30 x 5-1 1 mm, oblong to elliptic or oblanceolate, paler beneath, thickly chartaceous; apex rounded-obtuse, margin plane, base rounded; venation: 3-4 pairs of main laterals from base and lower quarter of midrib, tertiary reticulation lax; laminar glands punctiform, pale (minute) and black, submarginal and distal; intramarginal glands black, dense. Inflorescence 7- 11 -flowered, from 2-3 nodes, with flowering branches from up to 4 nodes below, the whole subcorymbiform to condensed-cylindric; pedicels 0.7-3 mm (apical to 5 mm); bracts and bracteoles linear, entire. Flowers 1 5-20 mm in diam., stellate; buds elliptic, obtuse. Sepals 5, subequal, 3-5 x 0.7-2 mm, oblong-ovate to lanceolate or narrowly elliptic, acute to gradually acuminate, entire; veins 5, occasionally branched; laminar glands pale, linear to striiform, sometimes also a few punctiform and occasionally a few black, distal, punctiform; intra- marginal and submarginal glands black, spaced. Petals 5, 'yellow', 8.5-12 x 2.5-3.5 mm, 2.5-3 x sepals, obovate to oblanceolate, subsymmetric, distally not or scarcely crenate; laminar glands pale, linear to punctiform and sometimes a few black, distal, punctiform. Stamens c. 40, '3'-fascicled, longest 7-8 mm, 0.7-0.85 x petals; anther gland black. Ovary 3-locular, c. 3.5 x 2 mm; styles 3, free, 4- 5 mm, 1.2-1.4 x ovary, 'erecto-patent' ; stigmas narrowly capitate. Capsule and Seeds not seen. 2n= ? Habitat not recorded. Japan (central Honshu- two localities). JAPAN. Honshu. Nagano: Mt. Hachibuse, 18 July 1930 (fl), 5. Momose s.n. (TI). Aichi: Atsumi-gun, Fukue, Kantekiyama, 3 August 1939 (fl), S. Yamamoto s.n. (TI). In habit and distribution H. momoseanum is intermediate between the southern H. tosaense and the northern H. yezoense. Like the former species it has a very restricted distribution, being confined to two stations in central Honshu (Nagano and Aichi Prefectures). Kimura (1959) described var. atumense as having stems unbranched or almost so and larger flowers than var. momoseanum, in which the stems are branched (as in H. yezoense). The differences do not seem to warrant taxonomic recognition. 1 1 . Hypericum yezoense Maxim, in Bull. Acad. Imp. Sci. Saint- Petersbourg 31: 16 (1886), Mel. Biol. Bull. Acad. Imp. Sci. Saint-Petersbourg 12: 420 ( 1 886); Matsumura, Index pi. jap. II- 2: 370 (1912); Miyabe & Miyake, FL. Saghalin: 79 (1915); Y. Kimura in Bot. Mag. (Tokyo) 51: 701, ff. 20-21 (1937), in Nakai & Honda, Novafl. jap. 10: 216, f. 73.1 (1951), Enum. Sperm, jap. 3: 1 86 ( 1 954), in J. Jap. Bot. 34: 323, ff. 1 , 2 ( 1 959), in Asahi ... shokobutsu, no. 64: 1508 (1977); Sugawara, Ill.fl. Saghalien 3: t. 600 (1940); Gorschkova in Shishkin & Bobrov, Fl. URSS 15: 238 (1949); Ohwi, Fl. Japan (Engl. trans.): 632 (1965); Lauener in Notes Roy. Bot. Card. Edinburgh 27: 5 (1966); Kitamura & Murati, Clrd Ills Herb. Pis Japan 2: 66 (1972). Type: Japan, Yezo [Hokkaido, S. Oshima], circa Hakodate, 20 July 1878 (fl), R. Yatabe s.n. (LE-holotype; TI!-isotypes). Fig. 11, Map 18; also Kimura (1959: 324, f. 1). H. attenuatum [var.] fifruticulosum F. Schmidt in Mem. Imp. Acad. Sci. Saint-Petersbourg, 7e sen, 12(2): 119 (1868) ['fruticulosa']. Types: Russia, Sachalin, Sachkotau, 18 July 1860 (fl), Friedr. Schmidt s.n. (LE-syntype); loc. cit., late August 1860 (fr), Friedr. Schmidt s.n. (LE-syntype). H. procumbens R. Keller in Bull. Herb. Boissier 5: 639 (1897); H. Leveille in Bull. Soc. Bot. France 53: 502 (1906); Matsumura, Index pi. jap. II-2: 370 (1912); Miyabe & Miyake, Fl. Saghalin: 79 (1915); Makino & Nemoto, Fl. Japan: 546 (1925); non Desf. ex Willd. (1802) nee Michx. (1803). Type: Japan, Hokkaido, 'in 116 N.K.B. ROBSON Fig. 11 A. H. scouleri: (a) habit; (b) sepal; (c) petal; (d) capsule. B. H. scouleri 'nortoniae': (e) habit. C. H. yezoense: (f) habit; (g) sepal; (h) petal; (i) capsule (a, e, f x 2/3; c, d, h, i x 4; b, g x 8). A. (a) Copeland 426; (b, c) Kellogg & Harford 102; (d) Jones 5682. B. (e) Moore & Steyermark 37 1 1 . C. (f, g, h) Furitse 940; (i) Faurie 6905. STUDIES IN HYPERICUM 117 promentorio Gangenzaki', 7 June 1894 (fl), Faurie 13325 (G- holotype?; FI!, K!). H. mororanense R. Keller in Bull. Herb. Boiss. 5: 640 ( 1 897), in Bot. Jahrb. Syst 33: 553 (1904), op. cit. 44: 49 (1909), op. cit. 58: 194 (1923) in obs. ['morarense']; H. Leveille in Bull. Soc. Bot. France 53: 502 (1906), op. cit. 54: 595 (1908); Matsumura, Index pi jap. II-2: 367 (1912); Makino & Nemoto, Fl. Japan: 543 (1925), 2nd ed. 2: 753 (1931). Type: Japan, Hokkaido, Iburi, Mororan [Muroran], Faurie 10304 (Z?-lectotype, selected here); Hokkaido [Shiribeshi], Promontorium Asushi, Faurie 5061 (Z?- syntype; K!). H. oliganthemum R. Keller in Bot. Jahrb. Syst. 33: 553 (1904); H. Leveille in Bull. Soc. Bot. France 53: 502 (1906) ['iloganthum']; Matsumura, Index pi. jap. 11-2: 367 (1912); Makino & Nemoto, Fl. Japan: 544 (1925), 2nd ed.: 751 (1931). Type: Japan, precise locality not cited, Rein s.n. (Bt-holotype). H. mororanense forma tetragynum H. Leveille in Bull. Soc. Bot. France 53: 502 (1906); Matsumura, Index pi. jap.: 367 (1912). Type: Japan, Hokkaido [Soya], in rupibus maritimis Rebunshiri, 1 August 1889, Faurie 3086 (P?-holotype). Kimura (1937: 702) pointed out that flowers with abnormal numbers of parts, such as occur in the type of this variety, are not uncommon in northern or alpine regions of Japan. H. porphyrandrum H. Leveille & Vaniot in Repert. Spec. Nov. Regni Veg. 6: 330 (1909); Matsumura, Index pi. jap. II-2: 368 (1912); Makino & Nemoto, FL Japan: 544 (1925), 2nd ed.: 751 (1931); Gorschkova in Shishkin & Bobrov, Fl. URSS 15: 257 (1949). Type: Russia: Sakhalin I., in rupibus Tonaichan, 18 September 1908 (fl), Faurie 521 (P?-holotype; KYO!-isotype). ?//. pseudo-nikkoense H. Koidz. in J. PL Iwateken 2(2): 87 (1937). Type: Japan, Honshu, Prov. Rikutiu [Iwate], Geibikei, G. Toba in Herb. H. Koidz. 95578 (TSN?-holotype). ?//. yoitiense H. Koidz. in J. PL Iwateken 2(2): 101 (1937). Type: Russia, Kurile Is., Kunashir, Yoito, /. Yamamoto in Herb. H. Koidz. 102121 (TNS?-holotype). Icones: S. Sugawara, Illustr.fl. Saghalien 3: t. 600 (1940); Y. Kimura in J.jap. Bot. 34: 325, f. 2 (1959). Perennial herb 0.1-0.3(-0.35) m tall or long, erect to ascending or more rarely procumbent from creeping and rooting base, with stems numerous to few and caespitose or rarely solitary, branched or not. Stems 2-lined (if slender almost 4-angled), with black punctiform glands on lines only; internodes (4-) 12-28 mm, usually exceeding leaves. Leaves sessile; lamina 8-20(-25) x 3-9 mm, ovate (upper- most) to elliptic-lanceolate or ± narrowly oblong to oblanceolate, slightly paler beneath, chartaceous; apex obtuse to rounded, margin plane, base broadly cuneate to subamplexicaul; venation: 2-3 pairs of main laterals from lower quarter of midrib; laminar glands pale, punctiform, dense and black, punctiform, few, towards margin, or absent; intramarginal glands black, dense. Inflorescence 3-9-flow- ered from 1-2 nodes, without or with one pair of flowering branches from node below, corymbiform to subpyramidal; pedicels 0-1 mm (that of terminal flower to 4 mm in fruit); bracts reduced-foliar, bracteoles 4-5 mm long, linear-lanceolate, entire. Flowers 15-20 mm in diam., stellate; buds narrowly ovoid, subacute. Sepals 5, unequal, 4-6 x 1.2-2 mm, erect in bud and fruit, lanceolate to oblong-lanceolate, acute, entire; veins 3-5, unbranched; laminar glands pale, punctiform and sometimes a few black, punctiform, distal; intramarginal glands black, punctiform, rather sparse, distal. Petals 5, pale yellow, not? tinged red in bud, 8-12 x 3.5-5 mm, 2 x sepals, obovate-oblong, asymmetric, subentire; laminar glands pale, linear to punctiform, scattered; marginal and intramarginal glands black, irregular, locally dense. Stamens c. 30-50, '3'-fascicled, longest c. 8-10 mm, c. 0.9 x petals; anther gland black. Ovary 3- locular, 3-3.2 x 1 .5-2 mm; styles 3, free; stigmas narrowly capitate. Capsule 5-8 x 3? -4 mm, c. 1 .3 x sepals, broadly to narrowly ovoid; valves longitudinally vitiate. Seeds dark brown, 1-1.3 mm, cylin- dric; testa linear- foveolate. 2n=? Habitat not recorded. Japan (extreme northern Honshu, Hokkaido), Russia (southern Sakhalin; Kuriles - Kunashiri and Shikotan Is.). JAPAN. Honshu. Iwate: Motomura, Akka-mura, Shimohei-gun, 400 m, 13 June 1957 (bud), Shimizu 2052 (K). Aomori, Akita: see Kimura (1959: 324, f. 1). Hokkaido. Oshima: Eramachi, 22 July 1890 (1. fl), Miyabe & Tokubuchi s.n. (K); *Esan Isoya, 26 July 1926, Tatewaki 6110 (SAPS?). Shiribeshi: in Shiribeshi, August 1905 (fl), Faurie 6905 (BM); "Isoya, 18 July 1888, Y. Tokubuchi s.n. (TI?). Ishikari: Hassabu, August \9l6(fl\Kudo in HTU 77274 (TAI); *Mt. Tengu, 15 August 1931, Yamamoto s.n. (SAPS?). Iburi: *Mororan, Faurie 10304 (P?). Hidaka: *Iwayama, Samani, 18 June 1884, K. Miyabe s.n (TI?). Rumoi, Abashiri: see Kimura (1959: 324, f. 1). Soya: Rebun I., Funado-mari-choo, 13 August 1975 (1. fl), Furuse 9407 (K). Nemuro: Hanasaki, prope urbem Nemuro, 7 July 1909 (1. fl), Takeda s.n. (K). RUSSIA. Sakhalin: *Toyohara [Yuzhno-Sakhalinsk], Takinosawa, 2 August 1928, N. Hiratsuka s.n. (TI?); *Makunkotan circa Motodomari [Tomari?], 1 5 September 1 906, T. Miyake (TI?). Kurile Is.: *Kunasiri, 26-27 July 1935, Y. Takahashi s.n. (TI?); Shikotan, Notoro, 2 August 1931, Ohwi s.n. (KYO?). Hypericum yezoense is a reduced 'form' of the central Honshu 10. H. momoseanum, differing from it in habit and inflorescence size. Kimura (1937: 703) notes that the Korean record for this species is erroneous, as is Maximowicz's description of the ovary as one- celled. 12. Hypericum scouleri Hook., Fl. bor.-amer. 1: 111 (1831), PL Hartweg.: 301 (1848); Torrey & Gray, FL N. Amer. 1: 160 (1838); Torrey in Emory, Rep. U. S. Mex. bound. 2(1): 36 (1858); Greene, FL francisc. 1: 112 (1891), Man. hot. San Franc. Bay: 63 (1894); R. Keller in Engl. & Prantl, Nat. Pflanzenfam., 2nd ed., 21: 179 (1925); Peck, Man. pis Oregon: 481 (1941); Gillett & Robson in Publs. Bot. Natl. Mus. Nat. Sci. Canada no. 1 1: 28, t. 12, map 9 (1981); Scott, Alpine FL Rocky Mts 1: 458 + f. + map (1996). Type: U.S.A., [Oregon] 'Nw. coast of America, near the Columbia', n.d. [1825] (fl), Scouler s.n. (KMectotype, selected here; NY-isolectotype); [Washington?] grassy plains of the Columbia, 1825 (fl), Douglas s.n. (BM!- syntype); ex eodem loco, 1 826 (fl), Douglas s.n. (BM !-syntype). Fig. 11, Map 19. H. skouleri sensu Walp., Rep. hot. syst. 1: 387 (1842), sphalm. H. formosum sensu A. Gray, PI. wright. 2: 17 (1853); Torrey in Emory, Rep. U. S. Mexico bound. 2(1): 36 (1858); Kearney & Peebles, Arizona fl.: 556 (1951); Clark, Wildfls Pac. Northwest: 312, 322 (1976); Dorn, Man. Vase. Pis Wyoming (2): 818 (1977), Vase. Pis Wyoming: 184 (1988); Scoggan, FL Canada 3: 1096 (1978); Martin & Hutchins, FL New Mexico (2): 1280 + f., map 758 (1981); Welsh etal., Utah FL, rev. ed.: 353 (1993); non Kunth (1822). H. formosum var. scouleri (Hook.) Coult. in Bot. Gaz. 11: 108 ( 1 886); Abrams, ///. FL Pac. States 3:116, f. 3233 ( 195 1 ); Munz, Calif. FL: 192 (1959); Scoggan, Fl. Canada 3: 1096 (1978). H. simulans Rose in Contr. U. S. Natl. Herb. 10: 124 (1906); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed., 21: 179 (1925). Type: Mexico, Hidalgo, Canales Station, 1800 m, 8 August 1904 (fl), Pringle 8993 (US!-holotype; BM!, C!, F!, G!, GH!, K!, MEXU!, NY!, P!, W!-isotypes). H. collinum var. schmitzii R. Keller in Bull. Herb. Boissier II, 8: 1 87 118 (1908). Type: Mexico, Chihuahua, Potrero passada la Piedat, 1856? (fl), Schmitz s.n. (W!-holotype, F!, US!-photographs). H. nortoniae M.E. Jones in Bull. Univ. Montana no. 61, Biol. Ser. no. 15: 39, t. 5 (1910) ['Nortonae'}. Type: U.S.A., Montana, McDonald Peak, 2190 m, 2 September 1908 (fl), M.E. Jones s.n. (POMI-lectotype, selected here). H. formosum subsp. scouleri (Hook.) C.L. Hitchc., Vase. Pis Pac. N.-W. 3: 434 + f. (1961); Taylor & Macbryde, Vase. Pis Br. Columbia: 203 (1977); Thorne in Aliso 9: 193 (1978), independ- ent comb.; E. Murray in Kalmia 12: 21 (1982), independent comb.; Moss, Fl. Alberta, 2nd ed. rev. Packer: 404, map 765 (1983). H. formosum var. nortoniae (M.E. Jones) C.L. Hitchc., Vase. Pis Pac. N. W. 3: 434 + f. (1961) ['nortonae']; Scoggan, Fl. Canada 3: 1096(1978). H. formosum subsp. formosum sensu C.L. Hitchc., Vase. Pis Pac. N. W. 3: 434 ( 1 96 1 ); McVaugh, Fl. Novo-Galicia 3: 5 1 (200 1 ), pro parte quoad loc. s. w. US; non H. formosum Kunth (1822). H. formosum var. formosum sensu Dorn, Man. Vase. Pis Wyoming (2): 818 (1977), Vase. Pis Wyoming: 184 (1988); Scoggan, Fl. Canada 3: 1096 (1978); non H. formosum Kunth (1822). H. scouleri subsp. nortoniae (M.E. Jones) J.M. Gillett in Canad. J. Bot. 57: 185 (1979); Gillett & Robson in Publs. Bot. Natl. Mus. Nat. Sci. Canada no. 1 1 : 30, map 9 ( 1 98 1 ). Type as for H. norton- iae M.E. Jones. H. scouleri subsp. scouleri Gillett & Robson in Publs. Bot. Natl. Mus. nat. Sci. Can. no. 11: 30 (1981). Icones: Gillett & Robson in Publs. Bot. Natl. Mus. Nat. Sci. Canada no. 11: 29, t. 12 (1981); C.L. Hitchc., Vase. Pis Pac. N.-W. 3: 434 (1961). Perennial herb 0.05-0.6(-0.8) m tall, erect or ascending from creeping and branching rhizomatous and often stoloniferous base, with stems solitary or up to c. 10, not caespitose, sometimes with short ascending stem branches. Stems shallowly 2-lined or often partially 4-lined (with evident complete or incomplete subsidiary lines) or rarely almost terete, eglandular or sometimes with a few reddish punctiform glands usually on or near lines; internodes 15- 50 mm, usually exceeding leaves. Leaves all sessile or lower subpetiolate, spreading or rarely erect; lamina 12-28(-32) x 6-15(- 1 8) mm, oblong-elliptic or elliptic to triangular-ovate or suborbicular or (lower) obovate, paler beneath, chartaceous; apex obtuse to rounded, margin plane, base rounded to subcordate or (lower) cuneate; venation: 4-5 pairs of basal and lateral main veins from lower third of midrib; tertiary reticulation rather dense but obscure to lax; laminar glands pale, punctiform, small to large and some- times 1-2 or more (rarely all) black, distal and submarginal; intramarginal glands black, dense. Inflorescence (l-)8-20-flow- ered from 1-3(4) nodes, sometimes with flowering branches from 1-2(3) nodes below, the whole narrowly cylindric to pyramidal; pedicels 2-6 mm; bracts and bracteoles 3-6 mm long, ovate-lanceo- late to ovate, entire. Flowers 6-15(-25) mm in diam., stellate; buds broadly ellipsoid to subglobose, obtuse to rounded. Sepals 5, un- equal to subequal, 2.5-5.5 x 1-2 mm, erect in bud and fruit, c. 0.25 united, ovate to lanceolate or narrowly oblong, acute to rounded, entire or sometimes with a few sessile glands or regularly or irregu- larly shortly glandular-ciliate to -denticulate; veins 5, distally branched; laminar glands pale, striiform and sometimes black, punctiform to striiform or linear; intramarginal and marginal glands black, irregular or distally regular. Petals 5, golden yellow, some- times tinged red in bud, 7-12 x 3-7 mm, ± narrowly obovate, asymmetric, distally subcrenate to entire; laminar glands pale, linear to striiform, occasionally a few black, punctiform to striiform; N.K.B. ROBSON marginal glands black, distally dense. Stamens c. 50-90(-103), '3'- fascicled, longest 6-10 mm, c. 0.7-0.9 x petals; anther gland black. Ovary 3(4)-locular, 3-5 x 2-2.5 mm, ovoid; styles 3, free, (2-)3- 6(-8) mm, c. 1-1.7 x ovary, widely spreading; stigmas narrowly capitate. Capsule 6-10 x 3.5-6 mm, ovoid to narrowly ovoid- pyramidal; valves densely longitudinally vittate. Seeds brown, (0.5-)0. 7-0.8 mm, cylindric, not carinate or appendiculate, some- times? apiculate; testa linear-reticulate. 2n=16, n=8 (Kyhos, 1967; Ward & Spellberg, 1983). Wet meadows, bogs and grassy or rocky banks, stream and lake margins, thickets, coniferous forest, screes; Canada: c. 50-1800 m; U.S.A.: 400-2700 m; Mexico: 1800-3000 m. Canada (British Columbia (N. to c. 55°), extreme SW Alberta), western U.S.A. (E. to Montana, Wyoming, Colorado, New Mexico), Mexico (Baja California Norte and Sierra Madre Occidental from Sonora, Chihuahua and Coahuila to Michoacan, Mexico and Vera Cruz [Orizaba]). CANADA. British Columbia: Kokanee Glacier Park, 1800m, 15 August 1 938 (fl), Moir 276 (K); Selkirk and Rocky Mts, near 5 1 ° 30' N, Spillamasheen Valley, 1800 m, 30 July 1904 (fl), Heacock 41 1 (BM, JE, K, Z); Vancouver Island, Cameron Lake, near Port Alberni, 181 m, 5 August 1965, Ashlee 215 (H). Alberta, fide Gillett & Robson (1981: map 9). U.S.A. Washington: Douglas Co., near Egbert Spring, 390m, 2 July 1993 (fl), Sandberg & Leiberg 361 (BM, K, US*). Oregon: Klamath Co., summit of Cascade Mts, Crescent Lake, 1251 m, 17 August 1 934 (fl), Constance 977 (K);Harney Co., Shirk, 1500m, 19 July 1 996 (fl&e.fr), Leiberg 2598 (BM, US*). Montana: Gallatin Co., Bozeman, Middle Creek, 14 July 1905 (fl), Blankinship 98 (BM, H); Mineral Co., Fish Creek, near mouth of Cache Creek, 1050 m, 1 1 July 1923 (fl), Hitchcock 1696 (K). Idaho: Latah Co., 6.4 km E. of Farmington, 28 June 1892 (fl), Sandberg, MacDougal & Heller 512 (BM, K, Z); Shoshone Co., 6.4 km E. of Burke, c. 1620 m, 12 August 1949 (fl), Cronquist 6069 (G). Wyoming: Teton Co., Grand Teton National Park, cirque above Indian Paint Brush Canyon, 2750 m, 220 August 193 1 (fl & fr), Moore & Steyermark 3711 (BM, W); Park Co., Lewis R., Yellowstone National Park, 9 August 1 899 (fl), A. & E. Nelson 6388 1 (BM, K). Colorado: Montezuma Co., Mancos, 9 July 1 898 (fl), Baker, Earle & Tracy 889 (BM, K, W); El Paso Co., Colorado Springs, 1650 m, June 1879 (fl), 6.4 km NW of Brighton, 2220 m, 4 August 1964 (fl), Bennett 8440 (W). Utah: Pluto Co., Marysvale, Tate Mine, 2700 m, 2 August 1894 (fr), M.E. Jones 5862 (BM); Salt Lake Co., Big Cottonwood Creek canyon, Wasatch Mts, 6.5 km NW of Brighton, 2220 m, 4 August 1964 (fl), Bennett 8440 (W). Nevada: White Pine Co., southern Schell Creek Range, E. of Mt Grafton, 2250 m, 1 1 August 1 969 (f[), Holmgren 3869 (W); Douglas? Co., L.Tahoe, 1866m, 29 July 1931 (fl), L.S. Rose s.n. (JE). California: Butte Co., Jonesville, 1550 m, 18 July 1930 (fl), Copeland 426 (BM, H, K, TAI, W, Z); El Dorado Co., Sierra Nevada Mts, Placerville-Tahoe road, 0.8 km W. of Lover's Leap, 13 July 1935 (fl), Belshaw 1158 (K); San Bernardino Co., Lythe Creek canyon, 14 July 1902 (fl), Abrams 2727 (K, Z). Arizona: Coconino Co., Bill Williams Mtn, 2 1 00 m, 22 July 1 898 (fl), MacDougal 3 1 9 (G); Cochise Co., Chiricahua Mts, Barfoot Park, 2400-2475 m, 26 September 1906 (fr), Blumer 1426 (K, W, Z). New Mexico: Caltron Co., Gila National Forest, Mogollon, Willow Creek Rec. Area, 2640 m, 26 July 1963 (fl), Demaree 48702 (BM); Santa Fe Co., Santa Fe, Creekbottom, 1847 (fl), Fendler 53 (BM, K, W). MEXICO. Baja California Norte: Sierra San Pedro Martir, 2100 m, 21 August 1967 (e. fr), Moran & Thome 14453 (CAS, DS). Sonora: San Jose Mts, 8 km S. of Naco, 1 800 m, 6 July 1 928 (fl), Wolf 25 16 (GH). Chihuahua: Sierra Madre near Colonia Garcia, 2250 m, 5 August 1 899 (fl), Townsend & Barber 224 (BM, F, G, GH, K, MEXU, MICH, MO, NY, P). Coahuila: Muzquiz, Hacienda Mariposa, foot of E. slope of Sierra de Puerto Santa Ana, 23 June 1936 (fr), Wynd & Mueller 245 (K). Durango: Metates, N. of Cueva, 2650-2700 m, 29-30 August 1934 (fl), Pennell 18384 (US). Zacatecas: Sierra Madre, August 1897 (fl), Rosen 2382 (NY, US). Hidalgo: Sierra de Pachuca, 2850 m, 4 August 1898 (fl), Pringle 6941 (BM, C, F, G, GH, K, MEXU, MICH, MO, NY, US, W). Mexico: Lagunas de Zempoala Nat. Park, 3 August 1949 (fl), McAdams 88 (MICH). Distr. Fed.: Desierto de los Leones, 23 September 1938 (fl), Lyonnet 2586 (US). Michoacan: 6.4 km W. STUDIES IN HYPERICUM -•* • > * * ^ € • • /^ Map 19 12. H. scouleri •, other records O. of Hidalgo, c. 2000 m, 17 July 1940 (fl), Hitchcock & Stanford 7194 (F, GH, NY). Vera Cruz: Orizaba, 1846 (fl), Heller 143 (W). Many authors have included H. scouleri in the central Mexican H. formosum Kunth as a synonym, subspecies or variety, but the similarities in sepal form and glandularity of these species are due to convergence. Hypericum formosum is related to an undescribed subspecies of the Mexican H. oaxacanum R. Keller (sect. 9b. Graveolentia), whereas the affinities of H. scouleri are with the eastern Asian 6. H. attenuatum. Gillett (1979) and Gillett & Robson (1981) treated the dwarf, small-flowered alpine form of H. scouleri as a distinct subspecies, subsp. nortoniae (M.E. Jones) J.M. Gillett. Further work, however, has revealed a range of intermediate forms between the two 'subspe- cies' that prevents their recognition. Likewise, the differentiation of a southern U.S. population ('H. formosum subsp. formosum'' sensu C.L. Hitchc.) from a northern one ('H. formosum subsp. scouleri (Hook.) C.L.Hitchc.'), based on the 'broader, blunter and less glandular sepals' of the latter, does not appear to be warranted. The glandular, often acute sepals of this New Mexican form are an extreme development in H. scouleri, not an indication of affinity with H. formosum. In Mexico the sepals become more black- glandular south-eastwards, so that, in the plants known as H. simulans, they have black linear laminar glands; but the margin is always entire. ACKNOWLEDGEMENTS. Once again, as for Part 4(1), I am especially indebted to Dr Peter Raven and Dr Ihsan Al-Shebaz (MO) for help with loans and for hospitality during my studies of Chinese Hypericum in St. Louis, to Yang Chiang (NAS) for label translation and for compiling a database for me, and to Prof. Li Xiwen (KUN) for his publications on Chinese Hypericum. I am also very grateful to Prof. Hideaki Ohba (TI) for facilitating loans of Japanese specimens. My studies of western Eurasian species have been helped particularly by Dr Pavol Martonfi (KO), whose work on hybridity in the H. perforation group has thrown considerable light on aspects of this difficult complex; and Prof. Jacques Lambinon (LG), Prof. Fabio Garbari and Dr Daniela Ciccarelli (both PI) have all helped me to clarify the variation in H. perforatum itself. As before, I have obtained assistance from many colleagues at the BM, particularly from Mike Gilbert (computing matters, nomenclature and general discussion) and Prof. Chris. Humphries and Josephine Camus (mapping); and Malcolm Beasley and his colleagues in the Botanical Library have, as usual, been very helpful. My thanks are also due to the directors of the following herbaria for loans and/or study facilities: A, ANG, ANK, BASBG, BAY, BC, BD, BISH, BP, BRNM, C, CAS, CL, CLF, DS, E, EVIN, F, Fl, FR, G, GH, GL, GRM, GZU, H, HNWP, IBSC, JE, K. KO, KUH, KUN, KYO, L. LANC, LD, LE, LINN, LIV, MAK, MARS, MEXU, MICH, MO, NAS, NY, P, PE, POM, PR, RAW, RNG, S, SEV, SZ, TAI, TI, TNS, TO, U, UBC, UPS, US, W, WAG, WIS, WU, Z. I am again deeply indebted to Margaret Tebbs for the drawings. My wife Eve has.as always, been a help and support in many ways. Finally, I must thank the Keeper of Botany, Prof. Richard Bateman, and his immediate predecessors for study facilities in the Botany Department, and the Stanley Smith Trust for a grant towards illustration costs. REFERENCES Anthony, J. 1976. John Anthony's Flora of Sutherland (Ed. J.B. Kenworthy). Edin- burgh. Ascherson, P. 1 864. Flora der Provinz Brandenburg. Berlin. Axelrod, D.L. 1959. Geological history. In P. A. Munz & D.D. Keck, A California Flora: 5-9. Berkeley. Bonnet, E. 1879. Revision des Hypericum de la section Holosepalum Spach. Bulletin de la Societe botanique de la France 25: 274-282. Celakovsky, L. 1875. Prodromus der Flora von Bohmen 3. Prague. Chattaway, M.M. 1926. 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Holosepalum (Spach) Fourr. 73, 81 dubium (Leers) Fourr. (=lc) 73 quadrangulum (L.) Fourr. (=3) 81 Hypericum L. 61 Sect. Concinna N. Robson 61 Sect. Euhypericum Godr. 67 Subsect. Heterotaenium R. Keller 67 Series Perforata Gorschk. 67 Subsect. Homotaenium R. Keller 67 Series Acuta Gorschk. 67 Series Attenuata Gorschk. 67 Series Crispa Gorschk. 67 Series Elegantia Gorschk. 67 Series Kamtschatica Gorschk. pro parte 67 Series Quadrangula Gorschk. 67 Sect. Graveolentia N. Robson 61 Sect. Homotaenium Y. Kimura 67 Series Bilineata Y. Kimura 67 Series Crispa Y. Kimura 67 Series Quadrangula Y. Kimura 67 Sect. Hypericum 61, 62, 65, 66, 67 Subsect. Erecta N. Robson 61, 62, 66, 115 Subsect. Hypericum 61, 62, 65, 66 Series Hypericum 61-66,67 Series Senanensia 61, 62, 65, 66, 68 Sect. Roscyna (Spach) R. Keller 61, 62, 63, 65 acutum Moench (=3) 80, 81 subsp. boeticum (Boiss.) Cout. (=2b) 80 subsp. corsicum (Steud.) Rouy (=3b) 84 van rotundifolium sensu Briq. (=3b) 84 subsp. desetangsii (Lamotte) Rouy (=5xa) 104 var. genuinum (Bonnet) Rouy (=5xa) 104 var. imperforatum (Bonnet) Rouy (=lc) 73 subsp. eu-acutum Hayek (=3a) 81 subsp. tetrapterum (Fr.) Maire (=3a) 81 subsp. undulatum (Schousb. ex Willd.) Rouy (=2) 78 var. anagallidifolium (Boiss.) Post (=3c) 84 var. anagallidioides (Jaub. & Spach) Zohary (=3c) 84 var. corsicum (Steud.) Halacsy (=3b) 84 var. insulare (Foucaud & Mandon) Briquet (=3b) 84 var. rotundifolium (Willk.) Schinz (=2b) 80 var. typicum Gu§ul. (=3) 81 var. undulatum (Schousb. ex Willd.) Pau (=2) 78 forma humile (Boenn.) Hegi (=3) 81 forma patulum (Boenn.) Hegi (=3) 81 acutum x perforatum (=5x) 108 aetnaeum Tornab. (= 5c) 97 alatum Retz. (= 3?) 80 anagallidifolium Boiss. (= 3c) 84 anagallidioides Jaub. & Spach (= 3c) 84 ascyron L. 61, 62 subsp. ascyron 62 subsp. gebleri (Ledeb.) N. Robson 62 subsp. pyramidatum (Aiton) N. Robson 62 assurgens Peterm. (=5a) 89, 118 attenuatum Fisch. ex Choisy (6) 62, 63, 65-67, 88, 89, 108, 109*, 111, 114, 119 var. fruticulosum F. Schmidt (=11) 115 attenuatum sensu Kitamura & Murati (=6+8+10) 114, 115 baeticum sensu auct. plur. (=2b) 78 boeticum Boiss. (=2b) 79, 80 sensu Syme (=2) 78 borbasii Formanek (=3) 81 x carinthiacum A. Frohl. (=5xb) 105 forma maculatiforme A. Frohl. (=5xba) 65, 106 forma perforatiforme A. Frohl. (=5xbb) 65, 107 carpaticum Martonfi (=5xbb) 107 chrysostictum Webb (=5c) 96 collinum var. schmitzii R. Keller (=12) 117 commutatum Nolle (=lb) 71, 76 confertum (Choisy) G. Don (=3) 81 corsicum Steud. (=3b) 84 crispum L. (=4) 67, 84 decipiens H. Watson (=2) 78 deidesheimense Sch. Bip. ex Trev. (?=5a) 89 delphinense Vill. (=lb) 71, 73 x desetangsii Lamotte (5x) 63, 64, 76, 78, 102, 103* nothomorph carinthiacum (A. Frohl.) N. Robson (=5xb) 105 nothomorph desetangsii sensu N. Robson (=5xa) 104 nothomorph perforatiforme (A. Frohl.) N. Robson (=5xbb) 88, 107 nothosubsp. balcanicum N. Robson (5xc) 64. 103*. 104, 107 nothosubsp. carinthiacum (A. Frohl.) N. Robson (5xb) 64, 65, 104, 105 nothoforma maculatiforme (A. Frohl.) N. Robson (5xba) 65, 103*. 106 nothoforma perforatiforme (A. Frohl.) N. Robson, (5xbb) 65. 103*. 106. 107 122 N.K.B. ROBSON nothosubsp. desetangsii (5xa) 64, 103*. 104 nothovar. desetangsii sensu N. Robson (=5xa) 104 van genuinum Bonnet (=5xa) 104 var imperforatum Bonnet (=lc) 73, 74 desetangsii sensu Lloyd (=2) 78 x desetangsii x tetrapterum (5x x 3) 108 x dubioides Druce (=5xx) 108 dubium Leers (=lc) 65, 68. 71, 73, 74, 75 var. genuinum Syme (=lc) 73 var. maculatum sensu Bab. (=lc) 73 var. perforation (Tourlet) Pugsley (=lc) 74 dubium sensu Bab. (=lc) 73 dubium sensu Mauri (=lb) 71 elegans Stephan ex Willd. (7) 62, 63, 65, 109*, 111 var. elegans Jordanov & Koz. (=7) 112 var. genuinum Gus.ul. (=7) 1 12 var. pectinatum Celak. (=7) 1 1 1 var. pedunculatum Jordanov & Koz. (=7) 1 12 var. stepposum SIvul. & Rayss (=7) 1 12 elegans x perforatum (7x) 112 erectum Thunb. 62, 111 erosum (Schinz) O. Schwarz (=5xba) 104 erosum sensu O. Schwarz p. p. excl. typum (=lc) 74, 75 fallax Grimm (=lb) 71 subsp. fallax 71 var. fallax 71 forma punctatum O. Schwarz (=lb) 71 var. quadrangulare (L.) O. Schwarz (=3) 81 var. quadrangulare sensu O. Schwarz p. p. (=lb) 71 subsp. immaculatum (Murb.) O. Schwarz (=la) 70 foliosissimum Koidz. (=5d) 101 formosum Kunth 119 subsp. formosum sensu C.L. Hitchc. (=12) 118, 119 subsp. scouleri (Hook.) C.L.Hitchc. (=12) 118, 119 var. formosum sensu Dorn (=12) 118 var. nortoniae (M.E. Jones) C.L. Hitchc. (=12) 118 var. scouleri (Hook.) Coult. (=12) 1 17 formosum sensu A. Gray (=12) 117 hyssopifolium Vill. var. y sensu Choisy (=7) 1 1 1 var. pauciglandulosum Choisy (=7) 1 1 1 immaculatum (Murb.) Vierh. (=la) 70 insulare Foucaud & Mandon (=3b) 84 intermedium Bellynck (=lxb) 76 iwatelittorale H. Koidz. (9) 65, 114, 115 kamtschaticum Ledeb. var. pibairense Miyabe & Y. Kimura 65, 66, 68 var. senanense sensu Y.N. Lee (=6?)1 10 kohlianum Spreng. (=7) 1 1 1 komarovii Gorschk. (=5b) 89, 95 x laschii A. Frohl. (Ix) 75, 76, 102 nothoforma froelichii N. Robson (Ixb) 76 nothoforma laschii ( 1 xa) 76 leersii C.C. Gmel. (=lc) 73 lineolatum Jord. (=5a) 89 linnaeanum Callay & Gren. (=5xa) 104 maculatum Crantz (1) 62, 76, 88, 106, 1 1 1 subsp. desetangsiiforme A. Frohl. (=5xbb) 107 var. aporosum A. Frohl. (=5xbb) 107 subsp. erosum sensu Schinz & Keller (=lc) 74 var. imperforatum (Bonnet) A. Frohl. (=lc) 74 forma latisepalum A. Frohl. (=lc) 74 [sub]forma lucidum A. Frohl. (=lc) 74 [sub]forma nigrum A. Frohl. (=lc) 74 var. perforatum (Tourlet) A. Frohl. 74 subsp. erosum x acutum (=lxb) 76 subsp. erosum x perforatum (=5xa) 104 subsp. eu-maculatum Schinz & Thell. (=lb) 71 subsp. immaculatum (Murb.) A. Frohl. (la) 62, 63, 65, 66, 68, 69*, 71, 75, 108 var. epunctatum A. Frohl. (=la) 70 var. punctatum A. Frohl. (=la) 70 subsp. immaculatum x attenuatum ( 1 a x 6) 63 subsp. immaculatum x perforatum (=5xc) 108 subsp. maculatum (Ib) 62-65, 68, 69*, 71, 75, 76, 107, 1 1 1 forma punctatum (Schinz) Stjep.-Vesel. (=lb) 71 subsp. maculatum x perforatum (Ib x 5) 64, 102 subsp. maculatum x tetrapterum (Ib x 3) 64, 75 subsp. obtusiusculum (Tourlet) Hayek (Ic) 63-65, 68, 69*, 73, 75, 104, 106 var. imperforatum (Bonnet) A. Frohl. (=lc) 74 var. perforatum (Tourlet) A. Frohl. (=lc) 74 subsp. obtusiusculum x perforatum (=5xa) 64, 104 subsp. obtusiusculum x tetrapterum (Ic x 3) 76 subsp. quadrangulum (L.) Hayek (=3) 81 subsp. quadrangulum sensu Hayek (=lb) 71 subsp. styriacum A. Frohl. (=lc) 74 subsp. styriacum x acutum (=lc x 3) 76 subsp. typicum A. Frohl. (=lb) 71 var. genuinum (Schinz) A. Frohl. (=lb) 71 var. punctatum (Schinz) A. Frohl. (=lb) 71 forma luteum A. Frohl. (=lb) 71 subsp. typicum ' acutum (=lb x 3) 76 subsp. typicum x perforatum (=lb x 5) 105 subsp. undulatum (Schousb. ex Willd.) P. Fourn. (=2) 78 var babingtonii H. & J. Groves (=lc) 74 var. immaculatum (Murb.) Gu§ul. & Nyar. (=la) 70 maculatum x acutum (=1 x 3) 75 maculatum x perforatum (1x5) 64, 76, 102, 108 maculatum x perforatum forma sub-maculatum (= 5xba) 106 maculatum x perforatum forma sub-perforatum (=5xbb) 107 (maculatum x perforatum) x acutum (=5xxx) 108 maculatum x tetrapterum (Ix) 64, 75 marylandicum Biroli ex Colla (=5a) 89 x medium Peterm. (5xx) 64, 108 microphyllum Jord. (=5c) 96 x mixtum Du Moulin (=5xa?) 76, 104 momoseanum Makino (10) 65, 111, 115, 117 var. atumense Y. Kimura (=10) 115 var. momoseanum (=10) 115 mororanense R. Keller (=11) 117 forma tetragynum H. Leveille (=11) 117 nachitschevanicum Grossh. (=5c) 97, 98 neapolitanum Ten. (=3) 78, 81, 82 var. confertum (Choisy) Guss. (=3) 81 noeanum Boiss. (=5c) 97 nortoniae M.E. Jones (=12) 118 oaxacanum R. Keller 119 obtusum Moench (=lc) 73 officinale Gaterau (=5a) 89 officinarum Crantz (= 5a) 89 oliganthemum R. Keller (= 1 1 ) 117 ovalifolium Koidz. 62 perforato-acutum O. Kuntze & Focke (=5xx) 108 perforato-quadrangulum Lasch (=5xa) 104 perforatum L. (5) 62-66, 71, 75, 87, 88, 89, 90*, 98 101, 104-107, 1 10, 1 1 1 subsp. angustifolium (DC.) A. Frohl. (=5c) 96, 97 subsp. chinense N. Robson (5d) 66, 89, 90, 101 subsp. latifolium (Gaudin) A. Frohl. (=5xbb) 107 forma dentatum A. Frohl. (=5xbb) 107 forma fimbriatum A. Frohl. (=5xbb) 107 var. obscurum O. Schwarz (=5xba) 106 subsp. perforatum (5a) 65, 88, 89, 90*, 92, 94, 95, 96, 98 var. pellucidum O. Schwarz (=5a) 91 subsp. songaricum (Ledeb. ex Rchb.) N. Robson (5b) 65, 66, 89, 90*, 95, 96 subsp. veronense (Schrank) H. Lindb. (5c) 66, 88, 89, 90*, 94, 96, 101 subsp. vulgare (Schimp. & Spenn.) A. Frohl. (=5a) 91 forma brevisepalum A. Frohl. (=5a) 91 forma lineolatum (Jord.) A. Frohl. (=5a) 91 forma lucidum A. Frohl. (=5a) 91 var. P sensu Choisy (=5a) 89 var. y sensu Choisy (=5c) 96 var. albiflorum Choisy (=5a?) 89 var. alpinum Parl. (=5a) 89 var. angustifolium Borkh. (=5c?) 96 var. angustifolium DC. (=5c) 88, 89, 96 subvar. lineolatum Rouy (=5c) 97 var. angustifolium sensu Ohwi (=5d) 101 var. anomalum Frid. (=5a) 89 var. ceretanicum Sennen (=5a) 91 var. collinum Woron. (=5c) 89, 97, 101 var. confertiflorum Debeaux (=6) 101, 110 var. corioides Vokut (=5c) 97 var. decompositum Nyar. (=5a) 9 1 var. elatum Choisy (=5c) 96 var. ellipticum Durand & Pittier (=5a) 89 var. ellipticum Freyn (=5c) 97 STUDIES IN HYPERICUM 123 van floribundurn Sennen (=5c) 97 var. gracile Gruner (=5b) 89, 95 var. humile Stranski (=5a) 89, 9 1 , 93 var. latifolium Gaudin (=5xbb) 88, 106 subvar. lineolatum Rouy (=5xbb) 107 var. latifolium sensu W. Koch (=5a) 89 var. latiglandulosum Choisy (=5b) 95 var. lineolatum (Jord.) Hayek (=5a) 91 var. longicapsulum Jordanov & Koz. (=5c) 97 var. mediterraneanum Rouy (=5c) 97 var. microphyllum DC. (=5c) 96 var. microphyllum H. Leveille (=5d) 101 var. moesicum Velen. (=5c) 97 var. nanum Gaudin (=5c) 96 var. normale Woron. (=5a) 91 var. perforatum, N. Robson (=5a) 91 var. petiolatum Peterm. (=5a) 89 var. platycalyx Celak. (=5xbb) 107 var. semihumifusum Nyar. (=5a) 91 var. songaricum (Ledeb. ex Rchb.) K. Koch (=5b) 89, 95 var. stenophyllum (Opiz) Wimm. & Grab. (=5c) 96 var. typicum Beck (=5a) 89 forma microphyllum (DC.) Fiori (=5c) 97 var. veronense (Schrank) Ces. (=5c) 96 var. vulgare Schimp. & Spenn. (=5a) 89 forma anomalum (Frid.) Hegi (=5a) 91 forma brevisepalum (A. Frohl.) Hegi (=5a) 91 forma lineolatum (Jord.) Hegi (=5a) 91 forma lucidum (A. Frohl.) Hegi (=5a) 91 subvar. lineolatum (Jord.) Rouy (=5a) 89 perforatum x elegans (5x7) 64 perforatum x maculatum subsp. maculatum (5xlb) 105 perforatum x maculatum subsp. obtusiusculum (5 xlc) 104 (perforatum x maculatum) x tetrapterum (5x x 3) 108 perforatum x tetrapterum (5 x3) 108 perforatum group 1 1 9 porphyrandrum H. Leveille & Vaniot (=11) 117 procumbens R. Keller (=11) 115 przewalskii Maxim. 61 pseudo-nikkoense H. Koidz. (=11?) 117 pseudopetiolatum R. Keller 115 pulchrum Pallas ex M. Bieb. (=7) 1 1 1 quadrangulare sensu L. et auct. (=3) 71, 80 var. alatum (Retz.) Boenn. (=3) 81 var. humile Boenn. (=3) 81 \ar.patulum Boenn. (=3) 81 quadrangulum L. (=3) 67, 71, 80, 81 subsp. acutum (Moench) Goday & Carbonell (=3) 81 subsp. corsicum (Steud.) Fiori & Fiori (=3b) 84 subsp. desetangsii (Lamotte) Tourlet (=5xa) 104 subsp. erosum (Schinz) Schinz (=5xba) 106 var. epunctatum Schinz (=lc) 74 subsp. immaculatum (Murb.) Wettst. (=la) 70 subsp. maculatum (Crantz) Goday & Carbonell (=lb) 71 subsp. obtusiusculum Tourlet (=lc) 73 var. imperforatum (Bonnet) Tourlet (=lc) 73 var. perforatum Tourlet (=lc) 73 subsp. quadrangulum sensu Tourlet (=lb) 71 subsp. undulatum (Schousb. ex Willd.) Goday & Carbonell (=2) 78 var. |3 sensu Choisy (=lc) 73 var. y sensu Choisy (=lb) 71 var. 8 sensu Choisy (=3) 80 var. acutum (Moench) Fiori (=3) 81 var. ambiguum Schur (=lc?) 73 var. confertum Choisy (=3) 80, 81 var. dubium (Leers) Choisy (=lc) 73 var. erosum Schinz (=5xba) 106 var. genuinum Schinz (=lb) 71 var. immaculatum Murb. (=la) 70 var. macmphyllum Schur (=Ib) 71 var. maculatum (Crantz) Choisy (=lb) 71 var. occidentale Franch. (=lc) 73 var. punctatum Schinz (=lb) 71 var. subcorymbosum Schur (= 1 b) 71 var. tetrapterum (Fr.) Borg (=3) 81 forma undulatum (Schousb. ex Willd.) Borg (=2) 78 var. undulatum (Schousb. ex Willd.) Choisy (=2) 76 quadrangulum sensu Colm. (=2) 78 quadrangulum sensu Des Etangs (=lxb) 76 quadrangulum sensu Fr. et auctt.(= 1 b) 7 1 quadratum Stokes (=3) 80 quadrialatum Wahlenb. (=3) 81 repens Duby ex Nyman (=3b) 84 rotundatum Schur (=3) 81 schlosseri Heuffl. (=5c) 96 scouleri Hook. (12) 62, 65, 66, 111, 116*,117 subsp. nortoniae (M.E. Jones) J.M. Gillett (=12) 118 subsp. scouleri (=12) 118 simulans Rose (=12) 117 skouleri Walp. (=12) 117 songaricum Ledeb. ex Rchb. (=5b) 95 x sparsiflorum Schur (=5xx) 108 stenophyllum Opiz (=5c) 96 tenellum Tausch (=3b) 84 tetragonum Fr. (=lb) 71 tetraptero-quadrangulum Lasch (=lx) 75 tetrapterum Fr. (3) 61, 64, 65, 67, 71, 76, 77*, 78, 80, 81, 98 subsp. corsicum (Steud.) Zangh. (=3b) 84 subsp. tetrapterum, Zangh. (=3a) 82 subsp. undulatum (Schousb. ex Willd.) P. Silva (=2a) 78 var. anagallidifolium Boiss. (3c) 84 var. boeticum (Boiss.) Cout. (=2b) 78, 79, 80 var. corsicum (Steud.) Boiss. (3b) 84 var. intermedium Coss. & Germ. (=lxb?) 76 var. neapolitanum (Ten.) N. Terrace. (=3) 81 var. rotundifolium Willk. (=2b) 79, 80 var. tenellum (Tausch) Parl. (=3b) 84 var. tetrapterum (3a) 77*. 80, 82 var. undulatum (Schousb. ex Willd.) Cout. (=2a) 78 forma stigmaphyllum Woron. (=3) 81 tetrapterum group 71 tosaense Makino (8) 62, 65, 1 1 1, 112, 1 14, 1 15 var. atumense (Y. Kimura) Y. Kimura (=10) 1 1 1, 1 15 forma insulare Y. Kimura (=8) 1 14 forma tosaense Y. Kimura (=8) 112 triquetrifolium Turra (4) 62, 63, 65, 67, 77*, 84, 87 umbellatum Miel. ex Wohlf. (=lb) 71 undulatum Schousb. ex Willd. (2) 61, 64, 65, 76, 78, 87 var. boeticum (Boiss.) Lange (2b) 65, 79, 80 var. undulatum (2a) 65, 77*, 78, 80, 81 undulatum x tetrapterum (2 x 3) 80 veronense Schrank (=5c) 96 vulgare Bubani (=5a) 91 vulgare Lam. (=5a) 89 yamamotoi group 62 yamamotoi var. riparium Y. Kimura 62 yezoense Maxim. (11) 65, 115, 116*, 117 var. momoseanum (Makino) Ohwi (=10) 115 yezoense group 62 yoitiense H. Koidz. (=11?) 117 Bull. nat. Hist. Mm. Lond. (Bot.) 32(2): 125-136 Issued 28 November 2002 The natural history of reproduction in Solanum and Lycianthes (Solanaceae) in a subtropical moist forest STAGEY D. SMITH Department of Botany, 132 Birge Hall, 430 Lincoln Drive, University of Wisconsin, Madison WI 53706-1381, U.S.A. SANDRA KNAPP Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS Introduction 125 Methods 127 Study site 127 Taxa studied 127 Floral phenology 127 Flower and fruit production 129 Flower visitors 129 Pollen load analysis 129 Results 129 Floral phenology 129 Flower production 131 Fruit production 131 Flower visitors 131 Pollen load analysis 133 Discussion 133 Floral phenology 133 Flower production 133 Fruit production 135 Flower visitors 135 References ... 136 SYNOPSIS. The phenology and pollination of seven understory species of buzz-pollinated Solanaceae (Solanum erythrotrichum, S. lanceifolium, S. rudepannum, S. cordovense, S. nudum, Lycianthes hypoleuca and L gorgonea) were investigated at the end of the dry season in the subtropical moist forest at the Las Cuevas Research Station, Chiquibul Forest Reserve, Cayo District, western Belize. Three phenological phenomena were tracked: the opening and closing of flowers, flower production and fruit production. The large short-lived white flowers of S. lanceifolium, S. rudepannum, Lycianthes hypoleuca and L. gorgonea opened around sunrise and closed at sunset. The purple flowers of 5. erythrotrichum and the small white flowers of S. nudum and S. cordovense opened more or less randomly. All seven study species flowered at least once during the months of May, June and July; there was substantial overlap in the flowering of some species. Four species, S. rudepannum, S. cordovense, S. lanceifolium and S. erythrotrichum, developed mature fruit during the monitoring period while the remaining species possessed immature fruit at the termination of the study. Thus, it appeared that these seven solanaceous species would provide a fairly constant supply of mature fruit during the rainy season. During observations of pollinators, 17 different bees in the families Colletidae. Halictidae and Apidae were found to visit the buzz-pollinated flowers of Solanum and Lycianthes. Analysis of the pollen loads revealed that bees were highly constant to Solanaceae although it was not possible to determine their constancy to particular species. Very few visits were observed to 5. cordovense and L. gorgonea. INTRODUCTION The Solanaceae is an economically important, cosmopolitan family with over 2500 species in some 90 genera. The family has members occurring in all habitats, and their habit ranges from canopy trees to minute ephemeral herbs. Flowers in the family Solanaceae exhibit a wide array of forms (Knapp, 2002), and are pollinated by a similarly © The Natural History Museum, 2002 wide variety of organisms, including bees, hummingbirds and bats (Cocucci, 1999). In Mesoamerica, the family is an important com- ponent of the forest understory in a variety of habitats, and in Belize, it is the eleventh most diverse vascular plant family overall (Balick et al., 2000). Solanum L. and Lycianthes (Dunal) Hassl. are the two largest genera in the family (D'Arcy, 1991). Solanum comprises 1500 or more species, 800-900 of which occur in the New World (D'Arcy, 126 S.D. SMITH & S. KNAPP Fig. 1 Study species of Solanaceae at LCRS. Photographs taken from vouchers cited in the text. A. Solanum erythmtrichum, B. 5. lanceifolium, C. S. rudepannum, D. S. cordovense, E. S. nudum, F. Lycianthes hypoleuca, G. L. gorgonea. Scale bar = 1 cm. REPRODUCTION IN SOLANUM & LYCIANTHES 127 1991; Nee, 1999). Lycianthes contains approximately 200 species distributed in both tropical America and Asia (D'Arcy, 1973, 1979; Symon, 1985), and despite being superficially similar to Solatium (see below), it is closely related to the genus Capsicum L. (Olmstead et al., 1999). Approximately ten species of Lycianthes and 30 species of Solarium occur in Belize (Balick et al., 2000; Knapp et al., in press). Although phylogenetically distant, members of Solarium and Lycianthes share similar, convergent, floral morphology. The flow- ers possess a five-parted gamopetalous corolla. The five equal stamens are fused to the corolla tube, and the bright yellow tubular anthers form a cone around the style (typical 'solanoid' flowers, Endress, 1994; Fig. 1). The flowers of Solanum and Lycianthes are buzz-pollinated (Buchmann, 1983; Nevers, 1986; Lester et al., 1999). When buzzing a flower, an insect grasps the cone of tubular anthers with the front pairs of legs, wraps the abdomen around the apical pores of the anthers, and vibrates the anthers by rapidly contracting the indirect flight muscles (Michener, 1962; Buchmann & Hurley, 1978). This audible action causes pollen to burst out of the anther and land onto the venter of the insect. As the flowers of these Solanum and Lycianthes species offer no nectar and little scent, pollen is the only reward available to their pollinators (Symon, 1979; D'Arcy et al., 1990; pers. obs.). Although the mechanism of buzz pollination in Solanaceae has been well characterized (Michener, 1962; Buchmann, 1983), detailed studies which identify both the buzz-pollinated plant species and their pollinators are few (but see Linsley & Cazier, 1963; Anderson & Symon, 1988). Here we examine the phenological patterns and pollination biol- ogy of seven sympatric buzz-pollinated species of Solanum and Lycianthes in the Chiquibul forest of western Belize. In studying their phenology, we tracked the daily opening and closing of flowers as well as the seasonal timing of flower and fruit production. Characterizing these daily and seasonal floral events in turn set the stage for investigations of pollination biology. In undertaking this study, we seek to expand our knowledge of Solanaceae in Belize, their natural history and range of pollinators. METHODS Study site The Chiquibul Forest Reserve, Cayo District, western Belize, is nested completely within the Chiquibul National Park (Fig. 2). The vegetation comprises deciduous semi-evergreen and deciduous sea- sonal forest with stands of Caribbean pine to the north (Wright et al., 1959). The Macal River roughly divides the Caribbean pine forest to the north from the broad-leaved tropical forest of the Chiquibul to the south. Las Cuevas belongs to the subtropical moist life zone (Holdridge et al., 1971). While largely protected, some selective logging of commercial species such as mahogany (Swietenia macrophylla) and cedar (Cedrela odorata) is carried out in the Chiquibul on a >40 year rotational basis. The area has also suffered hurricane damage in tl ; past (most recently from Hurricane Hattie in 1961), and the forest is a mosaic of different successional stages. Our field studies were carried out at the Las Cuevas Research Station (LCRS) (16°44' N, 88°59' E; altitude 550-600 m; Fig. 2) operated jointly by the Forest Department of the Government of Belize and The Natural History Museum, London. Las Cuevas is situated in the centre of the Chiquibul Forest Reserve on the north- western side of the Maya Mountains, and has an annual rainfall of c. 1500 mm. The dry season at Las Cuevas runs from December to May, while the rainy season begins in mid to late June and ends in January. This study began on 27 May 2000 and continued until 23 July 2000, thus covering the transition from the dry season to the wet season. The flowering of understory herbs including Solanum is known to peak during this transition period in Costa Rica and in Panama (Croat, 1969, 1975; Opler et al., 1980; Knapp, 1986), and in cerrado habitats in Brazil (Oliviera & Gibbs, 2000). Therefore, June and July were expected to be ideal months in which to study the reproductive biology of the understory Solanaceae in Belize. Though the study was conducted primarily along a trail to Monkey Tail River, pollinator behaviour was also studied at an observation tower located 1 km northwest of the station. Taxa studied The habit, flowers and fruit of the seven species studied at Las Cuevas are briefly described below. The Solanum species are listed in their respective infrageneric taxon (sensu Nee, 1999). The infrageneric classification of Lycianthes is not well understood, so this information was not included in the descriptions below. • Solanum erythrotrichum Fernald (Subgenus Leptostemonum) - An erect spiny shrub, with purple flowers (2 cm in diameter), tapering anthers and globose, slightly hairy green hard fruit, around 1 .5 cm in diameter. It is commonly found in partly shaded disturbed areas and tree falls. (Voucher: S.D. Smith 008, BRH, BM). Fig. 1A • 5. lanceifolium Jacq. (Subgenus Leptostemonum) - A herba- ceous weedy vine with short recurved spines. Its flowers are white, up to 2 cm in diameter with tapering anthers, and the small round fruit are 1 cm in diameter and bright red at maturity. It is found in thickets and in the canopies of small to medium trees. (Voucher: S.D. Smith 034, BRH, BM). Fig. IB • S. rudepannum Dunal (Subgenus Leptostemonum) - An erect spiny shrub, which produces large white flowers up to 2.75 cm in diameter and hard round berries which are about 2 cm in diameter and green at maturity. It prefers clearings such as road verges and fields. (Voucher: S.D. Smith 013, 037, BRH, BM). Fig. 1C • S. cordovense Sesse & Mo9- (Subgenus Solanum) - A clamber- ing woody shrub with small white flowers (1 cm in diameter). The fruit is a globose juicy black berry, 1-1 .25 cm in diameter. It is common along partly shady trails or in thickets. (Voucher: S.D. Smith 002, BRH, BM). Fig. ID • S. nudum Dunal (Subgenus Solanum) - A ubiquitous, entirely glabrous shrub with small white flowers similar to those of S. cordovense. Its fruit are yellowish-green berries of 1 cm in diameter. It is abundant in sunny areas along trails and roads. (Voucher: S.D. Smith 038, BRH, BM). Fig. IE • Lycianthes hypoleuca Standl. - A vigorous climbing shrub with large rotate white flowers (1.8-2 cm in diameter) and round red fruits just under 1 cm in diameter. It is found occasionally in small forest gaps or on hillsides, either in part-shade or full sun. (Voucher: S.D. Smith 026, BRH, BM). Fig. IF • L. gorgonea Bitter - A delicate scandent shrub with flowers similar to those of L. hypoleuca. Its fruit are slightly ovate, red berries of about 1 cm in diameter. It is quite infrequent, occurring only in the shady understory. (Voucher: S.D. Smith 012, BRH, BM). Fig. 1G Floral phenology Information was systematically collected on the time of opening, the time of closing and the extent to which the corollas were open (e.g., partly open to fully open with petals reflexed). These data were used to characterize the movements of the corolla and the longevity of the 128 S.D. SMITH & S. KNAPP JS ^ MAYA MOUNTAINS 0 10 20km Fig. 2 Map of southern Belize (Belize shown in inset) showing position of Chiquibul Forest Reserve and the Las Cuevas Research Station (filled circle). REPRODUCTION IN SOLANUM & LYCIANTHES 129 flowers. One mature individual was selected from each of the study species, and up to 25 flowers per individual were tagged and monitored for their entire lifespan. At the beginning of the study, the flowers were checked every four hours. Once it was determined whether the flowers of a given species close at night or not, flowers were checked only from 0500 hours to 2 1 00 hours for the remainder of the study. The position at 0100 hours was interpolated from observations at 2100 hours and 0500 hours. For example, if the corolla was 50% open at 2100 hours and was 100% open at 0500 hours, its position at 0100 hours was recorded as 75% open. At Las Cuevas, the sun rises around 0500 hours and sets at 1845 hours. The position of the petals was recorded as follows: 0 - unopened bud (beginning) or flower dead (end); 0. 1 =closed but petals not appressed as in the bud; 0.25=flower 25% open; 0.50=flower half-open; 0.75=flower 75% open; 1.0=flower open with petals perpendicular to the pedicel; 1.25=flowers with petals reflexed. After the flower bloomed, 0.1 indicated that the flower had closed but remained attached whereas 0 indicated that the flower had fallen off. A flower was considered open to visitors when it was 40% open; below that level the aperture was too small for the bees to access the reproduc- tive organs (pers. obs.). Flower and fruit production All mature individuals (up to a maximum of 25) of the seven study species present in the first two kilometers of the trail to Monkey Tail River were tagged, and (starting June 1 ) every three days, the number of inflorescences, flowers and fruit was recorded. An inflorescence was counted from the moment it was visible at the tip of the branch to the abscission of its last flower. Lycianthes gorgonea and L. hypoleuca were not included in the counts of inflorescences because the flowers occurred singly. A flower was counted if its petals were open sufficiently so that the cone of anthers could be easily seen. Fruit were only counted when the ovary had doubled in size, i.e., once they were so large that it was clear that they would not be aborted. Approximate canopy cover was also recorded for each individual in the study in order to make a preliminary assessment of its effect on these species. The percent canopy cover above each individual was estimated visually and recorded in one of the following five categories: 0% cover, 25% cover, 50% cover, 75% cover and 100% cover. Flower visitors Observations were taken from 0400 hours to 1900 hours and spo- radically throughout the night. Flower visitor activity was most intense between sunrise and noon, so most observations took place between 0500 hours and 1200 hours. Observations included time of visit, number of flowers visited, length of visit and activities on the flowers. Each study species was observed for 1 8 to 20 hours or more. Every different visitor was collected using a sweep net and trans- ferred directly into an eppendorf tube containing 1 ml isopropanol. Bees too large to fit into a tube were immediately washed in isopropanol to remove pollen, and this pollen sample was kept for pollen analysis (see below). When possible, several specimens of each visiting species were captured, so that later pollen load analysis could give some indication of the overall constancy of the species. Chris O'Toole (Oxford University) identified each bee to genus level and to species when possible. Pollen load analysis To remove pollen, each bee was transferred from its eppendorf tube to a 1 2 ml glass tube. The eppendorf tube was rinsed out with an additional one ml of isopropanol, which was added to the glass tube. Each bee was then shaken vigorously for 30 seconds to free the pollen from the body. The body was visually checked afterwards to assure that the pollen had been removed, and washed further if necessary. The tubes containing the pollen suspended in isopropanol were centrifuged at 5000 x g for five minutes to pellet the pollen. The isopropanol was poured off and the tubes inverted on a paper towel to drain. Then, 0.3 ml melted glycerol jelly was added to the pellet and stirred. One drop of this mixture was poured onto a glass slide and topped with a slide cover. Three hundred pollen grains were counted on each slide, and the percentage of solanaceous pollen calculated from this count. Solanaceous pollen was distin- guishable from other pollen grains because it is small, smooth and tricolporate. It was not possible to distinguish between the study species with the light microscope. Density of pollen grains was estimated at 40x magnification and was coded as follows: l=one pollen grain per view or less; 2=1 to 5 pollen grains per view; 3=5 to 10 pollen grains per view; 4=10 to 20 pollen grains per view; 5=greater than 20 pollen grains per view. RESULTS Floral phenology Based on the data collected, the seven species could be divided into two basic groups: those whose flowers opened at sunrise and those whose flowers opened throughout the day and night. The flowers of Solanum lanceifolium, S. rudepannum, L. hypoleuca and L gorgonea opened around 0500 hours, sunrise at Las Cuevas, and closed between 1700 and 2100 hours (Fig. 3B, C, F, G). The flowers of 5. cordovense and S. nudum opened more or less randomly throughout the day and night and remained open through the night, creating the potential for nocturnal pollination (Fig. 3D, E). The corolla of some S. cordovense flowers closed partially during the night although never sufficiently to exclude pollinators entirely. Solanum erythrotrichum (Fig. 3A) was an intermediate between these two basic groups as its flowers only opened between 0600 and 1300 hours (instead of right around sunrise) and closed at night. Great variation in the synchrony of flower movements was observed among the seven species. Fig. 3 shows the average position of a corolla of a given species throughout the day. For each data point, standard deviation was calculated to show the variation in corolla position between flowers of the same species. Several species show extreme asynchrony in opening and closing, such as Solanum cordovense and S. nudum, both of which had long-lived (Fig. 3D, E), small, white flowers less than 1 cm in diameter. This is consistent with the observation that their flowers open randomly throughout the day and night. The large showy white flowers of 5. rudepannum, Lycianthes hypoleuca and L. gorgonea, ranging from 1 .75 to 2 cm in diameter, opened much more uniformly, particularly on their first day (Fig. 3C, F, G). Flowers of the latter normally opened for one day only (Fig. 3C, F, G), with occasional flowers opening on a second day (perhaps due to lack of pollination). The flowers of 5. lanceifolium usually opened to some extent for a second day although they were rarely visited by the bees, which could apparently discern the older flowers (Fig. 3B) (see below). The petals of 5. nudum remained reflexed for only the first 4 to 12 hours, then the flowers slowly closed over the next day. The petals of 5. cordovense flowers were reflexed for fewer consecutive hours although they were capable of returning to the reflexed position for several days in a row. This suggests that they continued to be receptive to pollinators for several days after opening or that the reflexed petals act as a pollinator attractant. 130 S.D. SMITH & S. KNAPP cd 1 0 Day 1 Day 2 Day 3 Day 4 Day5 S. erythrotrichum (N=6) Fig. 3 Corolla movements. Data from up to 25 flowers was combined to produce the graphs. The position of the petals on each flower monitored was recorded every 4 hours using the following scale: 0 = unopened bud (beginning) or flower dead (end); 0.1 = closed but petals not appressed as in the bud; 0.25 = corolla 25% open; 0.50 = corolla 50% open; 0.75 = corolla 75% open; 1.0 = open with petals perpendicular to the pedicel; 1 .25 = flowers with petals reflexed. For heterostylous species (Solarium lanceifolium, S. cordovense and S. nudum), only the data from long-styled flowers are shown. Error bars around each point are ± one standard deviation. Where no bars appear, the standard deviation is zero. REPRODUCTION IN SOLANUM & LYCIANTHES 131 Solarium erythrotrichum was an exception to these generaliza- tions. It had flowers of 2 to 2.5 cm in diameter that opened well after sunrise, but they were long-lived, persisting for up to 80 hours (over three days) (Fig. 3 A). Each of its flowers opened briefly to the fully reflexed position during late afternoon of its first day before closing for the night. Its flowers then returned to the reflexed position on the second day and occasionally on the third day as well. Old flowers could be distinguished from younger flowers by the purple petals, which darken with age. Flower production All the species flowered at least once during the two-month period of June and July. Two of the species flowered continuously through- out the period: Solarium nudum and S. erythrotrichum (Fig. 4A). Solanum cordovense, S. lanceifolium and S. rudepannum all flow- ered in late May, paused in mid-June and began to flower again in late June or July (Fig. 4A). The two species of Lycianthes were very different from each other in their flowering phenology. Lycianthes hypoleuca flowered only during early June (and presumably in late May before monitoring had begun) (Fig. 4A). Lycianthes gorgonea flowered for most of June and sporadically in July and still had buds present in late July. With relation to intensity of flower production, the most prolific bloomers of the seven species were Lycianthes hypoleuca and Solanum nudum. On average, mature individuals of these species produced around 200 or more flowers (Table 1 ). Despite the fact that L. hypoleuca possesses much larger flowers, the display produced by S. nudum overwhelmed that of L. hypoleuca because of the sheer number of individuals flowering. While only six of the L. hypoleuca individuals monitored flowered, all 25 5. nudum individuals flow- ered, producing a total of 4936 flowers over the 52-day monitoring period. Solanum lanceifolium produced the fewest flowers during the monitoring period (Table 1 ). Considering the number of buds present on individuals at the end of the study, S. lanceifolium probably produced quite a large display in August. Regression analyses revealed significant relationships between canopy cover and flower production for several species. Individuals of both Solanum cordovense and S. lanceifolium in the shade produced significantly fewer flowers (p=0.04, p=0.02 respectively). As colonizers of disturbed areas and secondary forest, these species probably have a low tolerance for shady conditions. Increased canopy cover was also related to a greater proportion of male (short- styled) flowers in the andromonoecious S. nudum (Smith & Knapp, in prep.). Fruit production The first species to produce fruit was Solanum rudepannum. Its round, green berries were becoming mature in late May and were almost entirely gone by mid-June (Fig. 4B). The fruit of S. cordovense were the next to mature, turning from green to a deep purple-black in late June (Fig. 4B). The soft, sweet berries were consumed, probably by birds, in June and July. The fruit of S. lanceifolium matured in July, shortly after those of S. cordovense (Fig. 4B). Its scarlet berries were not nearly so sweet but quite piquant (pers. obs.). When the berries of S. lanceifolium were fully ripe, the fruiting vines attracted large numbers of birds (N. Bol, pers. comm.). Solanum erythrotrichum, unlike the previous three species, had mature fruit available for the majority of the monitoring period (Fig. 4B). Starting in mid-June, its ripe green fruit began to be taken, although new fruit quickly replaced them. The fruit of the remaining three species were not yet ripe when the monitoring period con- cluded although it appeared that the fruit of 5. nudum would mature first in August, followed by Lycianthes gorgonea then L. hypoleuca (Fig. 4B). The amount of fruit produced during June and July varied greatly among the seven species (Table 1). Solanum nudum produced the largest number of fruit per individual on average, closely followed by 5. rudepannum. Lycianthes gorgonea produced the fewest, around 1 3 per individual on average, which was anticipated considering its small floral display (Table 1). Flower visitors Bees of the families Colletidae, Halictidae and Apidae (Michener, 2000) were the only insects observed to visit, vibrate (buzz) and presumably pollinate the flowers of Solanum and Lycianthes (listed in Table 2). Pollination is normally effected only by floral visitors that vibrate or buzz the flowers in buzz-pollinated flowers, so we have generally assumed that the visitors we observed were legiti- mate pollinators of these species. Flowers bagged as part of another study (Smith & Knapp, in prep.) never set fruit. Members of the three bee families we observed at LCRS comprise most of the pollinators of melittophilous Solanaceae (Linsley & Cazier, 1963; Sazima et al., 1993; Raw, 2000). The frequency of visits by the bee species to the study taxa is shown in Table 3. The bees were most active in the morning, and this activity tapered off quickly as the day progressed, a foraging pattern commonly observed in tropical bees (Knapp, 1986; Roubik, 1989). The earliest visitor by far was Megalopta sp., which began foraging on L. hypoleuca between 04 1 5 and 0430 hours. It was also noted that larger bees tended to be most active early in the morning while smaller bees were active through- out the morning and sometimes into the afternoon (Table 2). Most bees extracted pollen by buzzing the flower although some small bees obtained pollen by digging into the pores and scavenging on floral parts, as has been observed elsewhere (Anderson & Symon, 1988; Storti, 1988). Bees occasionally dug into the pores after first attempting to buzz the flower, a behaviour that appeared to widen the aperture, allowing the pollen to escape. Neither bees nor other insects were seen to 'rob' pollen by cutting holes into the anthers. In general, large bees visited for shorter periods of time than smaller bees (Table 2), an observation also made by Anderson & Symon (1988). In general, small bees visited small flowers (e.g., those of Solanum nudum) while larger bees visited the large flowers (e.g., those of Lycianthes hypoleuca). All bees appeared to locate flowers visually, and were often seen approaching from afar. Once on the flower, bees positioned them- selves so that their venter covered the pores of the anthers, and buzzed the anthers to extract the pollen. While buzzing, medium to small bees rotated up to 5 or 6 times to obtain the maximum amount of pollen from each anther. Large bees rotated only once or twice and sometimes not at all. The weight of the large bees was sufficient to invert the flowers, further assisting pollen extraction (Linsley & Cazier, 1963). Bees rarely visited the same flower twice (also noted by Shelly & Villalobos, 2000) and recognition of 'buzzed' flowers may have been assisted by physical changes to the flower. For example, damage by tarsal claws caused bruising on the abaxial anther surface and bees preferentially buzzed flowers free from marks. This preference was tested by bagging flowers on several plants, and allowing the other flowers to be visited and buzzed. When the bags were removed and bees had a choice of buzzed and unbuzzed flowers, only unbuzzed flowers were visited. The bees observed during this study varied greatly in constancy, as measured by visits to non-conspecifics on a single foraging trip. Paratetrapedia sp. was by far the most promiscuous and was observed visiting members of the Melastomataceae, Lamiaceae, 132 S.D. SMITH & S. KNAPP - - S. erythrotrichum S. lanceifolium • S. rudepannum S. cordovense S. nudum L. hypoleuca L. gorgonea 55 ,£ •$> ^ V Date Fig. 4 Flowering (A) and fruiting (B) phenology from 1 June 2000 to 2\ July 2000. Shown is the proportion of flower or fruit present on a given date. This proportion was calculated by summing the total number of flowers or fruit present on all the individuals of a species on a given date and dividing by the total number of flowers or fruit present on all the individuals of a species during the entire monitoring period. Green-fruited species have been given dashed lines; red- or black-fruited species have solid lines. REPRODUCTION IN SOLANUM & LYCIANTHES Table 1 Flowering and fruiting intensity from 1 June 2000 to 21 July 2000. 133 Percentage of mature individuals monitored which flowered Average number of flowers produced per flowering individual Percentage of individuals monitored which fruited Average number of fruits produced per fruiting individual 5. erythrotrichum S. lanceifolium S. rudepannum S. cordovense S. nudum L hypoleuca L gorgonea (N=25) (N=25) (N=10) (N=25) (N=25) (N=10) (N=5) 52% 32% 50% 52% 100% 60% 80% 15.85 4.88 51.60 16.54 197.44 220.00 18.50 40% 60%* 40% 44% 80% 60% 80% 13.62 22.27 63.50 18.55 74.20 33.83 12.75 A number of individuals of Solarium lanceifolium were fruiting when the survey began, remained in fruit and never flowered during the study period. Piperaceae, and Asteraceae in addition to the Solanaceae. Paratetra- pedia sp. also flew between study species on a single foraging trip, particularly those which grew side by side, such as Solarium rudepannum and 5. nudum, potentially resulting in interspecific pollen transfer. Some bees visited only solanaceous species, but were not constant to any individual species. For example, Colletes sp. visited S. rudepannum, S. nudum and 5. cordovense in one foraging trip, but was never observed visiting nearby non-solana- ceous species in flower. At the other end of the spectrum were bees like Xylocopa anthophorides and Xylocopa sp., which were only observed to visit a single study species (Table 2). Pollen load analysis Analysis of pollen loads corroborated field observations that sug- gested varying levels of constancy among bees. The percentage of pollen from Solanaceae varied from 69.95 to 1 00.00. More promis- cuous pollinators such as Paratetrapedia sp. carried lower amounts of solanaceous pollen whereas constant pollinators like Xylocopa carried nearly 100% (Table 2). Overall, visitors to the seven study species carried a very high percentage of solanaceous pollen. Unfor- tunately, it was not possible to distinguish between the pollen of the seven study species. The size of the pollen load found on the pollinator species generally depended on the size of the bee and how well developed its corbiculae were. Larger bees with well-developed corbiculae such as Xylocopa spp. and Eulaema sp. carried the most pollen (Table 2), followed by small- and medium-sized apid bees such as Exomalopsis sp. 2 and Melipona sp. with large corbiculae and hairy bodies (Table 2). The less hairy halictids, which lack corbiculae, carried small amounts of pollen, although Megalopta sp. was an exception in foraging only on newly opened, pollen-rich flowers of Lycianthes hypoleuca. DISCUSSION Floral phenology Patterns of flower movements (i.e., opening and closing of the corolla) often relate to pollination syndrome. Most flower visitation took place in the morning hours, when fresh flowers were presented. Some individuals of L. hypoleuca opened their flowers before dawn, allowing pollination by the night-foraging Megalopta sp. (Hopkins et al., 2000). Only a few bees foraged throughout the day and these tended to visit plant species that were offering fresh flowers through- out the day, such as S. nudum. Flower movements varied slightly depending on location; e.g., flowers of the individuals of Lycianthes hypoleuca near the Las Cuevas field station opened around 0500 and closed between 1900 and 2 1 00 while those near the observation tower were open between 0430 and 1500 to 1700. The light from the LCRS generator at the former site may account for this difference. Individuals in the sun occasionally had slightly different phenology from those in the shade; e.g., S. cordovense in the sun kept their flowers open for a greater number of consecutive days than those in the shade. Primack (1985) compared the longevity of flowers in major habitat classes and found that most tropical forest species have one- day flowers, regardless of their pollinators. Furthermore, he suggested that within these classes, some variation in longevity within related taxa may be due to flower size. Primack postulated that large flowers would be expected to last longer than small flowers because plants expend more energy to produce them. The present study indicated that the exact opposite is true. The largest-flowered species, with the exception of S. erythrotrichum, were the most short-lived. This suggests a trade-off between size and longevity where a plant can either maintain energetically-expensive large flowers for a short time or small energetically-inexpensive flowers for a longer time. Flower production Available surveys of flowering phenology in the seasonal tropics indicate that most species exhibit a definite peak or series of peaks in flowering and fruiting at some time during the year (Croat, 1969; Opler et al., 1980; Tanner, 1982; Newstrom et al., 1994). Flowering times for tropical understory herbs and shrubs, like Solanum and Lycianthes, are often clustered around the transition period from the dry to the wet season (Augspurger, 1983; Knapp, 1986; Oliviera & Gibbs, 2000). This pattern may relate to plants' resource needs such as amount of precipitation (Croat, 1969; Augspurger, 1980, 1983) or their use of environmental cues like temperature to time reproduc- tive activities (Levin, 1978). In the present study, it was only possible to track the study species for two months, an insufficient time to draw conclusions about the phenology over the course of a year or even a season. Locating large numbers of mature individuals in the study area was problematic for the two species of Lycianthes and for 5. rudepannum. Thus, the data presented here provide a preliminary picture of flower production for these seven species. Plant guilds, i.e., sympatric species sharing a pollination syn- drome, may exhibit staggering of flowering times (e.g., Heithaus et al., 1974; Newstrom et al., 1994). Although the buzz-pollinated Solanaceae at LCRS can be considered a guild (Sakai et al., 1999, Murray et al., 2000), there did not appear to be a sequence of flowering times within the study period. Knapp (1986) described a similar situation with eight species of Solanum section Geminata in Monteverde, Costa Rica, which had randomly distributed and largely overlapping flowering peaks over a year, with some species bloom- ing more than once. Our finding that the multiple buzz-pollinated 134 S.D. SMITH & S. KNAPP 8, pajisu sgpads "'7 "7 uinpnu •$• umuuvdaptu •$ - it L "" O "rt "O u. 3 U 1> T3 o •" 'ff u S £ =3 118 5 as ^11 1 \ \\NSS S S S S S S S \ S S S S N S S S S so OO vOO oinoooooo-^-o Ot^OOOOOiO>OO E 3 _ I II so^o pjo^S ^3 O SO U E REPRODUCTION IN SOLANUM & LYCIANTHES \ 35 Table 3 Frequency of pollinator visits on Solatium and Lycianthes species at Las Cuevas. For each study species, flower hours was calculated from a series of observations (1 to n) using the following equation: O = (F, x H,) + (F,x H2) + . . . (Fnx Hn), where O equals the number of flower-hours, H equals the number of hours a given plant was observed, and F equals the number of flowers present on the plant at the time of observation. Species Total number of hours observed Total number of flowers observed Number of flower-hours observed Pollinating visitors per flower-hour Common pollinators (number which visited during hours observed) 5. erythrotrichum S. lanceifolium S. rudepannum S. nudum S. cordovense L. hypoleuca L gorgonea 21.40 19.13 18.10 49.45 20.52 19.52 19.00 61 79 242 1077 82 1190 28 160.53 125.55 595.93 2175.77 116.25 2017.02 71.03 0. 1 3 1 Paratetrapedia sp. ( 1 7) 0.0 1 6 Xylocopa anthophorides ( 1 0) Paratetrapedia sp. (22) Eufriesia sp. ( 1 0) Euglossa sp. (4) 0.037 Paratetrapedia sp. ( 1 6) Colletes sp. (5) 0.0 1 8 Paratetrapedia sp. (27) Colletes sp. (5) Augochloropsis sp. (5) 0.017 Colletes sp. (2) 0. 1 1 6 Megalopta sp. ( 1 6) Xylocopa sp. (9) Xylocopa cf. omata (13) Eulaema sp. (8) Melipona fasciata (12) Melipona sp. ( 1 3 1 ) 0.014 inconclusive species in the Solanaceae, which share many of the same pollinators, flower simultaneously is consistent with Feinsinger's (1987) obser- vation that pollinator-sharing between sympatric species does not always cause temporal segregation of flower production. Fruit production Vertebrates are the dispersal agents for the majority of Neotropical plant species (Howe & Smallwood, 1982; Janzen, 1983; Murray et al., 2000). For species dependent on biotic dispersal agents, optimi- zation of seed dispersal may involve staggering the time of fruit maturation to avoid interspecific competition for seed-dispersers (Levin, 1978; Fleming, 1985). Staggering of fruiting season has been recorded in the tropical forests of Trinidad in species of Miconia (Melastomataceae), whose fruits are dispersed by frugivo- rous birds (Snow, 1965). At LCRS, it appeared that time of fruit maturation in species potentially sharing the same dispersal agents was somewhat temporally separated. For instance, the brightly- coloured fruit of Solarium cordovense matured in late June and early July, those of S. lanceifolium in mid- July and those of Lycianthes gorgonea in August. By separating the time of fruit maturity, Solanum and Lycianthes species may maximize seed dispersal and provide a more constant supply of food for frugivorous birds and bats. However, several years of monitoring would be needed to test the staggering of fruit production and dispersal, as variability in seasonal patterns can be high (Wheelwright, 2000). Flower visitors Bees, particularly Xylocopa, Eufriesia, Euglossa, Eulaema, Melipona, Trigona and Exomalopsis species, are extremely import- ant pollinators in neotropical forests (Endress, 1994). At La Selva, Costa Rica, for instance, Kress & Beach (1994) record that bees pollinate 38.4% of all plant species, making them the most numeri- cally important pollinators. Our observations at LCRS revealed a diverse array of bees in the Colletidae, Halictidae and Apidae visiting Solanum and Lycianthes, including many genera observed to pollinate Solanaceae in other tropical habitats (Knapp, 1986; Storti, 1988). Large showy white-flowered species, like S. lanceifolium and L. hypoleuca, attracted mainly medium and large bees. As the flowers of these two species usually lasted for one day only and opened around sunrise, flower visitors could depend on fresh flowers full of pollen each day and responded by returning daily to forage early in the morning. Longer-lived flowers did not draw such a loyal following. The flowers of 5. erythrotrichum and S. nudum, for example, attracted mostly Paratetrapedia sp., the small promiscuous bee. Because the flowers opened more or less at random and remained open for several days, bees could not depend on a reliable supply of fresh pollen at a certain time of day. Thus, foraging activity on these species with 'long-lived' flowers was spread throughout the morning and early afternoon. The pollination ecology of two study species remains unclear. Very few visits were observed on individuals of Solanum cordovense. Although its flowers are morphologically similar to 5. nudum, its small display failed to attract a similar suite of flower visitors. No visitors to Lycianthes gorgonea were observed during the study. Its few scattered individuals also produced a small display, never more than ten flowers at a time. Pre-dawn pollination, as observed in Lycianthes hypoleuca, is not unknown in buzz-pollinated Solanaceae. Linsley & Cazier (1963) recorded pollination of Solanum elaeagnifolium Cav. and S. rostratum Dunal by Ptiloglossa and Caupolicana (Colletidae) up to an hour before dawn in desert habitats in Arizona. Our finding that flowers of Lycianthes hypoleuca open in the darkness and are pollinated before sunrise is particularly interesting because some members of the genus appear to open exclusively at night. Many white-flowered species of Lycianthes are nocturnal or crepuscular bloomers (Benftez & D' Arcy, 1997), and flowers are closed during the day (Nee, 198 1 ). No studies of flower longevity or movements have been undertaken with these species however, so it is unclear whether L. hypoleuca is unusual in its floral phenology. Members of Lycianthes section Meizodontae, a small group of primarily Mexican herbs, have flowers that open at dawn and close during the day (Dean, pers. comm., July 2000). These flowers are pollinated by small solitary bees, and open and close over the course of several days. This diversity of floral phenologies in the genus Lycianthes suggests adaptive radiation to access different suites of pollinators. 136 S.D. SMITH & S. KNAPP ACKNOWLEDGEMENTS. We thank Chris Minty and Nicodemas 'Chapal' Bol for facilitating fieldwork at LCRS; George Else (The Natural History Museum) and Chris O'Toole (University of Oxford) who kindly identified the flower visitors; the Belizean Ministry of Natural Resources for permis- sion to collect voucher specimens; and the helpful comments provided by two anonymous reviewers. SS would like to express her immense gratitude to Dr Barbara Pickersgill for patient and critical review of her Master's thesis on this subject, to Dr Roger Smith for his support and editorial assistance during the development of this paper, and to Stephen Hague for technical assistance at LCRS during part of the study. 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A hypothesis about the timing of flowering and fruiting in competing tropical trees. In N.M. Nadkarni & N.T. Wheelwright (Eds), Monteverde: ecology and conservation of a tropical cloud forest: 281-282. New York. Wright, A.C.S., Romney, D.H., Arbuckle, R.H. & Vial, V.E.. 1959. Land in British Honduras. London. Bull. nut. Hist. Mus. Land. (Bot.) 32(2): 137-151 Issued 28 November 2002 The diatom type slides and bibliography of John Carter (1908-1993) DAVID M. WILLIAMS AND GERALDINE REID Botany Department, The Natural History Museum, Cromwell Road, London, SW7 5BD (DM. Williams@nhm.ac.uk) CONTENTS Introduction Carter's new taxa: types and illustrations Bibliography References ... 137 137 150 150 SYNOPSIS. An inventory of John Carter's diatom type slides is given along with a complete bibliography. Details of the whereabouts of Carter's type slides are included along with their geography, references to sources of additional illustrations, and nomenclatural notes. INTRODUCTION 'In the heighday of the amateur naturalist nearly everyone who possessed a microscope, however humble the instrument, treasured in his cabinet of slides a few diatom specimens' (Carter, 197 \b: 66). So wrote John Carter in a short but significant piece on the role of amateur diatomists and the legacy of good specimen collections (Carter, 1971ft). After his death in 1993 (Marson, 1994; Hartley, 1994), Carter's diatom collection was donated to The Natural His- tory Museum, where it now resides. Carter's collections have added significantly to the data on the British diatom flora, of which Carter's collection is largely composed. In all, the collection con- sists of some 5000 microscope slides and 4000 bottled samples, along with a variety of notebooks and a card index. The early part of the collection consists of representative material from Rev. Dingley Fuge, an early inspiration for Carter. This material consists of some 1 200 bottles that were never made into slides for Carter's collection. Fuge's collection will be the subject of a further paper. The documentation included with the material makes the collec- tions relatively easy to work with and, along with the recently published collection of drawings (Sims, 1996), forms a major record for the British freshwater diatom flora, spanning a period of six decades (1930- 1990). Carter did not just study the British diatom flora but made a contribution to diatom taxonomy in many areas of the world, includ- ing places that have never received attention before. As a consequence of these activities, as well as the detailed study of the British flora, Carter inevitably described a large number of new taxa. This paper presents a compilation of all new taxa Carter described with details of type specimens and slides. In most cases it was relatively easy to trace the holotype specimen(s) if it was not stated at the time of publication. In Carter's earlier papers where the holotype was not usually cited, slides that may be types have been noted. In most cases a lectotype has not been selected, as that task is best left to those more familiar with the particular diatom under consideration. In some cases, for instance, the comprehensive Sttsswasserflora von Mitteleuropa (Krammer & Lange-Bertalot, 1986-1991), lectotypes have already been selected and these are indicated in the text below. In some early papers, Carter noted that some slides were sent to the Quekett Microscopical Society Club. A preliminary exam- ination of those collections has not revealed the material. We have marked these types as 'holotype?' simply because, if found, the Quekett material may very well be the holotype. In most cases we have discovered identical material in the NHM collections which may serve as lectotypes should the need arise. The nomenclature has been revised in cases where necessary. CARTER'S NEW TAXA: TYPES AND ILLUSTRATIONS Achnanthes Bory 1822 A. aperta J.R. Carter in J.R. Carter & Bailey-Watts 1981 : 529, pi. 2, fig. 8. Holotype: 'collection Carter, Shet. No. 68a' = BM 87975. Locality: 'In Chumlie Loch, Shetland.' Additional illustrations: Lange-Bertalot & Krammer (1989: 113, taf. 89, figs 13-13') as Achnanthes oestrupii var. aperta (J.R. Carter) Lange-Bert., '...Holotypus, Coll. Carter Shetland 65A [sic, 68a], Chumlie Loch'; Krammer & Lange-Bertalot (199 la: 82, 344, taf. 48, figs 15-16) as, Achnanthes oestrupii var. aperta (J.R. Carter) Lange-Bert., '...Holotypus'; Sims (1996: 18-19, fig. 3; 2 images). Note: Lange-Bertalot & Krammer (1989: 1 1 3) place this taxon in Achnanthes oestrupii var. aperta (J.R. Carter) Lange-Bert. A. atalanta J.R. Carter 1966: 444, pi. 1, figs 15-16, 19-20. Syntypes:EM 7759 1-2. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae ... Sample No. 6. Crater Lake, on moss, 6500 feet above sea level.' [BM 77591] 'Sample No. 7. As number 6 but on algae.' [BM 77592]. Additional illustrations: Lange-Bertalot & Krammer (1989: 23, 63, 204, taf. 1 9, figs 2 1 -25' ), as a possible synonym of Achnanthes helvetica (Hust.) Lange-Bert., Typenprap. BM 77591, Tristan da Cuhna.' © The Natural History Museum, 2002 138 D.M. WILLIAMS AND G. REID A. cassida J.R. Carter 19706: 608, pi. 1, figs 17-18. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer (1989: 37, 133, taf. 48, figs 9-21), '...Holotypus (von A. cassida) BM 77780. . .' as Achnanthes rossi Must. Note: Bukhtiyarova & Round (1996: 12) place this taxon in Psammothidium rossi (Hust.) Bukht. & Round. A. curtissima J.R. Carter 1963: 200, pi. 1, fig. 3. Holotype: Not given. Lectotype: BM 93606. Carter 2630 'Fell Loch Wigstown-shire, 3/63', see below. Locality: 'Fell Loch, Wigtownshire. Fresh water, probably not a rare form.' Additional illustrations: Krammer & Lange-Bertalot ( 1 99 la: 2 1 , 344, taf. 11, fig. 8), 'Holotypus, coll. Carter 2630, Schottland' = BM 93606. Slide marked as lectotype by Carter. It is worth noting that on slide 4774 (BM 96179, 'Sheigra, Shetland, Bog') Carter wrote in his index 'I still have difficulty with this form [A. curtissima]. These specimens are my idea of it.' This material is not related to type material; Sims (1996: 20-21, pi. 2, fig. 13, 4 images). A. didyma f. gibbosa J.R. Carter in J.R. Carter & Bailey- Watts 1 98 1 : 530, pi. 23, figs 17-18. Holotype: Not given. Locality: Not given. Additional illustrations: Sims (1996: 22-23, pi. 3, fig. 3A, 2 images). Note: As synonym of A. rosenstockii Lange-Bert. (Lange-Bertalot & Krammer, 1989: 131, no illustrations). A. exiloides J.R. Carter in J.R. Carter & Bailey-Watts 198 1 : 532, pi. 1, figs 40-43. Holotype: 'collection Carter, Shet. No. 35' = BM 87947. Locality: 'In Kirk Loch, Shetland.' Additional illustrations: Lange-Bertalot & Krammer (1989: taf. 67, figs 20-23') as 'A. expressa Carter (syn. A. exiloides Carter). . .Holotypus, Coll. Carter, Kirk Loch/Shetland' ; Krammer & Lange-Bertalot (199 la: taf. 31, figs 13-18), Typenpraparat von A. exiloides Carter nom. nov. = A. expressa . . . ' ; Sims ( 1 996: 22-23, pi. 3, fig. 10, 2 images). Notes: Noted in Lange-Bertalot & Krammer (1989: 55) as 'A. expressa Carter in lift. ' and in Krammer & Lange-Bertalot ( 1 99 la) as 'A. expressa Carter nom. nov.' but, in fact, the correct citation is A. expressa J.R. Carter in Hartley (see below). A. expressa J.R. Carter in Hartley 1986: 606 = A. exiloides J.R. Carter in J.R. Carter & Bailey-Watts (1981) non A. exiloides Cholnoky(1963). A.fugeii J.R. Carter 1963: 203, pl.l, fig. 7. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: 'Estuary of the Forth. Probably a brackish water form.' Additional illustrations: Lange-Bertalot & Krammer (1989: 58, taf. 32, figs 24-24'), 'Coll. Carter 39A, Burkwell/Shetland.' Not type material; Sims (1996: 24-25, pi. 4, fig. 7, 2 images). Note: There are no relevant slides in Carter's collection. A.fulla J.R. Carter 19706: 608, pi. 1, figs 19-20. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic . . . collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer (1989: 59, taf. 67, figs 1-6), 'Holotypenprap. BM 77780, Pyrenaen/ Andorra.' A. hirta J.R. Carter 19706: 609, pi. 1, figs 10-11. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer (1989: 65, taf. 40, figs 10-12), 'Holotypus, BM 77780, Andorra', taf. 31, figs 3-4, '...Population aus den Pyrenaen/Andorra (leg. J. Carter)'; Krammer & Lange-Bertalot (1991a: taf. 18, figs 1^), 'Holotypus von A. hirta Carter, Andorra.' A. interrupta J.R. Carter 19616: 18, figs 49-50. Holotype: Not given. Locality: 'Found in Strath Farrar in a sphagnum bog at 2000 feet above sea level.' Notes: Three slides in Carter's BM collection seem appropriate for selection of a lectotype: BM 93089, 'W Strath Farrar, 2276', BM 93090, 'Strath Farrar, 2276A, BM 93091, 'N Strath Farrar, 2276F'. These slides are from the same material (Carter 2276) although in the accompanying notebook the drawings given in the publication are not included in the notes for sample 2276. A. investians J.R. Carter 1966: 445, pi. 1, figs 23-28. Lectotype: BM 77587. After Lange-Bertalot & Krammer (1989; see below). Syntypes: BM 77588, BM 77593, BM 77594. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 2. Above Burntwood, stream at 1800 feet above sea level.' [BM 77587] 'Sample No. 3. As number 2 but a little upstream in a pool' [BM 77588] 'Sample No. 8. Salt beach at sea level.' [BM 77593] 'Sample No. 9. Hottentot Gulch, stream pool, 2000 feet above sea level.' [BM 77594]. Additional illustrations: Carter (1966: 483) as A. investians n. sp. forma, pi. 8, figs 13-14 '...form from Gough Island sample 12...Gough Island sample 1006, Steam Gonydale, 1400 feet a.s.l.' [BM 77600]. Lange-Bertalot & Krammer (1989: 71, taf. 18, figs 27-29'), 'Typenprap. BM 77587, Tristan da Cunha'. Note: Bukhtiyarova & Round (1996: 26) place this taxon in Psammothidium investians (J.R. Carter) Bukht. & Round. A. lanceolata f. rhombica J.R. Carter 1966: 446, pi. 1, figs 17-18. Holotype: BM 77586. Locality: 'Sample No. 1. Pigbite, stones in running water at sea level.' A. lintonensis J.R. Carter 1961a: 325, figs 15-16. Holotype: Not given. Locality: 'This species is rare and local to areas of the Cheviots.' Additional illustrations: Sims (1996: 26-27, pi. 5, fig. 11, 2 images). Note: One slide in Carter's collection might be an appropriate lectotype: BM 94111, 'Cheviots, 3019'. A. lucana J.R. Carter 19706: 609, pi. 1, figs 8-9. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' A. microcephala f. scotia J.R. Carter in J.R. Carter & Bailey-Watts 1981:534, pi. 1, fig. 31. CARTER DIATOM TYPE SLIDES Holotype: Not given. Locality: Not given. Lectotype: BM 94890 (Carter 3665), after Lange-Bertalot in Lange-Bertalot & Krammer (1989: 300). Carter noted on slide 3665 'fo. scotia in plenty.' Locality: Ardislaigh Lochan, June '76. Additional illustrations: Lange-Bertalot & Krammer ( 1 989: 106, taf. 55, figs 1 1-12), as Achnanthes minutissima var. scotica (J.R. Carter) Lange-Bert., 'Achnanthes microcephala f. scotia Carter, Coll. Carter 3665.' = BM 94890 and Taf. 67, figs 35-36, 'Coll. Carter 3665 Ardislaigh Lochan/Schottland.' = BM 94890; Krammer & Lange-Bertalot (199 la, taf. 31, figs 1-6), 'Coll. Carter aus Schottland und Brunnsee/Oberbayern.' [Not indi- cated which figures are from Carter material.] Notes: Lange-Bertalot & Krammer (1989: 106) placed this taxon in Achnanthes minutissima var. scotia (J.R. Carter) Lange-Bert. and later elevate it to species-level, as Achnanthes scotia (J.R. Carter) Lange-Bert. (Lange-Bertalot, 1993: 8). However, Achnanthes scotia had already been used by Flower & Jones (1989: 228) hence Lange-Bertalot proposed the substitute name Achnanthes caledonica Lange-Bert. in Lange-Bertalot & Moser (1994:95). A. minima J.R. Carter 196 \b: 21, figs 65-66. Holotype: Not given. Locality: 'I have found this species in Perthshire and Galloway but only as single cells.' A. miota J.R. Carter & Denny 1982: 286, pi. 1, fig. 9. Holotype: BM78107. Locality: 'River Jong (Taia) at Njala [Sierra Leone] ... phyto- plankton.' Additional illustrations: Lange-Bertalot & Krammer (1989: 107, taf. 87, figs 5-9) as Achnanthes lanceolata var. apiculata R.M. Patrick, '...Typenprap. BM 78107, Sierra Leone'; Krammer & Lange-Bertalot (1991a: taf. 46, figs 19-21), 'Typenprap. Sierra Leone/Afrika.' Note: As Achnantheiopsis miota (J.R. Carter) Lange-Bertalot (1997: 207) and Planothidium miotum (J.R. Carter) Lange- Bertalot (1999: 278), the latter being the current, accepted name. A. natrata J.R. Carter 1966: 446, pi. 1, figs 1-2. Holotype: BM 77593. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae . . . Sample No. 8. Salt beach at sea level.' A. parallela J.R. Carter 1963: 200, pi. 1, fig. 4. Holotype?: Slides in Quekett Microscopical Society Collections. Syntype?: BM 93945 = Carter 2904, 'Lindean - on Burrweed', specimen marked. Locality: 'Well distributed but never in plenty in the hill lochs of the Southern Scottish uplands and the mountainous districts of south-west Scotland.' Additional illustrations: Krammer & Lange-Bertalot ( 1 99 1 a: taf. 37, figs 24-27), 'Coll. Carter 2904, Schottland' as A. petersenii Hust.; Sims (1996: 32-33, pi. 8, fig. 5, 4 images). Notes: A. sonyda J.R. Carter = A. parallela J.R. Carter non A. parallela Castr. ( 1 886). As Achnanthes petersenii Hust. in Lange- Bertalot & Krammer (1989: 117). Carter 2904 is noted on Carter's card file as 'type material' and in the accompanying notebook as 'Pub as parallela JQMC 1963 Vol. 29.' A. perfida J.R. Carter 1963: 200, pi. 1, fig. 2. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: 'Loch Hempton on the surface of peat, fresh water, not common.' 139 Additional illustrations: Sims (1996: 30-31, pi. 7, fig. 5, 2 images). Notes: See Navicula perfidissima Lange-Bert. There are two slides in Carter's collection which may also be type material: BM 93425, 'Loch Hempton, 2497, BM 93597, 'Loch Hempton, on Weed, 2625, 5/63'. A.pericava J.R. Carter 1966: 447, pi. 1, figs 5-8. Holotype: BM 77593. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 8. Salt beach at sea level.' Additional illustrations: Lange-Bertalot & Krammer ( 1 989: 1 1 6, 162, taf. 79, figs 40-44'), 'Holotypus, BM 77593, Tristan da Cuhna.'; Krammer & Lange-Bertalot (199 la: taf. 40, figs 14- 16), 'Holotypus, Tristan da Cuhna.' Note: As Achnantheiopsis pericava (J.R. Carter) Lange-Bert. (1997: 207) and Planothidium pericavum (J.R. Carter) Lange- Bert. (1999: 278), the latter being the current, accepted name. A. piafica J.R. Carter & Denny 1982: 286, pi. 1, fig. 10. Holotype: BM 78108. Locality: 'River Jong (Taia) at Njala [Sierra Leone] ... phyto- plankton.' Additional illustrations: Lange-Bertalot & Krammer (1989: 119, taf. 85, fig. 9 and taf. 87, fig. 10), both Typenprap. BM 78108, Sierra Leone.' Note: Possible synonym of Achnanthes lanceolata var. rostrata (0strup) Hust. according to Lange-Bertalot & Krammer (1989: 90). A. pictii J.R. Carter 1963: 203, pi. 1, fig. 8. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: 'Solway mud flats, not uncommon.' Notes: Synonym of Achnanthes linkei Hust. (Lange-Bertalot & Krammer, 1989: 97, see also p. 119). There are two slides in Carter's BM collection which may also be type material: BM 93583, 'Solway, Mud flats, 2623, 5/63', BM 93584, 'Solway, Mud flats, 2623a, 5/63.' A. plana J.R. Carter 1963: 201, pi. 1, fig. 6. Holotype: Not given. Lectotype: BM 93605 = Carter 2629, 'L. Eidrig, 5/63', marked on Carter's card file as 'type slide'. Locality: 'Loch Eidrig, fresh water, very rarely.' Additional illustrations: Lange-Bertalot & Krammer (1989: 90, 120, taf. 88, figs 22-23), '...Holotypus, Coll. Carter 2629 Loch Eidrig/Schottland' as Achnanthes lanceolata var. robusta Hust.; Sims (1996: 30-31, pi. 7, fig. 7, 2 images). A. pseudoswazi J.R. Carter 1963: 201, pi. 1, fig. 5. Holotype: Not given. Locality: 'Perhaps widely distributed in the larger northern lakes but always odd cells. I have it from regions as far apart as L. Tummel in the north to the Border Country.' Additional illustrations: Lange-Bertalot & Krammer (1989: 25, taf. 45, figs 37-38), not type material; Krammer & Lange- Bertalot (1991a: taf. 24, figs 1-2), 'Holotypus.' Krammer & Lange-Bertalot do not identify a holotype and, as it was not given in the original description, these illustrations may not be from type material; Sims (1996: 30-31, pi. 7, fig. 12, 3 images). Note: Bukhtiyarova & Round (1996: 20) place this taxon in Psammothidium pseudoswazi (J.R. Carter) Bukht. & Round. A.pyropa J.R. Carter 19706: 610, pi. 1, figs 1-2. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the 140 D.M. WILLIAMS AND G. REID only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer ( 1 989: 1 28, taf. 67, figs 19-19'), 'Coll. Carter 13926, Pyrenaen/Andorra' = BM 77780. A. saccula J.R. Carter in J.R. Carter & Bailey- Watts 1981 : 535, pi. 24, fig. 6. Holotype: 'collection Carter, Shet. No. 25.' = BM 87942. Locality: 'At Funds, Shetland.' Additional illustrations: Lange-Bertalot & Krammer (1989: 134, taf. 27, figs 20-20'), 'Holotypus, Coll. Carter Shetland 25, "Funds'" = BM 87942; Krammer & Lange-Bertalot (199 la: taf. 15, figs 19-20), '...Holotypus, Shetland.'; Sims (1996: 32-33, pi. 8, fig. 1, 2 images). Note: Bukhtiyarova & Round (1996: 27) place this taxon in Psammothidium saccula (J.R. Carter) Bukht. & Round. A. sonyda J.R. Carter in Hartley 1986: 606 = A. parallela Carter non A. parallela Castr. (1886). A. sphacelata J.R. Carter 19706: 611, pi. 1, figs 41-44. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer (1989: 143, taf. 67, figs 10-13), 'Holotypenprap. BM 77780'. A. sumara J.R. Carter 1966: 447, pi. 1, figs 21-22. Holotype: BM 77586. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 1. Pigbite, stones in running water at sea level.' Note: As Achnanthes petersenii Hust. in Lange-Bertalot & Krammer (1989: 117). A. sutura J.R. Carter 19706: 61 1, pi. 1, figs 39^0. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer (1989: 145, 149, taf. 21, figs 4-4'), '...Typenprap.BM 77780' as Achnanthes subatomoides (Hust.) Lange-Bert. & R.E.M. Archibald. A. taiaemis J.R. Carter & Denny 1982: 288, pi. 1, figs 11-15. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' Additional illustrations: Lange-Bertalot & Krammer (1989: 150, taf. 67, figs 25-28), 'Holotypus, BM 78107, Sierra Leone'. A. tuma J.R. Carter 19706: 612, pi. 1, figs 35-36. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer (1989: 154, taf. 67, figs 14-18), 'Holotypenprap. BM 77780.' A. umara J.R. Carter 19706: 612, pi. 2, figs 1-2. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Additional illustrations: Lange-Bertalot & Krammer (1989: 145, 154, taf. 21, figs 3-3'), '...Typenprap.BM 77780' asAchnanthes subatomoides (Hust.) Lange-Bert. & R.E.M. Archibald; Sims (1996: 32-33, pi. 8, fig. 13, 4 images). A. wickenensis J.R. Carter 1963: 199, pi. 1, fig. 1. Holotype?: Slides in Quekett Microscopical Society Collections. Lectotype: BM 93539 = Carter 2591 (see below). Locality: 'Found very sparingly in Wicken Lode. Fresh water July.' Additional illustrations: Lange-Bertalot & Krammer (1989: 122, 156, taf. 41, figs 17-17'), '...Coll. Carter 3591 [sic, 2591] "Wicken",' as Achnanthes ploensis var. gessneri (Hust.) Lange- Bert.; Sims (1996: 32-33, pi. 8, fig. 14, 2 images). Note: There are four slides in Carter's BM collection which may also be type material: BM 93586, 'Wicken Lode, Cambs'. BM 93587, 'Wicken Lode, Cambs', BM 93588, 'Wicken Lode, Cambs', BM 93539, 'Wicken Lode, 259 la.' Amphora Ehrenb. ex Kiitz. 1 844 A. eximia J.R. Carter in Haworth 1974: 48, figs 3, 14. Holotype: BM 77902. Locality: 'Loch Leven, 20cm.' Additional illustrations: Sims (1996: 52-53, pi. 18, figs 8-9, 5 images). A. incurvata J.R. Carter 1966: 448, pi. 3, figs 29-31. Holotype: BM 77593. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample no. 8. Salt beach at sea level.' Anomoeneis E. Pfitzer 1 87 1 A. serians var. brachysira f. in i nut a J.R. Carter 196 Id: 75, fig. 13. Holotype: Not given Locality: '. . .from the bed of Loch na Cuilic.' Note: There are a number of slides in the collection from this locality. A possible lectotype is BM 93364, which is the sample cleaned by Mr S. Meakin which 'he has separated the very light forms' (Carter 196 Id: 74) although in Carter's notebook there is no mention of a new form in relation to this slide. Another possible lectotype is BM 93297-8. These are the slides Carter prepared from the cleaned material and in his notebook he wrote 'list for publication... list sent 17/6/61.' The list does not include f. minuta but does note the occurrence of a new variety. Asterionella Hassall 1850 A. contorta J.R. Carter & Denny 1987: 234, pi. 1, figs 7-8. //0/ory/*?:BM81159. Locality: '. . .coastal strip of South- Western Sierra Leone. . .Sample 68 (16th April 1976)...Phytoplankton horizontal haul... Lake Mape.' Berkella R. Ross & P. A. Sims 1978 B. alpina J.R. Carter ex D.M. Williams nov. sp. Description: In Carter (1993a: 106, figs 24-25). Holotype: BM 94636 = Carter no. 3462. As Berkella alpina (Amosse) J.R. Carter in Carter (1993a: 106, figs 24-25) and Berkella spicula (Amosse) J.R. Carter in J.R. Carter & Bailey- Watts (1981: 539, pi. 2, fig. 38, pi. 24, fig. 12). Locality: 'Ettrick Valley, S. Scotland; August 1974.' Additional illustrations: Sims(1996: 84-85, figs 8- 15, 16images as Berkella spicula var. alpina Amosse); Krammer & Lange- Bertalot (1986: 634, fig. 7), 'Shetlands, Pr. Carter') as Frustulia spicula Amosse. Notes: Ross & Sims (1978: 1 56) described the new genus Berkella R. Ross & PA. Sims basing it on specimens of B. linearis R. Ross CARTER DIATOM TYPE SLIDES & P.A. Sims, discovered in Loch Poit na h'l. Mull, Scotland. Although they write that their species 'was first described by Amosse under the name Frmtulia spicula van alpina . . . ' his name was invalid as 'he did not designate a type'. They continue: '[we have] adopted a new epithet to make it quite clear that we are not basingourspeciesonAmosse'svariety'(Ross&Sims, 1978: 158). Carter concluded, after examination of a variety of specimens, that Frustulia spicula var. alpina was indeed another species of Berk- ella, distinct from B. linearis. Carter described his Berkella alpina as a new combination but as Frustulia spicula var. alpina had no designated type, it should have been described as a new species. Caloneis Cteve 1894 C. borealis J.R. Carter 197 la: 660, pi. 1, figs 1-3. Syntypes: EM 77777-8. Locality: 'Devil's Hole Cave. ' '...Kincraig Hill at the south- western extremity of Kilconquhar parish, Fife. . .collected by the late Mr. Geo. West of University College, Dundee, in the second decade of this century [1920s].' Additional illustrations: Sims (1996: 96-97, pi. 40, figs 9-10, 6 images). C. continua J.R. Carter 1964: 225, pi. 2, fig. 9. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: 'Loch Maben Dumfriesshire, fresh water, seemingly somewhat rare.' Note: There are nine slides in Carter's BM collection which may be type material: BM 92993, 'Loch Maben, 2204', BM 93261, 'Loch Maben, Mud, 2371', BM 93262, 'Mabon, 2371 A, BM 93263, 'Loch Maben, Mud, 237 1C', BM 93264, 'Loch Maben, 2371P', BM 93265, 'Loch Maben, 2371R', BM 93282, 'Loch Maben, Mud, 2387', BM 93569, 'Loch Maben, 2612, BM 93570, 'Loch Maben, 2613'. C. inliberalis J.R. Carter & Denny 1992: 164, pi. 1, fig. 23. Holotype: BM 81522. Locality: 'Lake Popei... sample 54 [Sierra Leone].' C. lauta J.R. Carter in J.R. Carter & Bailey- Watts 198 1 : 540, pi. 23, figs 6, 8. Holotype: 'collection Carter, Shet. 11.' = BM 87928. Locality: 'In Burga Water, Shetland.' Additional illustrations: Krammer & Lange-Bertalot (1986: 389, taf. 173, fig. 3), 'HolotypusShetlands, Burga Water, Coll. Carter'; Sims (1996: 100-101, pi. 42, fig. 8, 4 images). C. lepidiformis J.R. Carter & Denny 1987: 235, pi. 1, fig. 12. //0/ofy/7:BM81509. Locality: 'In sample 5 quite commonly [Lake Sonfon, stream, Sierra Leone].' N. bacillum f. deticatula J.R. Carter & Denny 1982: 302, pi. 5, fig. 175. Holotype: Not given. Locality: Not given. Note: This taxon should belong in the genus Sellaphora Mereschk. N. belsesiensis J.R. Carter 1984: 134, figs 5-7a. Holotype: BMSmS. Locality: '...small ditch near Belses on the Scottish side of the Border.' N. bintila J.R. Carter & Denny 1982: 302, pi. 6, fig. 190. Locality: 'River Jong (Taia) at Njala [Sierra Leone] . . .from stones.' N. bottnica f. acuta J.R. Carter & Denny 1992: 181, pi. 5, fig. 2. Holotype: BM 81508. Locality: '...in sample 4... [Lake Sonfon, Sierra Leone]' N. brockmannii f. concava J.R. Carter 1966: 460, pi. 3, fig. 14. Holotype: BM 77590. Locality: 'Sample No. 5. Big Waterburn, stones in stream, 100 feet above sea level.' N. caenosus J.R. Carter 1.. ;.R. Carter & Bailey- Watts 198 1 : 574, pi. 11, fig. 41. Holotype: 'collection Carter, Shet. No. 16a.' = BM 87933. Locality: 'In Houlma Water, Shetland.' Additional illustrations: Sims (1996: 282-283, pi. 133, fig. 5, 2 images; 356-357, pi. 170, fig. 11, 1 image). N. carterii VanLand. (1975: 2520) see N. dispersa J.R. Carter. N. chaspula J.R. Carter & Denny 1982: 304, pi. 6, fig. 191. Syntypes: BM 78108, BM 78112. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from phytoplankton.' [BM 78108] 'River Jong (Taia) at Njala [Sierra Leone]... from rotting leaves.' [BM 78112]. N. claytoni J.R. Carter 1987: 633, figs 1-10. Syntypes: BM 8 14 13-4. Locality: 'In Gala Water inventa...In February 1986 Mr John Clay ton... took a sample of diatoms from the tributary of the River Tweed, the Gala Water.' Additional illustrations: Sims (1996: 284-285, pi. 134, fig. 5, 2 images). N. cocconioides J.R. Carter 1964: 227, pi. 2, fig. 13. Holotype?: Slides in Quekett Microscopical Society Collections. Syntypes: BM 93582, 'Madingly Brick Pit, Cambridge, 2622, 5/ 63'; BM 93794, 'Maddingly, Brick Pit, on Liverworts, 2777, 5/ 63.' Locality: 'Madingly Brick Pit, Cambridge.' Additional illustrations: Sims (1996: 286-287, pi. 135, fig. 2, 1 image). Notes: non Navicula cocconeoides (Rabenh.) Ralfs in A. Pritch. (1861: 909). Although the spelling of the specific epithet for Carter's taxon is similar to that of Ralfs, the two taxa would be hard to confuse. While there may be justification in re-naming Carter's taxon, we have chosen to retain it. N. concentrica J.R. Carter in J.R. Carter & Bailey-Watts 198 1 : 576, pi. 11, fig. 7. Holotype: 'collection Carter, Shet. No. 2.' = BM 87920. Locality: 'In Blister Funds, Shetland and several other locations.' Additional illustrations: Sims (1996: 286-287, pi. 135, fig. 6, 3 images). Note: non N. concentrica Bailey (1844: 139, pi. 3, figs 13-15). Inspection of Bailey's illustrations of his Navicula concentrica clearly show specimens of Goniothecium Ehrenb. while Carter's Navicula concentrica is certainly a species of Navicula sensu stricto. Nevertheless, Bailey's name still has priority hence Carter's taxon is renamed here as Navicula johncarterii D.M. Williams, nov. nom. N. crepitacula J.R. Carter & Denny 1982: 304, pi. 6, fig. 189. Locality: 'River Jong (Taia) at Njala [Sierra Leone]. . .from stones.' N. cuvella J.R. Carter & Denny 1982: 306, pi. 6, figs 193-194, 218. Holotype: BM78108. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from phytoplankton.' N. decissa J.R. Carter 1966: 461, pi. 3, fig. 11. Holotype: BM 77587. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 2. Above Burntwood, stream at 1800 feet above sea level'. N. dispersa J.R. Carter 1964: 225, pi. 2, fig. 10. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: 'Loch Hempton in the peat, fresh water, not common.' Additional illustrations: Sims (1996: 282-283, pi. 133, fig. 12, 1 image). Note: N. carterii VanLand. (1975: 2520) = N. dispersa J.R. Carter non N. dispersa Grove & Sturt (1887: 132, pi. 10, fig. 10) nee N. dispersa Manguin (1942: 137, pi. 2, fig. 32). Should the Quekett slides not materialize then BM 93597 will be a suitable lectotype: 'Loch Hempton, on Weed, 2625, 5/63, Na- vicula carterii.' N. diverta J.R. Carter 1966: 461, pi. 3, figs 19-20. Holotype: BM 77591. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 6. Crater Lake on moss, 6500 feet above sea level.' N. eidrigiana J.R. Carter 1979: 78, figs 58-64, 70-72. Holotype: BM 78083. CARTER DIATOM TYPE SLIDES 145 Locality: 'Loch Eidrig, south west Scotland. . .' (Carter, 1979: 78). Additional illustrations: Sims (1996: 296-297, fig. 1,9 images); Lange-Bertalot (2001: 34, pi. 43, figs 1-6). N.friska J.R. Carter 1966: 462, pi. 3, figs 7-10. Syntypes: BM 77587-8. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 2. Above Burntwood, stream at 1800 feet above sea level.' [BM 77587] 'Sample No. 3. As number 2 but a little upstream in a pool' [BM 77588]. N. glomus J.R. Carter in J.R. Carter & Bailey-Watts 1981 : 578, pi. 13, fig. 17. Holotype: 'collection Carter, Shet. No. 39a' = BM 87950. Locality. 'At Burkwell, Shetland.' Additional illustrations: Krammer & Lange-Bertalot (1986: taf. 71, fig. 39), 'Typenprap. Coll. Carter Shetland 39 a.' N. gralana J.R. Carter & Denny 1982: 306, pi. 6, fig. 195. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from rot- ting leaves.' N. lurinda J.R. Carter & Denny 1982: 308, pi. 6, fig. 201. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from phytoplankton.' N. insepta J.R. Carter & Denny 1982: 307, pi. 6, figs 197-198. Holotype: BM1SIOS. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from phytoplankton.' N. inexpectans J.R. Carter 1964: 227, pi. 2, fig. 12. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: 'Loch Mochrum fresh water as isolated cells.' Additional illustration: Sims (1996: 308-309, pi. 146, fig. 10, 1 image). Note: Two additional slides in the Carter collection may be type material: BM 93573, 'Loch Mochrum, on stones, 2616, 5/63'; BM 93578, 'Loch Mochrum, mud, 2619, 5/63.' N. involata J.R. Carter & Denny 1982: 307, pi. 6, fig. 199. Syntypes: BM 78 11 1-2. Locality: 'River Jong (Taia) at Njala [Sierra Leone] . . .from stones.' [BM 78111] 'River Jong (Taia) at Njala [Sierra Leone]... from rotting leaves.' [BM78112]. N. johncarterii DM. Williams, nov. nom. = N. concentrica J.R. Carter in J.R. Carter & Bailey- Watts (1981: 576, pi. 1 1, fig. 7) non N. concentrica Bailey (1844: 139, pi. 3, figs 13-15). N. laevissima var. semi-aperta H.G. Barber & J.R. Carter 197 \b: 82, fig. 97. Holotype?: Typus in collectione Quekett Microscopical Club.' Locality: '. . .fossil diatomaceous earth situated in Gordon Road, Auckland, New Zealand at co-ordinates 225,110E, 656,720N, Square G.7 of New Zealand M.S. 17, 1962 Edition 'Auckland'.' Note: This taxon should belong in the genus Sellaphora Mereschk. N. lapila J.R. Carter in J.R. Carter & Bailey-Watts 198 1 : 582, pi. 14, figs 34-35. Holotype: 'collection Carter, Shet. No. 4' = BM 87922. Locality: 'In Moo Water, Shetland.' Additional illustration: Sims (1996: 310-311, pi. 147, fig. 14, 1 image). N. lapsa J.R. Carter & Denny 1992: 184, pi. 5, figs 11-15. Holotype: BMS1509. Locality: 'In sample 5 [Lake Sonfon, stream, Sierra Leone].' N. lupula J.R. Carter & Denny 1982: 308, pi. 6, fig. 200. Holotype: BM 18112. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. malica J.R. Carter & Denny 1982: 308, pi. 6, figs 202-204. Syntypes: BM 78 107-8. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' [BM 78107] 'River Jong (Taia) at Njala [Sierra Leone]... phytoplankton.' [BM 78108]. N. menda J.R. Carter in J.R. Carter & Bailey-Watts 1981: 582, pi. 24, fig. 27. Holotype: 'collection Carter, Shet. No. 12' = BM 87929. Locality: 'In Voxterby, Shetland.' Additional illustrations: Sims (1996: 312-313, pi. 150, fig. 2, 2 images). Notes: Haworth & Offer (1986: 446) suggested two further slides as lectotypes '[BJecause the original description is based upon just one specimen...'. This reason is not a valid justification for the designation of lectotype material as Carter did identify the holotype slide and that has to remain so, regardless of how many specimens were consulted for the description. The two slides suggested by Haworth & Offer (BM 81209-10) may act as useful, additional non-type examples. N. mintracta J.R. Carter & Denny 1982: 309, pi. 6, fig. 205. Holotype: BMimi. Locality: 'River Jong (Taia) at Njala [Sierra Leone] . . .from stones.' N. nabrona J.R. Carter & Denny 1982: 309, pi. 6, fig. 206. Holotype: BM7&IV7. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. nabrona f. ovalis J.R. Carter & Denny 1982: 309, pi. 6, fig. 207. Holotype: Not given. Locality: Not given. Lectotype: BM 78107 ('River Jong (Taia) at Njala [Sierra Leone] . . .phytoplankton.'). Note: Carter writes 'occurs mixed with the type.' This seems to suggest that BM 78107 is the most suitable lectotype. N. nienta J.R. Carter 1966: 464, pi. 3, fig. 17. Holotype: BM 77590. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 5. Big Waterburn, stones in stream, 100 feet above sea level.' N. nutata J.R. Carter & Denny 1982: 309, pi. 6, figs 208-210. #0/007*-: BM 781 11. Locality: 'River Jong (Taia) at Njala [Sierra Leone] . . . from stones. ' N. orbita J.R. Carter 1966: 464, pi. 3, fig. 12. Holotype: BM 77594. Locality: 'In aquis dulcibus insulae Tristan da Cuhna dictae... Sample No. 9. Hottentot Gulch, stream pool, 2000 feet above sea level.' N. orbis J.R. Carter in J.R. Carter & Bailey- Watts 1981: 584, pi. 24, fig. 34. Holotype: 'collection Carter, Shet. No. 39a' = BM 87950. Locality: 'At Burkwell, Shetland.' Additional illustration: Sims (1996: 320-321, pi. 152, fig. 11,1 image). 146 N.patrina J.R. Carter & Denny 1982: 310, pi. 6, fig. 211. Holotype-.BMlSlll. Locality: 'River Jong (Taia) at Njala [Sierra Leone]. . .from stones.' N. perfidissima Lange-Bert. in Lange-Bertalot & Krammer (1989: 162, taf. 70, figs 22-25', ?26, 27). Holotype: BM 93708 = Carter 2716, 'Devils Hole (Tay Dist.) West Coll. A.W. Round no. 35' (Carter, 1971a: 659,Achnanthes perfida), two specimens marked (Lange-Bertalot & Krammer, 1989: taf. 70, figs 22-25), 'Als Achnanthes perfida Carter in Coll. Carter 2716, Devil's Hole/Schottland.' BM 93708 is not type material of A. perfida, see above. N. petilina J.R. Carter & Denny 1982: 310, pi. 6, fig. 212. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. pseudoclamans J.R. Carter in J.R. Carter & Bailey-Watts 1981: 585, pi. 14, fig. 7. Holotype: 'collection Carter, Shet. No. 4a' = BM 87922. Locality: 'In Moo Water, Shetland.' Additional illustrations: Sims (1996: 328-329, pi. 156, fig. 12, 2 images). N. pseudomenisculus J.R. Carter in J.R. Carter & Bailey- Watts 1981: 585, pi. 14, fig. 8. Holotype: 'collection Carter, Shet. No. 36' = BM 87948. Locality: 'In Loch of Cliff, Shetland.' Additional illustration: Sims (1996: 328-329, pi. 156, fig. 19, 1 image). Note: non N. pseudominiscula Fusey (1950: 156, fig. 22). Although the spelling of the specific epithet for Carter's taxon is remarkably similar to that of Fusey, these two taxa would be hard to confuse. While there may be justification in re-naming Carter's taxon, we have chosen to retain it. N. pseudostundlii H.G. Barber & J.R. Carter 19716: 83, fig. 122. Holotype?: Typus in collectione Quekett Microscopical Club.' Locality: '...fossil diatomaceous earth situated in Gordon Road, Auckland, New Zealand at co-ordinates 225,110E, 656J20N, Square G.7 of New Zealand M.S. 17, 1962 Edition 'Auckland'.' N. resecta J.R. Carter in J.R. Carter & Bailey-Watts 1981: 586, pi. 12, fig. 55. Holotype: 'collection Carter, Shet. No. 36' = BM 87948. Locality: 'In Loch of Cliff, Shetland.' Additional illustrations: Sims (1996: 336-337, pi. 160, fig. 4, 2 images). N. retried J.R. Carter & Denny 1982: 311, pi. 6, figs 214-215. Holotype: BM 18101. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. retrocurvata J.R. Carter ex R. Ross & P.A. Sims 1978: 159, figs 14-16. Holotype: BM 78070. Locality: '...ex rivulo 'on the links behind Sandside Bay, Main- land' in Orcades, J.R. Carter 3369 lectus.' Additional illustrations: Sims (1996: 312-313, pi. 148, fig. 11,3 images; 336-337, pi. 160, fig. 6, 2 images). Note: = Hippodonta lesmonensis (Hust.) Lange-Bert., Metzeltin &Witkowski( 1996: 261). N. riparta J.R. Carter 1966: 465, pi. 8, fig. 19. Holotype: BM 77599. D.M. WILLIAMS AND G. REID Locality: 'In aquis dulcibus insula Gough Island dictae. . .Sample No. 1 1 Gough Island sample 1003, Cave Bell Rocks, sea level.' N. rotesta J.R. Carter 19706: 620, pi. 2, figs 14, 32-33. Holotype: BM 77783. Locality: 'Sample No. 5 from the Rio d'Envalira... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' N. sarolata J.R. Carter & Denny 1982: 311, pi. 6, figs 216-217. Holotype: Not given. Lectotype:BM 18101. Locality: 'A few specimens seen. 11, 20' = 'River Jong (Taia) at Njala [Sierra Leone]... phytoplankton.' [Sample 11, BM 78107] 'River Jong (Taia) at Njala [Sierra Leone]... phytoplankton.' [Sample 11, BM 78108]; 'River Jong (Taia) at Njala [Sierra Leone]... from stones.' [Sample 20, BM 78111]. N. sarolata f. capitata J.R. Carter & Denny 1992: 185, pi. 5, figs 38- 39. Holotype: BM 81506. Locality: 'In sample 3 only [Lake Sonfon, Sierra Leone].' N. scalpia J.R. Carter 1966: 466, pi. 8, fig. 15. Holotype: BM 77600. Locality: 'In aquis dulcibus insula Gough Island dictae. . .Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' N. scela J.R. Carter & Denny 1982: 312, pi. 6, figs 187-188. Holotype: BM18U2. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from rot- ting leaves.' N. scirpus J.R. Carter in J.R. Carter & Bailey- Watts 1981: 586, pi. 24, figs 29-30a. Holotype: 'collection Carter, Shet. No. 26' = BM 87943. Locality: 'In North Vementry, Shetland.' Additional illustrations: Sims (1996: 340-341, pi. 162, fig. 8, 5 images). N. sequens H.G. Barber & J.R. Carter 19716: 83, fig. 110. Holotype?: 'Typus in collectione Quekett Microscopical Club.' Locality: '. . .fossil diatomaceous earth situated in Gordon Road, Auckland, New Zealand at co-ordinates 225,110E, 656J20N, Square G.7 of New Zealand M.S. 17, 1962 Edition 'Auckland'.' N. sponsa J.R. Carter 1964: 226, pi. 2, fig. 11. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: 'I have found this form in three locations in the Chevi- ots in every case on the surface of mud in high level springs.' Additional illustration: Sims (1996: 344-345, pi. 164, fig. 13, 1 image). N. subexilissima J.R. Carter 19706: 620, pi. 2, fig. 13. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' N. subvasta J.R. Carter 19706: 621, pi. 2, fig. 20. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' Notes: non N. subvasta Hust. Hustedt illustrated his N. subvasta in Schmidt's Atlas (1936: pi. 403, figs 49-50) but provided no description. While Hustedt did eventually provide a description, first he placed it under the name Navicula rivularis Hust. (1942: CARTER DIATOM TYPE SLIDES 147 112) and second, in 1945 (930, pi. 41, figs 38^0), he included it in Navicula helensis P. Schulz. Navicula helensis is now cor- rectly referred to as Fallacia helensis D.G. Mann (in Round et al. 1990: 668). As neither of Hustedt's descriptions can be under- stood as validly published and as the illustration in Schmidt's Atlas is after 1 908, Carter's name is the first valid use of Navicula subvasta. N. sulta J.R. Carter & Denny 1982: 314, pi. 8, fig. 290. Holotype: BM 78110. Locality: 'River Jong (Taia) at Njala [Sierra Leone]. . .from sub- merged wood.' N. suspica J.R. Carter & Denny 1992: 186, pi. 5, fig. 45. Holotype: BM 81522. Locality: 'In sample 54 rarely [Lake Popei, Sierra Leone].' N. tauta J.R. Carter & Denny 1982: 314, pi. 8, fig. 293. Holotype: BM78108. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. toxa J.R. Carter 19606: 284, fig. 6. Holotype: Not given. Lectotype: BM 9298 1 = Carter 2198, 'Woodhead Loch, Ancrum, Small Nav....' Locality: 'Abundant in Woodhead Loch, Roxburghshire, usually on the surface of mud.' N. variolinea J.R. Carter 1971a: 668, pi. 2, fig. 71, pi. 3, fig. 95. Syntypes: BM 77777-8. Locality: 'Devil's Hole Cave.' '...Kincraig Hill at the south- western extremity of Kilconquhar parish, Fife. . .collected by the late Mr. Geo. West of University College, Dundee, in the second decade of this century [1920s].' Additional illustrations: Sims (1996: 350-351, pi. 167, fig. 1, 3 images). N. vatinata J.R. Carter & Denny 1982: 315, pi. 5, fig. 180. Holotype: BM18U2. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from rot- ting leaves.' N. vincita J.R. Carter & Denny 1982: 315, pi. 5, fig. 182. Holotype: BM 78107. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. virosa J.R. Carter & Denny 1992: 188, pi. 4, fig. 43. Holotype: BM 81506. Locality: '...in sample 3... [Lake Sonfon, Sierra Leone]'. N. visitanda J.R. Carter 19706: 621, pi. 2, figs 15-16. Holotype: BM 77780. Locality: 'Sample No. 2 from a ditch near Llorts which is in the only side valley of the republic... collected by Mr. J. Broadhead during the summer of 1966 and 1967.' N. vitila J.R. Carter & Denny 1982: 316, pi. 5, figs 176-77. Holotype: BM 78108. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. vula J.R. Carter 1971a: 669, pi. 2, fig. 84. Holotype: 'Typus: praeparatum No. 27 1 6C in collectione Carter' = BM 93709. Locality: 'Devil's Hole Cave' '...Kincraig Hill at the south- western extremity of Kilconquhar parish, Fife. . .collected by the late Mr. Geo. West of University College, Dundee, in the second decade of this century [1920s].' Additional illustrations: Sims (1996: 350-351, pi. 167, fig. 9, 2 images). N. wavella J.R. Carter & Denny 1982: 316, pi. 5, fig. 178. Holotype: BM 18101 . Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N. wustula J.R. Carter & Denny 1982: 318, pi. 5, fig. 179. Holotype: BM 18112. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... from rot- ting leaves.' N. zipila J.R. Carter & Denny 1982: 318, pi. 5, fig. 181. Locality: 'River Jong (Taia) at Njala [Sierra Leone] . . .from stones.' Neidium Pfitzer 1871 N.juba J.R. Carter in J.R. Carter & Bailey-Watts 1981: 592, pi. 24, fig. 35. Holotype: 'collection Carter, Shet. No. 39a' = BM 87950. Locality: 'In Burkwell, Shetland.' Additional illustrations: Krammer & Lange-Bertalot ( 1 986: 269, taf. 99, fig. 11), 'Holotypus, Coll. Carter Shetland 39 A'; Sims (1996: 372-373, pi. 178, fig. 5, 1 image). N. lentis J.R. Carter & Denny 1987: 254, pi. 5, fig. 3. Locality: ' . . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)...Phytoplankton horizontal haul... Lake Mabesi.' N. miopsa J.R. Carter & Denny 1992: 189, pi. 6, fig. 26. Locality: 'In sample 9 rarely [Lake Sonfon, Sierra Leone].' N. plexum J.R. Carter & Denny 1987: 254, pi. 5, fig. 6. Holotype: BM 8U56. Locality: '... coastal strip of South- Western Sierra Leone . . . Sample 66 (3rd April 1976)...Phytoplankton horizontal haul... Lake Mabesi.' N. popeiensis J.R. Carter & Denny 1992: 189, pi. 6, fig. 27. Holotype: BM 81522. Locality: 'Not rare in sample 54... [Lake Popei, Sierra Leone].' N. tabletta J.R. Carter & Denny 1992: 190, pi. 6, figs 29-30. Holotype: BM&1522. Locality: 'Not rarely in sample 54. . . [Lake Popei, Sierra Leone].' N. taiaensis J.R. Carter & Denny 1982: 319, pi. 6, fig. 219. Holotype: 'BM 78 1 1 ' (sic) = BM 78 1 1 1 . Locality: 'River Jong (Taia) at Njala [Sierra Leone] . . .from stones.' N. testa J.R. Carter in J.R. Carter & Bailey-Watts 198 1 : 593, pi. 24, fig. 36. Holotype: 'collection Carter, Shet. No. 66' = BM 87973. Locality: 'At Housetter, Shetland.' Additional illustrations: Krammer & Lange-Bertalot ( 1 986: 268, taf. 99, fig. 10), 'Holotypus, Coll. Carter Shetland W66'; Sims (1996: 374-537, pi. 179, fig. 3, 1 image). Nitzschia Hassall 1845 N. disputata J.R. Carter 197 la: 670, pi. 1, figs 38^0. Syntypes: BM 77777-8. Locality: 'Devil's Hole Cave' '...Kincraig Hill at the south- 148 western extremity of Kilconquhar parish, Fife. . .collected by the late Mr. Geo. West of University College, Dundee, in the second decade of this century [1920s].' Additional illustrations: Krammer & Lange-Bertalot (1988: 51, taf. 40, figs 9-15), not type material, as N. epithemoides var. disputata (J.R. Carter) Lange-Bert. in Lange-Bertalot & Krammer (1987: 16); Sims (1996: 384-385, pi. 184, fig. 1, 3 images). N. eppressa J.R. Carter 1966: 467, pi. 6, figs 24-25. Holotype: BM 77590. Locality*: 'Tristan da Cunha inventa aqua dulci... Sample No. 5. Big Waterburn, stones in stream, 100 feet above sea level.' N. inimasta J.R. Carter & Denny 1982: 320, pi. 7, figs 230-230a. Holotype: BM78107. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' N.jamiesoni H.G. Barber & J.R. Carter 1972: 147, fig. 207. Holotype?: Typus in collectione Quekett Microscopical Club.' Locality: '...fossil diatomaceous earth situated in Gordon Road, Auckland, New Zealand at co-ordinates 225,110E, 656,720N, Square G.7 of New Zealand M.S. 17, 1962 Edition 'Auckland'.' N. navia J.R. Carter 1966: 468, pi. 7, figs 1, 7. Holotype: BM 77593. Locality: 'Tristan da Cunha inventa aqua dulci... Sample No. 8. Salt beach at sea level.' [BM 77593]. N. oscilla J.R. Carter 1966: 468, pi. 7, figs 21-22. Holotype: BM 77589. Locality: Tristan da Cunha inventa aqua dulci... Sample No. 4. Rookery Gulch, wet rock, with gelatinous algae (Nostoc sp.).' N. palmida J.R. Carter 1966: 468, pi. 9, figs 27-28. Holotype: BM 77600. Locality: 'Gough Island inventa aqua dulci... Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' N. petulla J.R. Carter 1966: 469, pi. 6, figs 18-23. Holotype: BM 77593. Locality: Tristan da Cunha inventa aqua dulci... Sample No. 8. Salt beach at sea level.' N. pistilla J.R. Carter 1966: 469, pi. 7, fig. 5. Holotype: BM 77590. Locality: Tristan da Cunha inventa in aqua dulci... Sample No. 5. Big Waterburn, stones in stream, 100 feet above sea level.' N. rotula J.R. Carter 1966: 470, pi. 7, fig. 19, 26. Holotype: BM 77586. Locality: Tristan da Cunha inventa aqua dulci... Sample No. 1. Pigbite, stones in running water at sea level.' N. salta J.R. Carter 1966: 470, pi. 7, fig. 24. Holotype: BM 77595. Locality: Tristan da Cunha inventa in aqua dulci... Sample No. 10. New lava field, stream and moss.' N. serpentiformis J.R. Carter 1966: 470, pi. 9, fig. 10. Holotype: BM 77599. Locality: 'Gough Island inventa aqua dulci... Sample No. 11 Gough Island sample 1003, Cave Bell Rocks, sea level.' N. sonora J.R. Carter 1966: 470, pi. 8, figs 21-22. Holotype: BM 77600. Locality: 'Gough Island inventa in aqua dulci... Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' D.M. WILLIAMS AND G. REID N. tantata J.R. Carter & Denny 1992: 192, pi. 7, fig. 17. Holotype: BM8\5Q9. Locality: 'In sample 5 only [Lake Sonfon, stream, Sierra Leone].' N. wodensis J.R. Carter 1964: 228, pi. 2, fig. 17. Holotype?: Slides in Quekett Microscopical Society Collections. Locality: Woden Loch on the surface of dead wood, fresh water, not very common.' Notes: If the Quekett slides are not available, BM 93522 ('Woden Loch, on dead wood, 2582, 9/62') would serve as lectotype. Opephora P. Petit 1888 O. taiaensis J.R. Carter & Denny 1982: 321, pi. 5, fig. 185. Holotype: BM78111. Locality: 'River Jong (Taia) at Njala [Sierra Leone] . . .from stones.' Pinnularia Ehrenb. 1843 P. carminata H.G. Barber & J.R. Carter 1978: 305, text-figs p. 306. Holotype: BM 78064. Locality: '...inter muscos in rivulo montoso ad Tai Newddion, Nant Ffrancon, Gwynedd', Cambriam, Barber - Cleaning No. 537 lectus.' Additional illustrations: Krammer (1992: 140, taf. 55, fig. 5), 'Loch Enoch, Schottland, leg. Carter. Prap. 574B, IOK' ; Krammer (2000: 153, pi. 133, fig. 5), 'Loch Enoch, Schottland, leg. Carter, slide 574B IOK'; Sims (1996: 434-435, pi. 209, fig. 3, 3 images). P. dispersa J.R. Carter 1966: 472, pi. 5, figs 7-9. Holotype: BM 77593. Locality: Tristan da Cuhna inventa in aqua dulci... Sample No. 8. Salt beach at sea level.' Note: non P. dispersa 0strup (1901: 273, fig. 37), see P. john- carterii D.M. Williams, nov. nom. P. extenda J.R. Carter 1966: 473, pi. 4, figs 3-4. Holotype: BM 77590. Locality: Tristan da Cuhna inventa in aqua dulci . . .Sample No. 5. Big Waterburn, stones in stream, 100 feet above sea level.' P. graphica J.R. Carter & Denny 1992: 194, pi. 8, fig. 5. Holotype: BM 81509. Locality: 'In sample 5 only, not common [Lake Sonfon, stream, Sierra Leone].' P. impassa J.R. Carter 1966: 473, pi. 4, figs 5-6. Holotype: BM 77590. Locality: Tristan da Cuhna inventa in aqua dulci . . .Sample No. 5. Big Waterburn, stones in stream, 100 feet above sea level.' P. johncarterii D.M. Williams, nov. nom. = P. dispersa J.R. Carter (1966: 472, pi. 5, fig. 7-9) non P. dispersa 0strup (1901 : 273, fig. 37). P. leptosomiformis J.R. Carter 1966: 474, pi. 8, fig. 6. Holotype: BM 77600. Locality: 'Gough Island inventa in aqua dulci... Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' P. mica J.R. Carter & Denny 1987: 258, pi. 5, figs 19-22. Holotype: 'BM91146' (sic) = BM 81146. Locality: ' . . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)... )...Periplankton scraped from Vossia stems... Lake Mape.' P. mondimia J.R. Carter & Denny 1992: 198, pi. 8, fig. 6. Holotype: BM 81522. Locality: 'In sample 54... [Lake Popei, Sierra Leone].' P. posita J.R. Carter 1966: 475, pi. 8, figs 7-8. CARTER DIATOM TYPE SLIDES 149 Holotype: BM 77599. Locality: 'Gough Island inventa in aqua dulci... Sample No. 11 Gough Island sample 1003, Cave Bell Rocks, sea level.' P. redunda J.R. Carter 1966: 475, pi. 8, figs 10-11. Holotype: BM 77600. Locality: 'Gough Island inventa in aqua dulci... Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' P. restituta J.R. Carter 1966: 475, pi. 8, figs 3-5. Holotype: BM 77599. Locality: 'Gough Island inventa in aqua dulci... Sample No. 11 Gough Island sample 1003, Cave Bell Rocks, sea level.' P. ritarda J.R. Carter 1966: 476, pi. 4, fig. 21. Holotype: 'in collectione Carter, Hawick, Scotland.' = BM 77590. Locality: 'Tristan da Cuhna inventa in aqua dulci... Sample No. 5. Big Waterburn, stones in stream, 100 feet above sea level.' P. seriata J.R. Carter 1966: 476, pi. 8, fig. 12. Holotype: BM 77600. Locality: 'Gough Island inventa in aqua dulci... Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' P. sistassa J.R. Carter 1966: 477, pi. 5, figs 3-6. Syntypes: BM 77586-7. Locality: 'Sample no. 1. Pigbite, stones in running water at sea level.' [BM 77586] 'Tristan da Cuhna inventa in aqua dulci... Sample No. 2. Above Burntwood, stream at 1800 feet above sea level.' [BM 77587]. Additional illustrations: Sims (1996: 474-475, pi. 229, fig. 6, 7 images). R statua J.R. Carter & Denny 1987: 260, pi. 8, figs 2-3, 10. Holotype: BM 8U54. Locality: '. . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)...Phytoplankton horizontal haul... Lake Mabesi.' P. subcapitata f.plana J.R. Carter 1966: 478, pi. 8, figs 2-3. Holotype: Not given. Lectotype: BM 77599, 'Sample 11 not rare.' Locality: 'Tristan da Cuhna inventa in aqua dulci... Sample No. 1 1 Gough Island sample 1003, Cave Bell Rocks, sea level.' R suplica J.R. Carter & Denny 1987: 262, pi. 7, fig. 8. Holotype: BM 81154. Locality: ' . . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)...Phytoplankton horizontal haul... Lake Mabesi.' R taiaensis J.R. Carter & Denny 1982: 324, pi. 8, fig. 265. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... surface Flotsam.' R trudis J.R. Carter & Denny 1992: 199, pi. 9, fig. 11. Holotype: BM 81508. Locality: 'In sample 4... [Lake Sonfon, Sierra Leone].' R trulla J.R. Carter & Denny 1987: 262, pi. 7, fig. 15. //0/ofy/>6>:BM81154. Locality: '. . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)...Phytoplankton horizontal haul... Lake Mabesi.' R unca J.R. Carter & Denny 1987: 263, pi. 8, fig. 14. Holotype: BM 81160. Locality: '. . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)...Phytoplankton horizontal haul from near Manobotma...Lake Mape.' P. vacua J.R. Carter & Denny 1992: 196, pi. 9, fig. 7. Holotype: BM 81521. Locality: 'In sample 54 only [Lake Popei, Sierra Leone].' R vapilla J.R. Carter 1966: 478, pi. 4, figs 1-2. Syntypes: BM 77593, BM 77595. Locality: 'Tristan da Cuhna inventa in aqua dulci... Sample No. 8. Salt beach at sea level.' [BM 77593] 'Sample No. 10. New lava field, stream and moss.' [BM 77595]. Pseudoeunotia Grunow in Van Heurck 1881 R duplex J.R. Carter 1966: 479, pi. 9, fig. 29. Holotype: BM 77600. Locality: 'Gough Island inventa in aqua dulci... Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' R linearis J.R. Carter 1966: 479, pi. 9, figs 15-19. Holotype: BM 77600. Locality: 'Gough Island inventa in aqua dulci... Sample No. 12. Gough Island sample 1006, Stream Gonydale, 1400 feet a.s.l.' Stauroneis Ehrenb. 1843 S. bulla J.R. Carter & Denny 1987: 264, pi. 9, figs 16-20, pi. 1 1, figs 14-15. //o/0fy/*>:BM81160. Locality: ' . . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)...Phytoplankton horizontal haul from near Manobotma...Lake Mape.' S. capula J.R. Carter & Denny 1992: 200, pi. 10, fig. 10. Holotype: BM. 81511. Locality: 'A few cells in sample 7 [Lake Sonfon, Sierra Leone].' 5. didacta J.R. Carter & Denny 1992: 200, pi. 10, fig. 9. Holotype: BM 81506. Locality: 'Occasional in sample 3, rarely in sample 54... [Lake Sonfon, Sierra Leone].' 5. impenda J.R. Carter & Denny 1992: 202, pi. 10, fig. 8. Holotype: BM 81509. Locality: 'In samples 5 [Lake Sonfon, Sierra Leone] and 54 [Lake Popei, Sierra Leone]....' Sample 5 is holotype material, sample 54 represents syntype material. S. singula J.R. Carter & Denny 1992: 202, pi. 10, fig. 5. Holotype: BM 81522. Locality: 'Not rarely in sample 54. . . [Lake Popei, Sierra Leone].' Surirella Turpin 1828 S. sutilis J.R. Carter & Denny 1992: 204, pi. 1 1, fig. 6. Locality: 'In sample 54... [Lake Popei, Sierra Leone].' S. taiaensis J.R. Carter & Denny 1982: 325, pi. 8, fig. 274. Holotype: BM 18108. Locality: 'River Jong (Taia) at Njala [Sierra Leone]... phyto- plankton.' S. terebra J.R. Carter & Denny 1987: 268, pi. 9, fig. 30. Holotype: BM81154. Locality: '. . .coastal strip of South- Western Sierra Leone. . .Sample 66 (3rd April 1976)...Phytoplankton horizontal haul... Lake Mabesi.' 150 Synedra Ehrenb. 1830 S. capilla J.R. Carter & Denny 1992: 204 = S.flliformis J.R. Carter & Denny non S. filiformis Grunow ( 1 880). S.flliformis J.R. Carter & Denny 1987: 270, pi. 9, fig. 32. Holotype: BM81159. Locality: '. . .coastal strip of South- Western Sierra Leone. . .Sample 68 (16th April 1976) ... Phytoplankton horizontal haul... Lake Mape.' BIBLIOGRAPHY 1. Burke, F. & Carter, J.R. 1943. A diatom new to Britain. Nature 151: 672-673. 2. Carter, J.R. 1947. Diatom notes. British fresh-water forms of the genus Gomphonema. The Microscope 6: 170-176. 3. Carter, J.R. 1960a. Diatom notes. British freshwater forms of the genus Gomphonema. The Microscope 12: 255-264. 4. Carter, J.R. 1960ft. Diatom notes. Some minute forms of the genus Navicula. The Microscope 12: 283-289. 5. Carter,J.R. 1961a. Diatom notes. The genus Achnanthes as it occurs in British fresh waters. The Microscope 12: 320-326. 6. Carter, J.R. 1 96 1 b. Diatom notes. The genus Achnanthes as it occurs in British fresh waters (continued). The Microscope 13:15-22. 7. Carter, J.R. 1 96 1 c. Diatom notes. The genus Achnanthes as it occurs in British fresh waters (continued). The Microscope 13: 37-45. 8. Carter, J.R. 196 Id. Diatom notes. Loch na Cuilc, a preliminary notice. The Microscope 13: 74-78. 9. Carter, J.R. 1962. Diatom notes. Some unusual British forms. The Microscope 13: 156-162. 10. Carter, J.R. 1963 ['1962']. Diatom notes. More unusual British forms. The Microscope 13: 231-237. 11. Carter, J.R. 1963. Some new diatoms from British waters. Journal of the Quekett Microscopical Club 29: 199-203. 12. Carter, J.R. 1964. Some new diatoms from British waters - Part II. Journal of the Quekett Microscopical Club 29: 225-228. 13. Carter, J.R. 1966. Some fresh water diatoms of Tristan da Cunha and Gough Island. Nova Hedwigia 11: 443-483. 14. Barber, H.G. & Carter, J.R. 1970. An account of fossil fresh water diatomaceous earth from Gordon Road Site, Auckland, New Zealand. Microscopy 31: 271-277. 15. Carter, J.R. 1970a. Observations of some British forms of Achnanthes saxonica Krasske. Microscopy 31: 313-316. 1 6. Carter, J.R. 1 970ft. Diatoms from Andorra. Nova Hedwigia, Beiheft 31 : 605-632. 17. Barber, H.G. & Carter, J.R. 197 la. An account of fossil fresh water diatomaceous earth from Gordon Road site, Auckland, New Zealand. Part 2. Microscopy 32: 24-28. 18. Barber, H.G. & Carter, J.R. 1971ft. An account of fossil fresh water diatomaceous earth from Gordon Road site, Auckland, New Zealand. Part 3. Microscopy 32: 82-89. 19. Carter, J.R. 197 la. Diatoms from the Devil's Hole Cave, Fife, Scotland. Nova Hedwigia 21: 657-68 1 . 20. Carter, J.R. 1971ft. Diatoms, the forgotten family. The Vasculum 56: 66-69. 21. Barber, H.G. & Carter, J.R. 1972. An account of fossil fresh water diatomaceous earth from Gordon Road site, Auckland, New Zealand. (Conclusion). Microscopy 32: 141-147. 22. Carter, J.R. 1972a. Some observations on the diatom Pinnularia acoricola Hustedt. Microscopy 32: 162-165. 23. Carter, J.R. 1972ft. Diatoms of Slapestone Sike, Upper Teesdale. The Vasculum 57: 35^41. 24. Barber, H.G. & Carter, J.R. 1978. Pinnularia carminata N. Sp. Microscopy 33: 305-307. 25. Carter, J.R. 1979. On the identity of Navicula cincta Ehrenberg. Bacillaria 2: 73-84. 26. Carter J.R. & Bailey- Watts, A.E. 1 98 1 . A taxonomic study of diatoms D.M. WILLIAMS AND G. REID from standing freshwaters in Shetland. Nova Hedwigia 33: 513-629. 27. Barber, H.G. & Carter, J.R. 1981 . Observations on some deformities found in British diatoms. Microscopy 34: 214-226. 28. Moss, M.O. & Carter, J.R. 1982. The resurrection of Achnanthes rostrata 0strup. Bacillaria 5: 157-164. 29. Carter, J.R. & Denny, P. 1982. Freshwater algae of Sierra Leone. HI. [sic, part IV] Bacillariophycaeae: Part (i) diatoms from the River Jong (Taia) at Njala. Nova Hedwigia, Beiheft 73: 281-331. 30. Carter, J.R. 1983. Diatoms from Tuffa at Hassendean Burn, Roxburghshire. The Vasculum 68: 47^48. 31. Carter, J.R. 1984. A new diatom, Navicula belsesiensis. Microscopy 35: 134-136. 32. Carter, J.R. 1986. Some little-known diatoms newly recorded in Brit- ain. Microscopy 35: 356-357, 389. 33. Preece R.C., Bennet K.D. & Carter, J.R. 1986. The Quaternary paleobotany of Inaccessible Island (Tristan da Cunha group). Journal of Biogeography 13: 1-33. 34. Carter, J.R. 1987. Navicula claytoni, a new diatom. Microscopy 35: 633-635. 35. Carter, J.R. & Denny, P. 1987. Freshwater algae of Sierra Leone. IV Bacillariophyceae: Part (ii) diatoms of the coastal region of the southern province. Nova Hedwigia 44: 229-275. 36. Carter, J.R. & Flower, R.J. 1 988. A new species ofEunotia, E. pirla sp. nov., from Woolmer Pond, an acid pool in the southeast of England. Diatom Research 3: 1-8. 37. Carter, J.R. 1 989. An unusual diatom community from ironstone waste heaps in Cleveland. Microscopy 36: 216-220. 38. Denys, L. & Carter, J.R. 1989. The diatom Navicula genustriata Hustedt: valve morphology, variability and notes on its ecology. Diatom Research 4: 9-19. 39. Carter, J.R. 1 990. A new Eunotia and its great morphological variations under stress caused by a habitat loaded with copper salts. In M. Ricard (Ed.), Ouvrage dedie a la Memoire du Professeur Henry Germain (1903-1989): 13-17. Koenigstein. 40. Hakansson, H. & Carter, J.R. 1990. An interpretation of HUSTEDT's terms 'Schattenlinie', 'Perlenreihe' and 'Hocker' using specimens of the Cyclotella rad/ora-complex, C. distinguenda HUST., and C. cyclopuncta nov. sp. Journal of Iowa Academy of Science 97: 153-156. 41. Carter, J.R. & Denny, P. 1992. Freshwater algae of Sierra Leone. IV. Bacillariophyceae: Part (iii) diatoms from the Lake Sonfon region and from Lake Popei. Nova Hedwigia 54: 159-21 1. 42. Carter, J.R. 1993a. A note on the genus Berkella Ross & Sims. Nova Hedwigia Beiheft 106: 101-107. 43. Carter, J.R. 1993ft. Observations on some newly described diatoms in the genus Cocconeis Ehrenberg now found to be members of the British flora. Quekett Journal of Microscopy 37: 39-41 . 44. Carter, J.R. & Round, F.E. 1993. Studies on freshwater Amphora species. V. A. montana and A. normanii. Diatom Research 8: 1-11. 45. Sims, P. (Ed.) 1996. An atlas of British diatoms based on illustrations by H.G. Barber and J.R. Carter; arranged by B. Hartley. Bristol. ACKNOWLEDGEMENTS. We would like to thank Karen Webb, whose initial investigations into John Carter's diatom collections instigated this work, and Dr E.Y. Haworth for information. REFERENCES Bailey, J.W. 1 844. Account of some new infusorial forms discovered in the fossil Infusoria from Petersburg, Va., and Piscataway, Md. American Journal of Science 46: 137-141. Bukhtiyarova, L. & Round, F.E. 1996. Revision of the genus Achnanthes sensu lato section Marginulatae Bukht. Sect. Nov. of Achnanthidium Kiitz. Diatom Research 11: 1-30. Castracane, A. 1886. Report on the scientific results of the voyage of H. M.S. Chal- lenger during the years 1873-1876; Botany, 2. London. Cholnoky, BJ. 1963. Beitrage zur Kenntnis des marinen Literals von Sudafrika. Botanica Marina 5: 38-83. CARTER DIATOM TYPE SLIDES 151 Cleve, P.T. & Grunow, A. 1880. Beitrage zur Kenntnis der artischen Diatomeen. Kongliga Svenska Vetenskaps-Akademiens Handlingar 17: 1-121. Flower, R.J. & Jones, V.J. 1989. Taxonomic descriptions and ocurrences of new Achnanthes taxa in acid lakes in the U.K. Diatom Research 4: 227-239. Fusey, P. 1950. Contribution a flore algologique du Jura. II. Diatome'es des R6gions de Montbenoit et du Russey (Doubs). Microscopic 2: 149-164. Greville, R.K. 1866. Description of new and rare diatoms from the tropics and Southern Hemisphere. Transactions of the Botanical Society of Edinburgh 8: 436- 441. Grove, E. & Sturt, G. 1887. On a fossil marine Diatomaceous deposit from Oamaru, Otago, New Zealand. Journal of the Quekett Microscopical Club 3: 63-78. Gutwinski, R. 1890. Materyjaly do flory glon6w (wodrostdw) Galicyi. Sprawozdanie Komisiji fizyograpficznej 25: 1-43. Hartley, B. 1986. A check-list of the freshwater, brackish and marine diatoms of the British Isles and adjoining coastal waters. Journal of the Marine Biological Associa- tion UK 66: 531-610. Hartley, B. 1994. John Ripley Carter (1908-1993). Diatom Research 9: 235-236. Haworth, E.Y. 1974. Some problems of diatom taxonomy in Scottish Lake sediments. British Phycological Journal 9: 47-55. & Offer, E.H. 1986. An amended description of the diatom Navicula menda Carter. British Phycological Journal 21: 445^447. Hustedt, F. 1936. In A. Schmidt et al. (Eds), Atlas der Diatomaceenkunde. Leipzig. 1942. Diatomeen aus der Umgebung von Abisko in schwedisch Lappland. Archiv fur Hydrobiologie 39: 87-174. 1945. Diatomeen aus Seen und Quellgebieten der Balkanhalbinsel. Archiv fur Hydrobiologie 40: 867-973. Krammer, K. 1992. Pinnularia eine Monographic der europaischen Taxa. Bibliotheca Diatomologica 26, 353pp. 1997a. Die cymbelloiden Diatomeen: eine Monographic der weltweit bekannten Taxa. Teil 1: Allgemeines und Encyonema. Jena. 1997i>. Die cymbelloiden Diatomeen: eine Monographic der weltweit bekannten Taxa. Teil 2: Encyonema, Encyonopsis and Cymbellopsis. Jena. 2000. Diatoms of Europe: diatoms of the European inland waters and compar- able habitats. 1. The genus Pinnularia. Ruggell. & Lange-Bertalot, H. 1985. Naviculaceae: neue und wenig bekannte Taxa, neue Kombinationen und Synonyme sowie Bemerkungen zu einigen Gattungen. Jena. 1986. Susswasserflora von Mitteleuropa, 2(1), Bacillariophyceae. Teil: Naviculaceae. Stuttgart. 1988. Susswasserflora von Mitteleuropa, 2(2), Bacillariophyceae. Teil Bacillariaceae, Epithemiaceae, Surirellaceae. Jena. 1991a. Susswasserflora von Mitteleuropa, 2(3), Bacillariophyceae. Teil: Centrales, Fragilariaceae, Eunotiaceae. Jena. 19916. Susswasserflora von Mitteleuropa, 2(4), Bacillariophyceae. Teil: Achnan- thaceae, Kritische Erganzungen zu Navicula (Lineolatae) und Gomphonema Gesamtliteraturverzeichnis. Jena. Lange-Bertalot, H. 1993. 85 Neue Taxa und iiber 100 weitere neu defmiterte Taxa erganzend zur SiiBwasserflora von Mitteleuropa. Vol. 2/1-4. Bibliotheca Diatomologica 27, 428pp. 1997. Zur revision der Gattung Achnanthes sensu lato (Bacillariophyceae): Achnantheiopsis, eine neue Gattung mit dem Typus generis A. lanceolata. Archiv fur Protistenkunde 148: 199-208. 1999. Neue Kombinationen von Taxa aus Achnanthes Bory (sensu lato). Iconographia Diatomologica 6: 270-283. 200! . Diatoms of Europe: diatoms of the European inland waters and compar- able habitats. 2. Navicula sensu stricto 10 genera separated from Navicula sensu lato. Ruggell. & Krammer, K. 1987. Bacillariaceae, Epithemiaceae, Surirellaceae. Neue und wenig bekannte Taxa, neue Kombinationen und Synonyme sowie Bemerkungen und Erganzungen zu den Naviculaceae. Bibliotheca Diatomologica 15, 289pp. 1989. Achnanthes: eine Monographic der Gattung mit Definition der Gattung Cocconeis und Nachtragen zu den Naviculaceae. Berlin. Metzeltin, D. & Witkowski, A. 1996. Hippodonta gen. nov. Umschreibung und Begriindung einer neuen Gattung der Naviculaceae. Iconographia Diatomologica 4: 247-274. & Moser, G. 1994. Brachysira. Monographic der Gattung. Bibliotheca Diatomologica 29, 2 1 2pp. Manguin, E. 1942. Contribution a connaissance des Diatome'es d'eau douce des A?ores. Travaux Algologiques 1: 115-160. Marson, J.E. 1 994. John Ripley Carter 1 908- 1 993. The Quekett Journal of Microscopy 37: 256-257. Ostrup, E. 1901. Freshwater diatoms from the Faeroes. In J.E.B. Warming (Ed.), Botany of the Faeroes 1: 260-290. Pritchard, A. 1 86 1 . A history of the Infusoria, living and fossil arranged according to 'Die Infusionsthierchen' of Ehrenberg. 4'h ed. London. Ross, R. & Sims, P.A. 1978. Notes on some diatoms from the Isle of Mull, and other Scottish localities. Bacillaria 1: 151-168. Round, F.E., Crawford, R.M. & Mann, D.G. 1990. The diatoms. Cambridge. Sims, P. (Ed.) 1996. An atlas of British diatoms based on illustrations by H.G. Barber and J.R. Carter: arranged by B. Hartley. Bristol. VanLandingham, S.L. 1975. Catalogue of the fossil and recent genera and species of diatoms and their synonyms. Part. V. Navicula. Vaduz. Zimmermann, S.S. 1915. Algunas Diatomaceas novas ou curiosas. Broteria 13 (Bot.): 33-36. Bull. nat. Hist. Mus. Land. (Bot.) 32(2): 153-156 Issued 28 November 2002 The typification, defining features and geographical range of Calymperes couguiense Besch. (Calymperaceae, Musci) LEN T. ELLIS Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD SYNOPSIS. A neotype is proposed for Calymperes couguiense Besch. and the features are outlined by which the species may be distinguished from C. subintegrum Broth, and C. porrectum Mitt. Owing to its confusion with these latter species, most reports of its occurrence in Malesia are erroneous. The distribution of C. couguiense includes NE Australia and the islands of the Pacific as far east as the Tuamotu Archipelago. Research for this paper was inspired by the discovery that, contrary to citations in the literature, the holotype of Calymperes couguiense Besch. is not in Bescherelle's herbarium (BM). An examination of many specimens identified as C. couguiense revealed a long-stand- ing confusion of this species with some forms of Calymperes porrectum Mitt, and Calymperes subintegrum Broth. These prob- lems of typification and identification are addressed below. Calymperes couguiense Besch. in Ann. Sci. Nat. Bot. ser. 5, 18: 206 (1873). Type: New Caledonia, Balade, Vieillard 1776 (BM!- neotype, designated here; S!-isotype?). Calymperes setosum Mull. Hal. in J. Mus. Godeffroy 3(6): 64 (1874). Type: Samoa, Upolu, Graeffe s.n. (BM!, S!-isotypes). Calymperes marginatum Dixon in Proc. Linn. Soc. New South Wales 55: 275 (1930). Type: Fiji, VanuaLevu, 3000 ft, December 1922, Greenwood 474 (BM!-holotype). Calymperes pseudopo dianum E.B. Bartram in Occas. Pap. Bernice Pauahi Bishop Mus. 10(10): 7 (1933). Type: Pacific Islands, Tuamotu Archipelago, Makatea, 3 March 1930, Emory s.n. (FH- holotype). Typification There has been some misapprehension with regard to the typifica- tion of Calymperes couguiense Besch. In the protologue, Bescherelle ( 1 873) cited a type specimen from New Caledonia - 'Ad cortices in monte Cougui (B ALANS A)'. Reese & Mohamed (1985) and Reese & Stone (1995) cited this specimen as being housed in BM. How- ever, the holotype of Calymperes couguiense (Balansa s.n.) is not in Bescherelle's herbarium (BM) and isotypes have not been found in BM or in other likely herbaria elsewhere (FH, H, L, MEL, MO, NY, PC, S). Hitherto, a non-type collection in BM, identified as C. couguiense by Bescherelle (1895), appears to have been mistaken for the type specimen. Although annotated as type material by recent authorities, it is clearly labelled 'Nov. Caled. Balade Vieillard No. 1776'. In Moller's herbarium (S) there is an apparent duplicate of Vieillard 1776 which has also been mistaken for type material. The outer packet of this specimen is marked 'Vieillard 1776' and stamped TYPUS'. Confusingly, the inner packet bears a label with the collection details of Vieillard 1776 followed immediately by those of Balansa s.n. This is possibly a rendition of the specimen citation published by Bescherelle (1895) rather than a specific reference to the contents of the packet. The true origin of this specimen remains uncertain. In the absence of material clearly identifiable as Balansa's collec- tion, Calymperes couguiense requires neotypification. The authen- tic material, Vieillard 1776 in Bescherelle's herbarium (BM), is here proposed as the neotype. Taxonomy Calymperes subintegrum Broth., C. porrectum Mitt, and C. couguiense Besch. all possess similar strongly dimorphic leaves (i.e. erect, linear gemmiferous leaves, largely consisting of thick costae (Fig. Ib) which project above a canopy of vegetative leaves with narrower costae and relatively broad laminal blades (Fig. la)). Collections of Calymperes subintegrum and of diminutive forms of Calymperes porrectum have often been misidentified as C. couguiense. This has led to a misrepresentation of the latter species both in herbaria and in recent literature. For example, the illustra- tions of C. couguiense by Eddy ( 1 990) mostly depict leaf features of C. porrectum, and Reese, Koponen & Norris (1986) cited only misidentified material in their account of the species. The distinguishing features of Calymperes couguiense, C. porrectum and C. subintegrum are outlined in a key and discussed in detail below. Key features of Calymperes porrectum Mitt., C. subintegrum Broth, and C. couguiense Besch. 1 Non-gemmiferous leaves recurved, costa smooth, cells of chlorophyllose lamina smooth and flat; leaf margin with intramarginal polystratose rib, serrate (Fig. Ir, t) Calymperes porrectum Non-gemmiferous leaves erect to spreading, costa scabrid to some degree, cells of chlorophyllose lamina projecting from the ventral and/or dorsal surfaces of the leaf; leaf margin with or without rib, subentire to denticulate 2 2 Cells of chlorophyllose lamina 7.5-15(->20) x 5-10(-12.5) \un, dorsally flat (occasionally unipapillose) (Fig. 1m, o) Calymperes subintegrum Cells of chlorophyllose lamina <5-10(-12.5) x 5-7.5(-10) urn, dorsally unipapillose (invariably) (Fig. Id, i) Calymperes couguiense For an illustrated account of Calymperes porrectum Mitt, with synonymy see Eddy (1990), and see Ellis (1991) for a similar treatment of Calymperes subintegrum Broth. Recent useful ac- counts of Calymperes couguiense Besch. include those by Reese & Mohamed (1985), Reese, Koponen & Norris (1986) and Reese & Stone (1995). © The Natural History Museum, 2002 154 L.T. ELLIS a, b 0. 5 mm 200 Lim d-k, m-t 50 jum Fig. 1 a-k. Calymperes couguiense Besch. a-c. leaves (a: non-gemmiferous leaf, b: gemmiferous leaf (gemmae missing), c: apex of gemmiferous leaf in lateral view); d-g. cells of leaf in surface view (d: chlorophyllose lamina (ventral view), e, f: margin of chlorophyllose limb (ventral view), g: margin at mid hyaline base), h-k. cross-sections of chlorophyllose limb (h: costa, i: lamina, j, k: margin). 1-p. Calymperes subintegrum Broth. 1. apex of gemmiferous leaf in lateral view, m, n. cells of chlorophyllose limb in surface view (m: lamina (ventral view), n: margin), o, p. cross-sections of chlorophyllose limb (o: lamina, p: margin), q-t. Calymperes porrectum Mitt, q, r. chlorophyllose limb in surface view (q: lamina, r: margin), s, t. cross- sections of chlorophyllose limb (s: costa, t: lamina and margin), a, c. Drawn from Society Islands, De Sloover 20962 (H). b, e, g-j. Drawn from New Caledonia, Vieillard 1776 (BM). d. Drawn from Australia, Stone 18577 (MEL), f, k. Drawn from Samoa, Graeffe s.n. (BM). 1, m, o. Drawn from Australia, Stone 23095 (MEL), n, p. Drawn from New Guinea, Zanten 149c (BM). q-t. Drawn from New Ireland, Eddy 6333 (BM). TYPICATION OF CALYMPERES COUGUIENSE 155 DISCUSSION. Shoots of Calymperes porrectum possess strongly recurved non-gemmiferous leaves with a smooth costa (Fig. Is) and chlorophyllose lamina (Fig. It). The laminal chlorocysts form a largely flat surface, but are sometimes very slightly protuberant ventrally. In surface view they reach <5-10(-12.5) x <5-10(- 12.5) jam, smaller on average, and more irregularly polygonal or rounded, than those in C. subintegrum (see below). A differenti- ated rib lies well within the margin of the leaf, and extends from the leaf base to near the apex. Above the hyaline lamina it be- comes polystratose and projects slightly from the dorsal leaf surface (Fig. It). Adjacent to this rib, a narrow, unistratose mar- ginal lamina, (2-)3-5(->6) cells wide, is formed, which at intervals gives rise to subtriangular, often multicellular, teeth (Fig. Ir). In gemmiferous leaves the costa is scabrid and slightly excurrent; gemmae are produced in a cluster from the costal apex. Adjacent to the cluster of gemmae some superficial costal cells become elongated and finger-shaped. In Calymperes subintegrum the non-gemmiferous leaves are erect to spreading and possess more or less scabrid costae. The cells of the chlorophyllose lamina are flat dorsally (occasionally unipapillose), but project acutely from the ventral surface of the leaf (Fig. lo). In surface view they reach 7.5-15(->20) x 5-10(-12.5) jam, and are mostly subrectangular (Fig. 1m). A narrow, intramarginal, unistratose (rarely bistratose) rib of linear cells is usually apparent. Adjacent to this, a single row of subrectangular cells (ventrally and dorsally flat) forms a subentire to denticulate margin (Fig. In, p). Unlike in C. porrectum and C. couguiense, the costa in the gemmiferous leaves of C. subintegrum ceases just below the leaf apex. This is formed by a narrow band of lamina, and gemmae are produced only from the ventral surface of the costal tip (Fig. 11). Adjacent to the clustered gemmae a few elongated, finger- like cells may protrude from the surface of the costa. Unfortunately, the disposition of gemmae and lamina is often obscured by damage, or in well-developed leaves, by the backward flexing of the leaf tip. In practical terms, this slightly limits the value of the apices of gemmiferous leaves as a reliable distinguishing feature. In Calymperes couguiense the non-gemmiferous leaves have a similar disposition to those in C. subintegrum and are never nota- bly recurved. They have scabrid costae (Fig. Ih), and the cells of the chlorophyllose lamina are ventrally acutely protuberant and dorsally unipapillose (Fig. li). Superficially they are visibly smaller than those of C. porrectum and C. subintegrum, reaching <5_10(-12.5) x 5-7.5(-10) (am (Fig. Id). Above the hyaline lamina, the leaf margins are often formed by a subentire to den- ticulate, polystratose rib (Fig. le, j, k). This sometimes appears to be intramarginal, particularly when seen in surface view. In the leaves of many specimens the marginal/intramarginal rib is poorly developed to absent (Fig. If)- The gemmiferous leaves possess apices similar to those of C. porrectum, with an excurrent, scabrid costa and gemmae produced in a cluster from the costal tip. The lamina narrows into the costal apex, its distal-most cells, and the distal-most superficial cells of the costa are elongated, finger-like with rounded ends; they form a regular to irregular fringe at the base of the crown-like cluster of gemmae (Fig. le). These sterile, projecting cells are a consistently well-developed feature in col- lections of C. couguiense. HABITAT. Calymperes couguiense Besch. evidently flourishes in habitats ranging from undisturbed forest to cultivated gardens, and has mostly been collected from sea level to below 500 m. Both Calymperes porrectum and Calymperes subintegrum occur within this altitudinal range, but records of these species extend to over 1000 m. Calymperes couguiense also shares some substrate preferences with C. porrectum and C. subintegrum, occurring on tree boles, twigs and dead wood. Well-developed turfs of C. couguiense have also been collected on sandy soil. DISTRIBUTION. Calymperes couguiense Besch. is largely a species of the Pacific islands and northeastern Australia. Reports of its occurrence in Java and New Guinea (Reese, Koponen & Morris, 1986; Reese & Stone, 1987, 1995) appear to be unsupported by specimens. Misidentified collections from the Malay Peninsula (Spare 3213b, BM), Java (Fleischer 70, BM, FH, NY; Schiffner 3, BM) New Ireland (Eddy 6165, BM), and New Guinea (Zanten 190a, NY) represent a diminutive form of Calymperes porrectum Mitt. A further wrongly determined collection from New Guinea (Zanten 149c, BM) is Calymperes subintegrum Broth. Of the material seen for this present study, only a single collection from New Britain (Streimann 40215, NY) indicates the presence of Calymperes couguiense in Malesia. Orban (1995) identified specimens from the Seychelles as C. couguiense and Reese & Stone (1987) cited Mauritius as a station. Unfortunately, the material on which the latter citation is presum- ably based (Balfour s.n., NY) represents a form of Calymperes hispidum Renauld & Cardot. SPECIMENS EXAMINED. Calymperes couguiense Besch. Australia. Queensland: Cook, Cairns, Crystal Cascades, 16° 55' S 145° 46' E, July 1979, Stone 15334 pro parte (MEL), 15349 pro parte (MEL); Cook, Bellenden Ker National Park, Goldsburgh Track, The Boulders, 6 km W. of Babinda, 17 July 1987, Stone & Reese 24451 (MEL); Cook, rainforest in saddle between Bartle Frere and Bellenden Ker ranges, 8 km W. of Babinda, 7 July 1987, Reese, Stone & Stone 17038 (NY); Cook, Mt Bellenden Ker, 17° 16' S 145° 51' E, 4 September 1986, Stone 24121 (MEL), 24122 (MEL), 24123 (MEL); Cook, Mossman Gorge, 16° 28' S 145° 21' E, 1 September 1986, Stone 24061 (MEL), 24045 (MEL); Cook, Mossman Gorge, May 1975, Stone 8837 pro parte (MEL), 12037 pro parte (MEL); Cook, Mossman, Rex Creek, 16° 28' S 145° 16' E, 30 June 1982, Stone 19642 pro parte (MEL); Cook, Wooroonooran National Park, Josephine Falls, 17° 25' 58" S 145° 51' 32" E, 19 August 1986, Stone 23904 (MEL); Cook, Cardstone, Tully, 17° 55' S 145° 55' E, 20 May 1985, Stone 23113 (MEL); North Kennedy, Kennedy Bay, 17° 58' S 146° 04' E, 16 May 1982, Stone 18573 pro parte (MEL), 18577 (MEL); 9 July 1982, Stone 19987 (MEL), 20000 (MEL); North Kennedy, Cardwell, Conn Creek, 18° 15' S 145° 43' E, 8 August 1980, Stone 16337 (MEL); North Kennedy, Cardwell, Stoney Creek, 18° 35' S 145° 48' E, 21 July 1979, Trovers 14876 pro parte (MEL), 20 May 1982, Stone 18666 pro parte (MEL); North Kennedy, Tully, Kooroomool Creek, 17° 52' S 145° 42' E, 25 May 1982, Stone 18803 pro parte (MEL). New Britain. Hoskins, town area, 5° 26' S 150° 25' E, 16 February 1989, Streimann 40215 (NY). Mariana Islands. Guam, limestone forest 6 miles N. of Agana, 1 3 No- vember 1945, Holland 21 (FH). New Caledonia. Balade, Vieillard 1776 (BM, S). Fiji. Viti Levu, Suva, 25 January 1923, Greenwood 477 (FH); Vanua Levu, December 1922, Greenwood 474 (BM). Samoa. Tutuila, Vatia, Bryan Jr s.n. (FH); Tau, 4 October 1939, Yuncker9243 (FH!); Tau, Judd's Crater, NW rim, 27 July 1938, Harris 413 (FH), 414 (FH); Upolu, Mataulu village, Lefaga Bay, 24 October 1953, Irwin 122 (FH). Tuamotn Archipelago. Makatea, 3 March 1930, Emory s.n. (FH). Society Islands. Tahiti: Gaugin Museum, 26 May 1988, Hegewald & Hegewald 11477 (MO), Insel Huahine, Hegewald & Hegewald 11442 (MO); Moorea, au- dessus du point de vue du Belvedere, 1 1 July 1975, De Sloover 20962 (H, NY). SPECIMENS MISIDENTIFIED AS CALYMPERES COUGUIENSE BESCH. Calymperes boulayi Besch. - Australia. Queensland: along 'Sullivan's Track', c. 6km WNW of Cardwell, 22 July 1987, Reese, Stone & Stone 17168 (MO). Calymperes hispidum Renauld & Cardot - Mauritius, Balfour s.n. (NY). Calymperes moluccense Schwagr. New Caledonia. Hot au Sud de Tile, May 1988, Martin s.n. (PC). Calymperes porrectum Mitt. - Malay Peninsula. Perak, Kampong Ijok, 156 L.T. ELLIS 25 August 1940, Spare 3213b (BM!). Java. Buitenzorg, Botanic Garden, March 1899, Fleischer 70 (BM, FH, NY), March 1894, Schiffner 3 (BM). New Ireland. East coast, Danfu Valley, 2 February 1970, Eddy 6165 pro parte (BM!). Papua New Guinea. Kau River, below Star Mountains, Woropko, Zanten 190a (NY). Calymperes strictifolium (Mitt.) G. Roth (forma) (C. franciH) - New Cal- edonia. Riviere Bleu Reserve, near bridge over Riviere Bleu, 7 September mi, Crosby 14325 (NY). Calymperes subintegrum Broth. - Australia. Queensland, Cook, Wooroonooran National Park, Josephine Falls, 17° 25' 58" S 145° 51' 32" E, 19 August 1986, Stone 23905 pro parte (MEL). Fiji. Viti Levu, hills between Navua and Suva, May 1943, Greenwood903 (BM), 976 (BM). Papua New Guinea. Ajerok, 24 May 1959, Zanten 149c (BM). ACKNOWLEDGEMENTS. I am grateful to the curators of FH, H, L, MEL, MO, NY, PC and S for the loan of specimens. REFERENCES Bescherelle E. 1873. Florule Bryologique de la Nouvelle Caledonie. Annales des Sciences Naturelles Botanique ser. 5, 18: 184-245. 1 895. Essai sur le genre Calymperes. Annales Sciences Naturelles Botanique s6r. 8, 1: 247-308. Eddy, A. 1990. A handbook of Malesian mosses. 2. London. Ellis, L.T. 1 99 1 . Calymperes schmidtii Broth, in J. Schmidt and C. subintegrum Broth. in J. Schmidt, two distinct species from Malesia. Journal of Bryology 16: 589-593. Orban, S. 1995. East African bryophytes, XV. Calymperaceae species collected in Seychelles Islands in 1993. Fragmenta Floristica et Geobotanica 40(1): 279-287. Reese, W.D., Koponen, T. & Norris, D.H. 1986. Bryophyte flora of the Huon Peninsula, Papua New Guinea. XIX. Calymperes, Syrrhopodon and Mitthyridium (Calymperaceae, Musci). Acta Botanica Fennica 133: 151-202. & Mohamed, H. 1985. A synopsis of Calymperes (Musci: Calymperaceae) in Malaysia and adjacent regions. Bryologist 88(2): 98-109. & Stone, I.G. 1987. New records of Australian Calymperaceae and keys to Australian species of Calymperes, Mitthyridium, and Syrrhopodon. Journal of Bryology 14: 487-498. 1995. The Calymperaceae of Australia. Journal of the Hattori Botanical Laboratory 78: 1-40. Bulletin of The Natural History Museum Botany Series Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. Volume 25 No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. 1995. 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(Ranunculaceae). R.M. Baldini and C.E. Jarvis. 2002. Pp. 7-1 1, 4 figs. Catalogue of the lichens of Thailand. PA. Wolseley, B. Aguirre-Hudson and P.M. McCarthy. 2002. Pp. 13-59, 1 fig. £43.40 JNTENTS 61 Studies in the genus Hypericum L. (Guttiferae) 4(2). Section 9. Hypericum sensu lato (part 2): subsection 1. Hypericum series 1. Hypericum N.K.B. Robson 125 The natural history of reproduction in Solanum and Lycianthes (Solanaceae) in a sub- tropical moist forest S.D. Smith and S. Knapp 137 The diatom type slides and bibliography of John Carter (1908-1993) D.M. Williams and G. Reid 153 The typification, defining features and geographical range of Calymperes couguiense Besch. (Calymperaceae: Musci) LT. Ellis CAMBRIDGE UNIVERSITY PRESS 0968-0446(20021 1 )32:2;1 -K Bulletin of The Natural History Museur BOTANY SERIES Vol. 32, No. 2, November 2002