ISSN 0968-0446 Bulletin of The Natural History Museum THE NATURAL HISTORY MUSEUM Botany Series THE NATURAL HISTORY MUSEUM VOLUME 31 NUMBER 2 29 NOVEMBER 2001 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: Dr R. Bateman Editor of Bulletin: Ms S.A. Henderson 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 for acceptance. 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The Natural History Museum, 2001 Botany Series ISSN 0968-0446 Vol. 31, No. 2, pp. 37-119 The Natural History Museum Cromwell Road London SW7 5BD Issued 29 November 2001 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. nut. Hist. Mus. Land. (Bot.) 31(2): 37-88 Issued 29 November 2001 Studies in the genus Hypericum L. (Guttiferae) 4(1). Sections 7. Roscyna to 9. Hypericum sensu lato (part 1) rTHENATURAL NORMAN K.B. ROBSON Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS Introduction 37 Further delimitation of Hypericum - exclusion of H. ellipticifolium H.L. Li 38 Relationship of H. elatoides R. Keller and sect. 8. Bupleuroides 40 Relationships of sects 7. Roscyna and 9. Hypericum sensu lato 41 Dismemberment of sect. 9. Hypericum 41 Key to sections, subsections and series in Part 4 43 Sect. 7. Roscyna 44 Characters and variation 45 Distribution and evolution 45 Sect. 9a. Concinna 45 Sects 9c. Sampsonia, 9d. Elodeoida and 9e. Monanthema 45 Characters and variation 45 Distribution and evolution 47 Systematic treatment 49 Sect. 3. Ascyreia Choisy - an additional species 49 Sect. 8. Bupleuroides Stef 49 Sect. 7. Roscyna (Spach) R. Keller 52 Sect. 9a. Concinna N. Robson 61 Sect. 9c. Sampsonia N. Robson 63 Sect. 9d. Elodeoida N. Robson 66 Sect. 9e. Monanthema N. Robson 75 References 85 Systematic index 87 SYNOPSIS. Following the removal of Hypericum ellipticifolium H.L. Li to a new genus, Lianthus N. Robson gen. nov., which is described, the variation, distribution and evolution of Hypericum sects 7-9 (all herbaceous) are discussed. It is shown that, whereas sect. 7. Roscyna is indeed derived from sect. 3 Ascyreia (as was indicated in Parts 1-3), sect. 8. Bupleuroides is not directly related to sect. 7 but to the Chinese suffrutex H. elatoides R. Keller, which, in turn, is shown to belong to sect. Ascyreia, not sect. Roscyna. Sect. 9. Hypericum is considered to be a complex of six sections each derived directly from sect. Roscyna, of which one (sect. 9a. Concinna N. Robson) has already been described. The others are: sect. 9 Hypericum (which is divided into two subsections: 9.1 Hypericum and 9.2 Erecta N. Robson, subsect. nov., with subsect. Hypericum further divided into two series, Hypericum and Senanensia N. Robson, series nov.), 9b. Graveolentia N. Robson, sect, nov., 9c. Sampsonia N. Robson, sect, nov., 9d. Elodeoida N. Robson, sect. nov. and 9e. Monanthema, sect. nov. A systematic treatment is then provided of H. elatoides and sects Roscyna, Bupleuroides, Concinna, Sampsonia, Elodeoida and Monanthema. INTRODUCTION Part 4 of the Hypericum monograph includes treatments of those sections derived directly or indirectly from sect. 3. Ascyreia in the affinity of the Nepali endemic H. podocarpoides N. Robson (see Part 3, Robson, 1985: 170, fig. 4). Part 4( 1 ) provides an introduction to the whole group and systematic treatments of sect. 7 and part of sect. 9. Also treated are H. bupleuroides Griseb., in the monotypic sect. 8 Bupleuroides, and H. elatoides R. Keller (its nearest relative in sect. 3. Ascyreia), which was wrongly included in sect. 7. Roscyna in Part 1 (Robson, 1977). The opportunity has also been taken to remove H. ellipticifolium H.L. Li to a new genus; by implication it has hitherto been included in the rather amorphous sect. 9. Hyperi- cum. Before turning to these taxa, however, it is desirable to draw attention to two potentially misleading errors that crept into Part 6: ( 1 ) On p. 1 2 1 , the record of 2n = 1 6 chromosomes by Love & Love refers to H. majus (A. Gray) Britton (sect. Trigynobrathys), not to H. ellipticum Hook. (sect. Myriandra). All records for the latter section, therefore, are n = 9 or 2n = 18. (2) On p. 1 80, the record for H. aethiopicum subsp. sonderi in the middle of Namibia (Map 32) is an error, as reference to the text will show. The dot became displaced from among the south- east African records during printing. The Natural History Museum, 2001 38 Finally, as it has not been possible to continue producing the distribution maps by hand, they have been made on computer. However, this has necessitated the omission of co-ordinates and Chinese provincial boundaries. Text figures of leaf, petal and dia- grammatic inflorescence for each species have also, unfortunately, had to be omitted because of time constraints. Further delimitation of Hypericum - exclusion of H. ellipticifolium H.L. Li Hui-lin Li's (1944) description of a suffruticose Hypericum with white flowers (from the border of Yunnan and Myanmar) indicated the need for further investigation. An examination of the type and other specimens revealed that, as suspected, the flower had three very small fasciclodes inserted between the stamen fascicles, i.e. that the affinities of this species were with Triadenum, not Hyperi- cum. In fact it appears to be basic to the Triadenum-Thornea group, which, in Parts 1 (Robson, 1977: 300) and 2 (Robson, 1981: 63), I placed with Eliea and Cratoxylum in the Hypericoideae-Cratoxyleae. However, following the discovery by Gibson (1980) that the sec- ondary xylem of Thornea, Triadenum and Hypericum is organised on a common design that is quite different from that of the Cratoxyleae, I re-examined the problem and now agree with him and with Stevens (in press) that the Triadenum group is most closely related to Santomasia and Hypericum but forms a clade in the Hypericeae distinct from them. Lianthus N. Robson, gen. nov. Triadeno Rafin. affine, a quo habito suffruticoso, foliis canaliculis et glandulosis pellucidis continentibus, petalis albis, staminibus plurimus, differt. Suffrutex, glabrous, without black glands, with stems caespitose or shortly rhizomatous, slender. Stems and branches terete, eglandular. Leaves opposite, entire, exstipulate, venation pinnate, dorsal, with translucent ('pale') glands only; glands in 2 systems: (1) dorsal - punctiform, (2) ventral - closely parallel, linear or interrupted, arching from near base. Inflorescence terminal, few-flowered, N.K.B. ROBSON cymose, umbelliform. Flowers bisexual, homostylous. Sepals 5, quincuncial, persistent. Petals 5, white, subsymmetric, apiculate, deciduous. Stamen fascicles '3' (i.e. 2 + 2 + 1), with filaments in each fascicle united at the base, deciduous, the stamens totalling 1 1- 15; fasciclodes 3, very small, inserted between stamen fascicles, persistent. Ovary 3-locular with loculi many-ovulate; styles 3, free, elongate, slender; stigmas narrowly capitate. Fruit capsular, septicidal, with valves longitudinally and narrowly vitiate. Seeds numerous, small, fusiform, carinate, with testa finely foveolate; embryo not examined. Pollen grains not examined. TYPUS GENERIS. L ellipticifolius (H.L. Li) N. Robson. Genus adhuc monospecificum. Lianthus ellipticifolius (H.L. Li) N. Robson, comb. nov. Type as for Hypericum ellipticifolium H.L. Li. Fig. 1, Map 1. Hypericum ellipticifolium H.L. Li in J. Arnold Arbor. 25: 307 (1944); Li Xiwen in Fl. R. P. Sinicae 50(2): 8, t. 1 ff. 5-6 (1990). Type: China, Yunnan, Taron-Taru divide, valley of Bucahwang, 2200 m, 4 September 1938 (fl & fr), T.T. Yii 20125 (A!-holotype; KUN!-isotype). Icon: Li Xiwen in Fl. R. P. Sinicae 50(2): 8, t. 1 ff.. 5-6 (1990). Suffrutex 0.3-0.6 m tall, erect, caespitose or from shortly creeping and rooting base, with stems few. slender, sometimes branched above. Stems terete, eglandular; internodes 30-45 mm, equalling or usually exceeding leaves. Leaves sessile; lamina 30-50 x 15-30 mm, elliptic, paler beneath, not glaucous, chartaceous; apex rounded to subretuse, often obtusely apiculate, margin entire, base rounded; venation: 2-3(4) pairs of main laterals from lower third of midrib, with tertiary reticulation lax, only main veins visible (and promi- nent) below; laminar glands (dorsal) very small, irregular, dense; laminar glands (ventral) linear, curved-parallel from base and some- times crossing veins, alternating with vein-like narrower, distally interrupted glands; marginal glands rather dense. Inflorescence 5-7- flowered, subumbelliform, from terminal node only; pedicels Map 1 Lianthus ellipticifolius O; Hypericum sect. 3: 16a. H. elatoides STUDIES IN HYPER1CUM 39 a b Fig. 1 Lianthus ellipticifolius: (a) habit; (b) half leaf, showing primary and secondary venation, with tertiary venation (below) and venation system crossing veins (above); (c) sepal with fasciclode and 3-stamened fascicle; (d) flower with fruiting capsule; (e) sepal; (f) petal; (g) capsule beginning to dehisce (a x 2/3, b x 2, c-g x 6). (a) T.T. Yii 20868, except base; (a) base and (b)-(g) 717: KM 20125. 40 c. 8 mm; bracts 1-1.5 mm long, narrowly triangular. Flowers c. 15 mm in diam., stellate?; buds elliptic, obtuse. Sepals equal, 6-7 x 2 mm, erect in bud, reflexed in fruit, narrowly triangular-lanceolate, narrowly acute to acuminate, entire; veins 5, outer sometimes branched; laminar glands linear, usually 2 between each vein; marginal glands absent. Petals white, 10-12 x 3-4 mm, c. 2 x sepals, oblong-obovate, entire, with apiculus lateral, very short; laminar glands linear to striiform; marginal glands absent. Stamens c. 11-15, '3'-fascicled (6 + 5 + 4? to 5 + 3 + 3), 8-9 mm, c. 0.75 x petals; anther gland amber. Fascidodes 3, very small, lenticular. Ovary 3- locular, c. 5 x mm, ovoid; styles 3, c. 2 mm, c. 0.4 x ovary. Capsule 7x4 mm, about equalling sepals, ovoid; valves narrowly vitiate. Seeds dark brown, c. 0.6 mm, fusiform, acuminate at extremities, carinate; testa finely foveolate. Grassy slopes; 1800-2200 m. China (Yunnan). CHINA. Yunnan: Taron-Taru Divide, Tangtehwang to Bucahwang, 1 800 m, 29 October 1938(fl&fr), 7! 71 Yii 20868 (1BSC, KUN); west of Dulongjiang, 1800 m, 17 November 1959 (fl), G.M. Feng 24299 (KUN). See also type. Lianthus is named primarily after Hui-lin Li, who described its sole species as a Hypericum, but this name also commemorates Li Xiwen, who has published helpful accounts of Hypericum and Triadenum for several Chinese Floras. It differs from Hypericum in several respects, e.g. in having white petals and fasciclodes and, most notably, in its two systems of foliar glands. Viewed from above, the leaf has venation and pale punctiform glands similar to those of Triadenum and many Hypericum species. From below, however, although the main venation system is visible (indeed prominent), it is crossed by what appears to be another venation system of numerous parallel veins interspersed with linear glands. 1 These 'veins' and glands form two arching groups originating at or near the base of the leaf and each following the curve of the adjacent leaf margin. On closer inspection, the 'veins' in this system, which have cross-connections and X junctions similar to a normal primitive venation system (see Melville, 1976), appear to be secretory, not true veins; and so the whole system is a 'fossil' venation that seems to have been replaced functionally by the dorsal veins described above. This situation is known elsewhere in the Guttiferae, especially in the Clusioideae (e.g. Garcinia - Robson, 1961 ; Jones, 1980), and in the Thymelaeaceae (Melville, 1983); and the transition from one system to the other is noticeable in the most primitive taxa of Hypericum (H. bequaertii De Wild, and H. revolutum subsp. keniense (Schweinf.) N. Robson), see Robson (1981:80). With this suite of foliar differences, together with the floral differences already mentioned, it is clear that Lianthus cannot be derived from any species of Hypericum. Its immediate relationships are with Triadenum and Thornea, genera which I now place in the Hypericoideae-Hypericeae (see above). They form a group in that tribe distinct from Santomasia and Hypericum. This group apparently lacks the yellow flavonoid that normally occurs in the petals of Hypericum, and therefore their petals are white to red (white in Lianthus, white or pink to carmine in Triadenum, pink or white and pink in Thornea); they have few (3-5) stamens in each of three fascicles, so that the 'double' antesepalous fascicles have fewer than double the number of stamens of the single antepetalous one, often the same number. The range of chromosome numbers in Triadenum is now known to include 2n = 36 (Probatova & Sokolovskaya, 1 983, 1986) as well as 2n = 38 (n = 19, Hoar & Haertl, 1932). In relation to Hypericum, where the original base number appears to be x = 12, N.K.B. ROBSON the numbers in Triadenum would seem to be part of a tetraploid reduction series based on n = 10, i.e. 2n = 40. On the other hand, the morphologically more primitive T.japonicum (Blume) Makino has 2n = 36 whereas T. virginicum (L.) Rafin. has 2n = 38, and so the series could be an ascending one. In terms of geographical and evolutionary trends, the Triadenum group fits together well. Lianthus, a suffrutex with a terminal inflorescence, 1 1-15 stamens, and sessile leaves with the remains of an ancient venation system, is confined to a very small area in western Yunnan near the Myanmar border. Triadenum, marsh herbs with terminal and axillary flowers or partial inflorescences, 9 sta- mens, and sessile to petiolate leaves with only the 'recent' vascular and gland system, has one species in Assam and south China, one in Japan, Korea and adjacent Siberia and China, and four in eastern North America, all in lowland areas. Thornea, evergreen shrubs with inflorescences terminal and terminating lateral branches, 9 stamens, and petiolate leaves with only the 'recent' vascular and gland system (and the smallest flowers in the group), has one species in southern Mexico (Chiapas) and Nicaragua (Jinotega) and a second (more reduced) in Mexico (Chiapas) and Guatemala (Huehuetenango), both in montane habitats. The group would there- fore seem to have spread from south China northward, across the Bering Bridge and southward through North America to Nicaragua, the southern Asian species of Triadenum (T. brevifolium (Wall, ex Dyer) Y. Kimura) being a development from the northern T. japonicum. From this view, the shrubby habit of Thornea appears to be the result of an evolutionary reversion, but, since the genus in all other characters is advanced in relation to Triadenum, I cannot see a satisfactory alternative hypothesis. Relationship of H. elatoides and sect. 8. Bupleuroides (Fig. 2) Although Keller had described H. elatoides in 1904, placing it in sect. Norysca (i.e. sect. 3. Ascyreid), he omitted it from the 2nd edition of Die naturlichen Pflanzenfamilien (Keller, 1925). On the basis of the limited material available to me when working on Part 1 , 1 decided that the petals and stamens were persistent and the habit herbaceous, characters that led me to place this species in sect. 7. Roscyna (Robson, 1977). Subsequent examination of more abun- dant material has revealed that the petals and stamens are tardily deciduous and the habit suffruticose. Keller was therefore correct; H. elatoides belongs to sect. 3. Ascyreia and appears to be a derivative of the variable H. monogynum L. The tardily deciduous petals and stamens, the large leaves, often cordate-amplexicaul at the base, and the sometimes minutely reddish-glandular-ciliate se- pal margin all indicate a trend to the monospecific sect. 8. Bupleuroides, where the stamen fascicles are '3' and the ovary trimerous, the petals and stamens are persistent, the leaves are perfoliate, and the sepals are sometimes minutely reddish- to black- glandular-ciliate. The distributional gap between north-west China and the south-east Pontic region is paralleled by that between the areas of another part of H. monogynum and the monospecific sect. 6. Inodora (see Robson, 1985: 314). It is thus necessary to add H. elatoides to sect. 3. Ascyreia as Species 16a, which can be inserted in the key (Robson, 1985: 207) as follows: 20(16) Leaves densely reticulate-veined beneath, (20-)30-110 mm long; inflorescence 1 -30-flowered, terminal on long shoots 20a Leaves without visible reticulate venation beneath, 10-31 mm long; inflorescence always 1 -flowered, terminal on long and on short lateral shoots 19. longistylum 'Lii & Hu (unpublished) provide support for the generic separation of H. ellipticifolium from a study of epidermal micro-morphology in Hypericum and Triadenum. STUDIES IN HYPER/CUM 41 20a(20) Petals and stamens tardily deciduous; sepal margin sometimes minutely reddish-glandular-ciliate; plant a suffrutex .... I6a. elatoides Petals and stamens soon deciduous; sepal margin entire; plant a shrub 21 21(20a) Leaves with base cuneate to rounded, or if cordate then apex rounded; leaf lamina usually broader at or above middle 16. monogynum Leaves with base cordate-amplexicaul, apex acute to acuminate; leaf lamina elliptic-ovate to broadly ovate 18. prattii Relationships of sects 7. Roscyna and 9. Hypericum sensu lato (Fig. 2) With the removal of Hypericum elatoides to sect. Ascyreia, sect. Roscyna consists of two variable herbaceous species with regular pentamerous flowers and persistent petals and stamens: H. ascyron L. and H. przewalskii Maxim. H. pedunculatum R. Keller, which was included with a query in Part 1 , cannot be distinguished from H. przewalskii. It is the nearest form of that species morphologically to H. ascyron. H. ascyron is distributed from the Altai region and adjacent Xinjiang through southern Siberia to Kamchatka and south to Japan, China (except Xizang and Qinghai), Taiwan and Vietnam, and also in eastern North America. It can be divided into three subspecies: (i) subsp. ascyron (Siberia from Ob and Altai eastward to Ussuri, Japan, Korea, China, Vietnam, Taiwan), (ii) subsp. gebleri (Ledeb.) N. Robson (Xinjiang, upper Altai region, montane and far-eastern parts of Siberia, northern Mongolia, northern China, Sakhalin, northern Kurile Is. and Kamchatka), (iii) subsp. pyramidatum (Aiton) N. Robson (eastern North America). Subsp. ascyron (the morpho- logically primitive taxon), having apparently given rise to the other two subspecies, is most closely related to H. podocarpoides (sect. 3. Ascyreia), endemic to central Nepal. The whole of the Himalayan massif and Tibetan Plateau thus separates the ranges of these species. H. przewalskii, which occurs in western China from Qinghai and Gansu south to Yunnan, differs from H. ascyron in its usually 2-lined to terete stem internodes and usually broader leaves with obtuse to retuse apex. '//. pedunculatum' is a narrow-leaved much-branched form of H. przewalskii from western Hubei that is linked to the more typical unbranched broad-leaved form by intermediates. With virtually no intermediate forms between the regular pentamery of the floral whorls of sect. Roscyna and the trimery and pseudotrimery of the inner whorls in sect. Hypericum sensu lato (apart from one or two aberrant cases of tetramery in the gynoecium (and androecium?) , e.g. H. kelleri H. Leveille (=//. ascyron) and H. paradoxum R. Keller (=H. kamtschaticum)), previous authors (e.g. Keller, 1925) have placed nearly all herbaceous species with trimerous inner floral whorls in a large section Hypericum or Euhypericum. The absence of black glands from sect. Roscyna species and their invariable presence at least somewhere in those of sect. Hypericum seemed to confirm their classification. We have already seen, however, that this unwieldy 'section' contains three distantly related groups - sects 9, 10-19 and 27 (Robson, 1977, 1996). It is perhaps, therefore, not surprising that sect. 9. Hypericum itself can be broken down further into sections and subsections. Dismemberment of sect. 9. Hypericum Sect. 9. Hypericum has been shown to comprise seven recognizable taxa. Although they are all related to species in sect. 7. Roscyna, six of them are not directly related to one another; so they have been treated as sections and the two interrelated ones as subsections (Figs 2-4). (i) The north Californian endemic H. concinnum Benth. has already been removed from sect. 9 to its own section, sect. 9a. Concinna N. Robson (Robson, 1981: 173), on the basis of its narrow concolorous, often conduplicate leaves, large sepals and amber anther gland. The only part of sect. Roscyna that approaches it morphologically is H. ascyron subsp. gebleri, in which the leaves are sometimes very narrow ('//. sachalinense H. Leveille') and the sepals are not very different from the narrowly to broadly ovate ones of H. concinnum. In addition. la. ascyron ssp. ascyron (22,20) 18(16) 16a. elatoides- 8. Bupleuroides 9a. Concinna Ib. ascyron ssp. gebleri 18 16 Ic. ascyron ssp. pyramidatum 16, 16,32,48 24,40 16,32 Fig. 2 Sections 7-9e. Relationships and chromosome numbers (2n), indicating derivations from sect. 3 Ascyreia. 42 N.K.B. ROBSON the presence of subsp. gebleri in Kamchatka and the northern Kurile Is. suggests that it or its derivative could well have crossed over to North America. (ii) The other North American group belonging to sect. Hyperi- cum occurs in the east, in extreme southern Canada, the U.S.A., eastern and southern Mexico and Guatemala. As this group proved difficult initially to separate from the Japanese species of this section, another trans-Pacific distribution appeared to be involved. Such a hypothesis turned out to be unnecessary, however, when the stem glandular patterns were compared. Furthermore, the acute sepals with linear laminar glands and the 4-lined young stem internodes of the most primitive species (H. graveolens Buckley) could have been derived only from the North American subspecies of H. ascyron, subsp. pyramidatum. The anther gland is exception- ally amber in one species, H. pseudomaculatum Bush. This group has thus been given sectional rank as sect. 9b. Graveolentia sect, nov. The five remaining groups can be divided into those that can be derived directly from H. ascyron subsp. ascyron (with stem internodes basically 4-lined) and those most closely related to H. przewalskii (with stem internodes basically terete). (iii) The basically 4-stem-lined group comprises one subgroup (iiia) in which the rootstock is herbaceous and stoloniterous and 2 or 4 stem-lines are always present, at least when young, and usually bear at least some dark glands, and another subgroup (iiib) with a woody or fibrous rootstock and eventually terete stems, from which stem-line glands are always absent. Since group (iii) contains the generic type, H. perforation L., it remains as sect. 9. Hypericum and subgroup (iiia) becomes subsect. 1 . Hypericum. Subgroup (iiia) (i.e. subsect. Hypericum) comprises two sub- groups with distinct distributions (a: Eurosiberia, and also Japan, and western North America; b: north Japan, Korea and central China), which have as respective basic taxathe Balkan H. maculatum subsp. immaculatum (Murb.) A. Frohl. and the northern Japanese H. kamtchaticum var. pibairense Miyabe & Y. Kimura, taxa that resem- ble one another closely. Subgroup (a) in turn has two centres of distribution (Europe-western Siberia and eastern Siberia-western North America), the parts of which are now linked by the Asiatic distribution of H. perforation, a presumed hybrid with one parent in each part, and H. elegans Stephan ex Willd., an eastern species that has spread westward from central Asia into Europe. The species of subgroup (b) lack black glands on the stem-lines, which eventually disappear in growth, whereas black-gland-bearing stem-lines are all but constant in subgroup (a). These two subgroups can thus be distinguished respectively as (iiiaa) series 1 . Hypericum and (iiiab) series 2. Senanensia series nov. The other main subgroup (iiib) contains the widespread and variable H. erectum Thunb. and has been named subsect. 2. Erecta subsect. nov. Stem lines are present only in the primitive taxa (e.g. H. yamamotoi var. riparium Y. Kimura) and, even then, rarely persist into maturity; the stems never bear dark glands except in H. nikkoense Makino, which sometimes has scattered black or reddish ones. Its distribution is centred in northern Japan, extending into the rest of Japan and adjacent areas of the Russian Federation, Korea and China as well as Taiwan, the Philippines and Borneo. (iv), (v) and (vi) Three groups are derived morphologically from H. przewalskii but appear to be related to different parts of this variable species. Although all have terete eglandular mature stem internodes (except sometimes when the stem is very slender, when they may be 2-6-lined and, in one species, rarely with some reddish glands on the lines), they are otherwise quite different, (iv) H. sampsonii Hance has a wide distribution in east Asia from southern Japan and Taiwan to central Myanmar; and a clearly derived species, H. assamicum Biswas, extends the area of this group to Assam (Meghalaya). These plants have leaves like those of '//. pedunculatuni , but they are perfoliate and broader at the point of union. The inflorescence, however, is multifloral, the flowers are much smaller with a subcupuliform base, and the three capsule valves bear elongate to punctiform amber vesicular glands. Taken together, these characters are quite diagnostic and indicate that these two species merit segregation into a separate section, 9c. Sampsonia sect. nov. The perfoliate leaves of H. bupleuroides and the some- times vesicular capsule valves of H. perforatum are parts of quite different character syndromes. There is no evidence supporting Kimura's (1951) allocation of H. sampsonii to the mainly Balkan- Mediterranean sect. 13. Drosocarpium on account of the vesicular capsule valves. (v) Another group of species related to H. przewalskii is linked to the narrow-leaved branching form, '//. pedunculaturn', through its most primitive species, H. seniawinii Maxim. In addition to the trimery of the inner floral whorls and the intramarginal foliar black glands, H. seniawinii differs from '//. pedunculatum' in having acute sepals with marginal black glands. It resembles the latter, however, in its relatively large and dense laminar leaf glands. Other species in this group have black-glandular fringes to auriculate appendages of the leaves or sometimes of the leaf itself. The distribution of the group extends from east and south China to north Vietnam, thence westward along the Himalaya to Kashmir. The section has been named after the widespread H. elodeoides Choisy, sect. 9d. Elodeoida sect. nov. (vi) The final group is related to the few-flowered broad-leaved form of H. przewalskii that occurs in Sichuan and adjacent Yunnan ('//. macrosepalum'). Like it, this group of species has small laminar leaf glands that are usually sparse and may be almost absent, and its primitive species have similar broad leaves. Its distribution extends from south and south-west China through north Laos, north Thai- land and north Myanmar along the Himalayan range to Pakistan, and it also occurs in south India and Sri Lanka. The section has been named after the widespread H. monanthemum Hook. f. & Thorns, ex Dyer, sect. 9e. Monanthema sect. nov. The subdivision of sect. 9. Hypericum sensu lato can be summa- rised as follows: Sect. 9. Hypericum. Mature stem internodes 2^-lined or terete; dark glands, when present, nearly always confined to lines. Leaves free, entire, exappendiculate, plane to marginally recurved; laminar glands dense to sparse or absent. Bracteoles entire or very rarely glandular-denticulate (H. elegans), reduced. Flowers stellate. Perianth 5-merous. Sepal laminar glands nearly always striiform to punctiform. Anther gland black. Capsule valves longitudinally vittate or rarely obliquely vittate to vesiculate, vesicles amber. 42 species. Subsect. 1. Hypericum (Part 4(2,3)). Rootstock herbaceous, stoloniferous. Mature stem internodes usually persistently 2-4- lined; dark glands usually present, nearly always confined to lines. Capsule valves nearly always longitudinally vittate, occasionally interrupted or with oblique swollen lateral vittae or vesicles. 19 species. Series 1. Hypericum (Part 4(2)). Stems persistently lined; lines prominent (except in 12. H. scouleri Hook.), sometimes almost winged, bearing dark glands (sometimes except 1 2. H. scouleri). 12 species. STUDIES IN HYPER1CUM 43 Series 2. Senanensia (Part 4(3). Stems becoming terete; lines weak, eglandular. 7 species. Subsect. 2. Erecta (Part 4(3)). Rootstock woody to fibrous, not stoloniferous. Mature stem internodes slightly 2(4)-lined or usu- ally terete; dark glands nearly always absent, when present not on lines. Capsule valves longitudinally vittate. 23 species. Sect. 9a. Concinna (Part 4( 1 ), p. 6 1 ). Mature stem internodes 2-lined to terete, eglandular. Leaves free, entire, exappendiculate, usually conduplicate; laminar glands small, dense. Bracteoles entire, reduced. Flowers stellate. Perianth 5-merous. Sepal laminar glands linear. Anther gland amber. Capsule valves longitudinally vittate. One species. Sect. 9b. Graveolentia (Part 4(3)). Mature stem internodes 2-4- lined and sometimes ancipitous above, nearly always terete below; eglandular or with dark (black) glands on or near lines or site of lines when young or scattered. Leaves free, entire, exappendi- culate, plane or recurved; laminar glands dense to sparse. Bracteoles entire or with prominent marginal glands or basally glandular-ciliate. Flowers stellate. Perianth 5-merous. Sepal laminar glands linear to punctiform. Anther gland black or rarely amber. Capsule valves longitudinally vittate or rarely elongate- to ovoid-vesiculate, vesicles amber or rarely black. 9 species. Sect. 9c. Sampsonia (Part 4( 1 ), p. 63). Mature stems terete, eglandular. Leaves perfoliate, entire, exappendiculate, plane; laminar glands dense. Bracteoles entire, foliar or reduced. Flowers basally cupuliform. Perianth 5-merous. Sepal laminar glands striiform to punctiform. Anther gland black. Capsule valves elongate- to punctiform-vesiculate, vesicles amber. 2 species. maturorum teretibus eglandularibus vel rarissime disperse rubiginoso-glandularibus, valvis capsularis semper longitudinaliter vittatis, differt. Typus: H. erectum Thunb. Sect. 9b. Graveolentia N. Robson, sect, nov.: a sect. Hypericum internodiis caulium maturorum persaepe basin versus teretibus, interdum eglandularis vel glandulis haud ad lineas prominentes limitatis, sepalorum glandulis laminaribus linearibus vel striiformibus vel raro punctiformibus, antherae glandula raro succinea, capsulae valvis longitudinaliter vittatis vel raro elongati- vel ovoidei- vesiculatis vesiculis rariore nigris, differt. Typus: H. graveolens Buckley Sect. 9c. Sampsonia N. Robson, sect, nov.: a sect. Hypericum internodiis caulium maturorum teretibus eglandularibus, foliis perfoliatis semper planis, floribus basi cupuliformibus, capsulae valvis disperse plusminusve elongati- vel punctiformi-vesiculatis, differt. Typus: H. sampsonii Hance Sect. 9d. Elodeoida N. Robson, sect, nov.: a sect. Hypericum internodiis caulium maturorum teretibus vel ubi tenuibus 2-lineatis eglandularibus, foliis interdum glanduloso-fimbriatis vel auriculatis auriculis glanduloso-fimbriatis, bracteolis semper glanduloso- fimbriatis, differt. Typus: H. elodeoides Choisy Sect. 9e. Monanthema N. Robson, sect, nov.: a sect. Hypericum internodiis caulium maturorum teretibus vel ubi tenuibus 2^(6)- lineatis eglandularibus vel rarissime lineis rubroglanduliferibus, foliis interdum glanduloso-fimbriatis, bracteolis interdum glanduloso-fimbriatis, differt. Typus: H. monanthemum Hook. f. & Thorns, ex Dyer Key to sections, subsections and series in Part 4 Sect. 9d. Elodeoida (Part 4(1), p. 66). Mature stem internodes terete or, when slender, 2-lined, eglandular. Leaves free, sometimes with gland-fringed auricles, occasionally lamina gland-fringed, plane; laminar glands relatively large, dense. Bracteoles often 1 Stamen fascicles and styles 5(4); black glands absent; shrubs, suffrutices with gland-fringed auricles, reduced. Flowers stellate or rarely or herbs infundibuliform. Perianth 5-merous. Sepal laminar glands linear Stamen fascicles '3'('4'); styles 3(4); black glands nearly always to punctiform. Anther gland black. Capsule valves longitudinally present; suffrutices or herbs 3 vittate. 5 species. 2( 1 ) Petals and stamens deciduous (sometimes tardily); shrubs or suffrutices 3. Ascyreia Sect. 9e. Monanthema (Part 4(1), p. 75). Mature stem internodes terete or, when slender, 2^(6)-lined, eglandular or very rarely Petals and stamens P ersistent; herbs - with reddish glands on lines. Leaves free, entire or gland-fringed 3( 1 ) Leaf pairs perfoliate and then with glandular-ciliate auricles, plane; laminar glands Leaf pairs free 5 relatively small, sparse to occasionally dense. Bracteoles with 4(3) plowers 2? ^ mm jn djam ste , late; styks appressed at base; capsule gland-fringed auricles and reduced or entire and then usually va , ves longitudina ii y v j tta te; black glands usually absent foliar. Flowers stellate. Perianth 5(4)-merous. Sepal laminar glands 8. Bupleuroides linear to punctiform. Anther gland black. Capsule valves longitu- ^^ ^ 15 mm in diam substd , ate with cupuliform base; styles dinally vittate. 7 species. outcurving from base; capsule valves vesiculate; black glands present 9c. Sampsonia The new taxa in the above summary are validated by the following diagnoses' ^(3) Leaves often conduplicate, very narrow (l:b = 4-10); anther gland amber; sepals imbricate 9a.Concinna Sect. 9. Hypericum subsect. Hypericum series Senanensia N. Leaves plane or recurved, broad to narrow but l:b = < 4; anther gland Robson, series nov.: a series Hypericum caulibus demum teretibus black (rarely except in Sect. 9b) ... 6 lineis debilibus eglandularibus, differt. 6(5) Stem intemodes persistently 2-4(6)-lined or -winged; dark glands on Typus: H. senanense Maxim. and usua n y confined to raised lines 7 Sect. 9. Hypericum subsect. Erecta N. Robson, subsect. nov.: a Stem internodes terete or usually becoming terete; dark glands absent or subsect. Hypericum caudice lignoso stolonifero, internodiis caulium not on raised lines 9 Descending numbers of character values in descriptions show the presumed direction of evolution. Thus 5(4) indicates that 5 is the primitive number and is usual, but that a reduction to 4 sometimes occurs. On the other hand, 3(4) indicates merely that 3 is the usual number but that 4 sometimes occurs. 44 N.K.B. ROBSON 7(6) Bracts and bracteoles exauriculate; stems usually stout 8 Sect. 7. RoSCyna Bracts and bracteoles auriculate; stems slender 9e. Monanthema (part) Characters and variation (Figs 2, 3) 8(7) Sepal laminar glands striiform to punctiform; styles 2-6 mm long ... MORPHOLOGY. The species in sect. 3. Ascyreia to which Hyperi- 9. Hypericum subsect. 1 . Hypericum series 1 . Hypericum cum ascyron is most closely related appears to be H. podocarpoides, which is almost endemic to Nepal but also occurs in Kumaun (Uttar Sepal laminar glands all or mostly linear; styles 5.6-1 2 mm long Pradesh). H. ascyron subsp. ascyron differs from it in its herbaceous 9b - Graveolentia (part) habit with one or a few erect stems; its usually broader leaves with 9(6) Stem internodes with dark glands scattered or in 2 ranks 10 densely reticulate, tertiary venation, the pale glands striiform to punctiform in the areoles; its persistent petals and stamen fascicles; Stem internodes eglandular 1 1 i ts i ar g er flowers with relatively broad, usually obtuse sepals and 10(9) Sepals obtuse to rounded; stems caespitose .... spathulate to subunguiculate, often curved petals deflexed after 9. Hypericum subsect. 2. Erecta (part) flowering; its united styles; and its larger ovoid capsules. Although most of these characters are advanced relative to those of H. podo- Sepals acute to obtuse or rarely rounded; stems creeping carpoides, the smaller flowers and condensed inflorescence axis of 9b - Graveolentia (part) t he latter species suggest that the relationship is not direct (Robson, 1 1 (9) Bracts and sometimes leaves with glandular or gland-fringed auricles ^ ' ^' "& *) 12 Of the two derivative subspecies, subsp. gebleri is a plant of more northern or montane regions of eastern Asia with narrower, cuneate- Bracts and leaves not auriculate; leaves and often bracts entire 13 based leaveS; smaller flowers with narrower sepals and shorter free 12(11) Leaves with laminar glands relatively large, dense; lamina narrowly st y les ' and a narrower capsule broadest at the middle. Subsp. oblong to linear or if relatively broader, then to 10 mm pseudopetiolate pyramidatum, on the other hand, is confined to north-eastern North 9d. Elodeoida America and differs from subsp. ascyron in its acute to acuminate sepals and almost free styles. Leaves with laminar glands relatively small, usually sparse; lamina An three subspedes have 4 _ lined stem intern odes and acute to broadly oblong to triangular-ovate or suborbicular or oblanceolate, up , ,- , ... . -,,. TI , , .. , . - on* obtuse lear apex, whereas in the west Chinese H. przewalskn the to 3 mm pseudopetiolate 9e. Monanthema (part) internodes are partially 2-lined or wholly terete when mature, and 13(1 1 ) Bracts foliar; perianth 4-merous 9e. Monanthema (part) the apex of the usually broader leaves is rounded to shallowly retuse. Other variation in those taxa is discussed in the systematic Bracts reduced; perianth 5-merous or variable (4-9-merous) 14 , s? 611 14(13) Rootstock herbaceous, stoloniferous; stems dispersed ... r^i^^, ,.,, x T - t /_, j > i . CYTOLOGY. Neither//, podocarpoides nor its near relative in sect. 9. Hypericum subsect. 1. Hypericum series 2. Senanensia , . ,.,. ,. -,, . Ascyreia, H. cordijolium Choisy, has a known chromosome number. Rootstock woody to fibrous, not stoloniferous; stems caespitose .... However, as the known diploid numbers in that section vary from 24 9. Hypericum subsect. 2. Erecta (part) to 20, and 22 is very rare, the number of H. podocarpoides is likely Stamen fascicles & styles 3(4) / 5 Mature stem internodes 2-4-lined i terete N X _ I!-. J _ styles JU--4--^~~~ free united or appressed Fig. 3 Sections 7-9e. Limits of certain characters. For names of sections and species see Fig. 2 (p."). STUDIES IN HYPERICUM 45 -16a. N.China S. Japan, Taiwan, C. & S. China Vietnam, Myanmar, Assam S. & E. Asia I 3 Kumaun & Nepal Turkey SW Georgia N California Siberia, Japan, Korea China, Taiwan, Vietnam SW China, Vietnam, Laos Thailand, Myanmar to Pakistan S. India, Sri Lanka NE Kazakhstan, Xinjiang Siberia, Mongolia, N. China Sakhalin I Kamchatka lc SE Canada NE United States Macaronesia to NW India Europe to Japan & China || Br. Columbia to C. Mexico SE Canada, E. United States Mexico, Guatemala Kamchatka to Sakhalin Japan. Korea, SE China Taiwan, Luzon I Sabah Fig. 4 Sections 7-9e. Distribution of species, showing major (II) and minor (I) disjunctions. For names of sections and species see Fig. 2 (p. 41 ). to be 2n = 24 or 20 (see Fig. 2). For a long time H. ascyron subsp. ascyron was recorded as having only 2n = 18, but Russian workers have more recently shown it to have a range of numbers (16-22?), although 18 remains the usual one (c. 20-22, Krasnoborov et al., 1980; 16, Krogulevich, 1978). The only record for subsp. gebleri is also 2n = 18 (Stepanov, 1994). Chromosome numbers for subsp. pyramidatum and for H. przewalskii have yet to be determined. No hybrids of species in sect. Roscyna have been reported. Distribution and evolution (Fig. 4) The ranges of H. podocarpoides (Nepal) and its closest herbaceous relative H. ascyron subsp. ascyron (southern Siberia) are separated by the higher Himalayan range and the Tibetan Plateau, which were elevated as the result of the collision of the Indian subcontinent with the mainland in the late Paleocene or early Eocene, 55-60 m.y. B.P., and the gradual widening of the contact area eastward during the Eocene (Axelrod et al., 1998: 44). The subsequent elevation of the Tibetan Plateau resulted in the spread of grassland vegetation into this region (Hsu, 1 983), an event that appears to have coincided with (caused?) the change in habit from shrub (H. podocarpoides) to herb (H. ascyron) and would have provided a suitable habitat for the latter. As the climate became increasingly desertified from the middle Miocene (15 m.y. B.P.) (Guo, 1981), the Tibetan Plateau would have become a barrier between the ranges of these species or their respective ancestors, confining H. ascyron to its northern rim in Siberia. The subsequent spread of H. ascyron westward to the Altai Region and eastward as far as Kamchatka, and the development of a higher altitude/latitude subspecies (subsp. gebleri) requires no special hypothesis, even though the distribution of the latter is now to some extent disjunct. The occurrence of H. ascyron in north- eastern North America, however, could be explained by a migration either eastward across the Bering Bridge or westward across Europe and the Atlantic. No fossil evidence for its occurrence in Europe has been found and the breaking of the Atlantic Bridge in the late Cretaceous (c. 81 m.y. B.P.; Raven & Axelrod, 1974: 544) was too early to allow the passage of//, ascyron to America from the east, as was the latest possible time for direct overland migration across the North Atlantic (49 m.y. B.P; McKenna, 1972). The Bering Bridge hypothesis is therefore to be preferred. The apparent evolution of subsp. gebleri from subsp. ascyron (and not vice versa), the postu- lated migration across the Bering Bridge of subsp. gebleri or the ancestor of H. concinnum is likely to have been later than the migration of subsp. ascyron. If so, this time difference would support the inference of Hui-lin Li (1952, cited by Kruckenberg, 1983: 594) that the western North American-eastern Asian disjunct group of species (in all families) is (i) more recent than the eastern North America-eastern Asian one and (ii) predominantly herba- ceous rather than woody. Sect. 9a. Concinna The apparent relationship of H. concinnum to H. ascyron subsp. gebleri has already been shown in Fig. 2 and the distributional and evolutionary connections discussed on p. 4 1 . These are shown in Figs 2-4. Note that the transition of the inner floral whorls from pentamery to pseudotrimery and trimery respectively again coincides (almost entirely) with a change of diploid number from 18 to 16 (Figs 2, 3). Sects 9c. Sampsonia, 9d. Elodeoida and 9e. Monanthema Characters and variation (Figs 5, 6) MORPHOLOGY. Sects Sampsonia, Elodeoida and Monanthema (ex- cepting the slender-stemmed species mentioned, p. 43) agree in having terete eglandular stem internodes but otherwise look very 46 N.K.B. ROBSON distinct. Quite apart from the perfoliate leaves, subcupuliform co- rolla and vesiculate capsule of Sampsonia, its species have leaves and sepals that are entire with intramarginal black glands, the sepals being oblong-spathulate to linear and rounded and never foliose. The marginal glands of the petals are sometimes on cilia, whereas in Elodeoida they are never more than subsessile and in Monanthemum at most one glandular cilium occurs. In its mostly rather narrow leaves H. sampsonii resembles the more primitive, ' pedunculatum form of//, przewalskii (sect. Roscyna); but the 1-7 large flowers of that species are replaced by 20-40 small ones, the leaf pairs are united, not free, the stamen fascicles are pseudotrimerous and the ovary trimerous with free outcurving styles. The somewhat incurved petals are said by Momiyama (1982) to have 'raised fissures' (raised veins?) at the base, a feature that I have not been able to observe on dried material. The differences between H. sampsonii and H. assamicum (the other species in sect. Sampsonia) are mainly of size, but the ovary in the latter is said to be unilocular rather than trilocular, a character that, again, I have been unable to confirm. The basic species of one clade of sect. Elodeoida, 1 . H. seniawinii, also has the narrow leaves of the 'pedunculatum' form of H. przewalskii, but the leaf pairs remain free and sessile, and their laminar glands are relatively large, round and dense, as in that form. The leaves in the other members of that clade (2. H. petiolulatum subsp. yunnanense (Franchet) N. Robson and subsp. petiolulatum respectively) become gradually smaller, relatively broad and, after developing a pseudopetiole, eventually suborbicular. The leaves, bracts and (usually) sepals remain entire. In the other clade, 3. H. hengshanense W.T. Wang also has narrow leaves with large, dense laminar glands; but the upper ones sometimes have marginal glan- dular cilia, which are constantly present in the bracts and sepals. Gland-fringed auricles, which are sometimes found in an early stage of development in H. seniawinii, are here constantly present in the upper leaves and bracts. The closely related 4. H. elodeoides is a usually smaller and acute-leaved derivative of//, hengshanense that has spread along the Himalayan range as far as Kashmir but is absent from Assam and Myanmar. The distributional gap is partly filled by subsp. wardii N. Robson, with obtuse to rounded leaves and a reversion to entire sepals and bracts. In the north of the same 'gap', in north Myanmar, Tibet and adjacent Arunachal Pradesh, is found a high alpine relative of subsp. wardii with much-branched stems, small broad leaves and irregularly gland-fringed to subentire sepals, H. kingdonii N. Robson. Whereas sects Sampsonia and Elodeoida are related to the more primitive ' pedunculatuni 1 form of//, przewalskii, sect. Monanthema is clearly related to a broad-leaved form of that species in which the flowering branches are few or absent. In the south-western part of its range (in south-western Sichuan and adjacent Yunnan), H. przewalskii occurs as an extreme development of such a form, having an unbranched stem and only 1-3 flowers, in which the sepals tend to be foliose ('//. macrosepalum' Rehder). In addition, the laminar leaf glands tend to be smaller and less dense than in the 'pedunculatum' form. All these characters, except foliose sepals, are found in the primitive (Yunnan) form of 1. H. monanthemum (la. subsp. monanthemum); foliose sepals and bracts occur in Ib. //. monanthemum subsp. filicaule (Dyer) N. Robson. Small, more or less sparse laminar leaf glands are characteristic of the whole section. Evolution in sect. Monanthema has apparently proceeded along three clades (Fig. 5). In the Monanthemum group the simple stem is retained until the later stages of development, e.g. the extreme small la. ascyron ssp. ascyron (22,20)18(16) 2. przewalskii 'pedunculatum' 'przewalskii' 9c. Sampsonia 1. sampsonii / 2. assamicum \. seniawini 2a. petiolulatum ssp. yunnanense 2b petiolulatum ssp. petiolulatum Also 9d. 4a * 9e. 4 H. elodeoides ssp. elodeoides x H. himalaicum -macmsepalum' 9e. Monanthema 6. daliense . - 3. trigonum 1 . wightianum 5. ludlowii la. monanthemum 4. himalaicum ssp. monanthemum \^ ~~~2. subcordatum 3. hengshanense Ib. monanthemum ssp. filicaule 4a. elodeoides ssp. elodeoides 16,32 4b. elodeoides ssp. wardii "^ ^^ 5. kingdonii Fig. 5 Sections 9c-9e. Relationships, chromosome numbers (2n) and presumed hybrid. STUDIES IN HYPER1CUM 47 Stamen fascicles and styles 3(4)7 5(4) black or red glands absent/ present Capsular glands vitiate--- -""' Leafp"affr free entire ...'' / Sepals ..-"'' 2a ..- .*** ^ ** gland-fringed' \arge\ small leaf laminar glands Fig. 6 Sections 9c-9e. Limits of certain characters. For names of sections and species see Fig. 5 (p. 46). diffuse forms of Ib. H. monanthemum subsp.filicaule; here also the inflorescence is rarely more than 5-flowered and almost always from the terminal node only. The leaves and bracts are always entire, and the transition to 4-merous sepals and petals in H. monanthemum subsp.filicaule is accompanied by a return to entire (foliose) bracts and sepals. Extreme forms of this subspecies have a 2-merous ovary. In the other (northward) development of this clade, the slender- stemmed 2. H. subcordatum has also developed entire sepals, but these are not foliose. In the other two clades of sect. Monanthema the inflorescence has become more branched, the branches more floriferous and the bracts and sometimes upper leaves auriculate. 3. H. trigonum Hand.-Mazz. (Trigonum group) is linked to H. monanthemum by a rather tall Sichuan population of the latter with an unbranched inflorescence but unusually long, slender styles. Its specific epithet refers to the triangular-ovate leaves that are characteristic of most of the group, as are glandular-ciliate bracts and sepals. There is, however, a reversion to entire sepals and bracts in extreme forms of 4. H. himalaicum N. Robson and the leaves in the Trigonum group are always entire. South of the Himalayan range, 4. H. himalaicum has developed from H. trigonum with smaller leaves and flowers and shorter styles, relatively and absolutely. To the north of that range, H. trigonum has given rise to 5. H. ludlowii N. Robson, in which the more erect stem has shorter branches and the small flowers have relatively longer styles, achieved by a greater reduction in the size of the ovary than in the length of the styles themselves. The frequency of red 'dark glands' in this species indicates a diminution in the concentration of hypericin, as was seen in H. elodes L. (sect. 28. Elodes; Part 6, Robson, 1996: 209). The third clade of sect. Monanthemum consists of two species, the widespread 7. H. wightianum Wall, ex Wight & Arn. and an endemic of the Dali and Lijiang ranges in Yunnan, 6. H. daliense N. Robson. In both these species the few- and large-flowered inflorescence of//. przewalskii and H. monanthemum is replaced by one with numerous small flowers; but the elliptic-oblong leaves are the most similar in this section to those of//, przewalskii ('macrosepalum' form). This clade is unlikely, therefore, to have been derived from either of the others (Fig. 5). Other developments in this clade that are unique in the section include the frequent occurrence of (a) a complete or basal glandular-ciliate margin in the upper leaves of H. wightianum and (b) marginal black glands between the glandular cilia in the sepals of both species but particularly those of H. wightianum. As in the Trigonum group, the bracts are glandular-auriculate and in the present group so are the gland-fringed leaves. The glandular fringes of the sepals vary from denticulate (in H. daliense) to ciliate and laciniate (in most of//, wightianum); but in the extremely reduced form of the latter species in south India and Sri Lanka, there is a reversion to entire sepals and bracts. The additional development of shortly petiolate leaves in this form has given rise to a plant that has been confused with the European H. humifusum L. (sect. 15. Oligostema) ever since the account of Hypericum was published in the Flora of British India (Dyer, 1874). CYTOLOGY. Practically no cytological data have been published on species in sects 9c-e. Chromosome counts have been made only on H. elodeoides (sect. 9d. Elodeoida), which has diploid and tetraploid populations on the base 8, i.e. 2n = 16, 32. No evidence regarding an association of this difference of ploidy level with morphological characters is available. Distribution and evolution (Fig. 7) 9c. S AMPSONIA. With its distinctive perfoliate leaf pairs, elongate entire sepals and vesiculate capsules, sect. Sampsonia would appear to have been an early development from H. przewalskii "pedunc- ulatum" (clade i); and the wide distribution of its main species, //. sampsonii, supports this idea. From a notional origin in west China (based on the present variation and distribution of//, przewalskii), it has spread north to eastern Gansu and Jiangsu, east to southern Japan and Taiwan, and south to the rest of China and extreme north Vietnam. Unlike members of sects 9d and 9e, it has not penetrated 48 N.K.B. ROBSON into Yunnan or the Himalayan massif but is found in eastern Myanmar; the very closely related H. assamicum extends the sectional distribution to Meghalaya. 9o. ELODEOIDA. The other two sections are more similar to each other than either is to sect. 9c, even though the morphological evidence suggests that 9d is related to 'pedunculatum , whereas the affinities of 9e are with 'macrosepalurn . The basic species in one clade (clade ii) of sect. Elodeoida, 1. H. seniawinii, has a rather similar mainland distribution to that of H. sampsonii, but is absent from Japan and Taiwan. To the west it intergrades geographically, though not morphologically, with 2a. H. petiolulatum subsp. yunnanense, the area of overlap including parts of Henan, Hubei, Hunan, Jiangxi, Guangxi and Guizhou. Subsp. yunnanense in turn is replaced in Sichuan and Yunnan by 2b. H. petiolulatum subsp. petiolulatum, which is distributed through the main Himalayan range to Nepal. The other clade (clade iii) has two closely related taxa at the base. 3. H. hengshanense has a limited relict distribution in south-east China (Jiangxi, Guangdong and northern Guangxi), whereas the more variable and morphologically more advanced 4a. H. elodeoides subsp. elodeoides occurs throughout southern China from Fujian, Jiangxi, Hubei and Sichuan southward and extends along the main Himalayan range as far as Kashmir and into Meghalaya. Although its range in China includes all that of H. hengshanense, this overlap should be interpreted as resulting from an invasion by H. elodeoides from the west, because the Chinese plants are morphologically advanced, the most primitive forms being in the central Himalaya. The other subspecies, 4b. H. elodeoides subsp. wardii, is a south- ward development into Manipur and central Myanmar (Mt. Victoria); this seems to have given rise to a small plant of extreme northern Myanmar, adjacent Xizang and extreme eastern Arunachal Pradesh, 5. H. kingdonii. 9E. MONANTHEMA. This section is related to the most advanced form of//, przewalskii from the southwestern-most part of its range (Sichuan and northern Yunnan), '//. macrosepalum ; the basic species of all three of its clades occur in or near this region. The basic taxon of clade iv, la. H. monanthemum subsp. monanthemum, has its most primitive form in northern Yunnan and has spread out to both south- west and north-east. South-westward there is a reduction trend in size through Yunnan and along the Himalayan range to Nepal, where there are some plants intermediate in form between it and Ib. //. monanthemum subsp. filicaule. This latter subspecies then shows a reduction trend in the opposite direction through Xizang and ending in Yunnan. North-eastward from northern Yunnan and through south-eastern Sichuan subsp. monanthemum remains almost uni- form, but there is a distinct morphological gap between it and the delicate derivative 2. H. subcordatum, which extends south in area from its centre in southern Shaanxi as far as Emei Shan. The second clade, clade v, originates in a species from a small area in north-west Yunnan, 3. H. trigonum, which is morphologically very similar to the form of 1 . H. monanthemum from nearby. It has given rise to two species. To the south of the main Himalayan range, 4. H. himalaicum occurs in the Arunachal Pradesh-Myanmar fron- tier area, then recurs in central Nepal, whence it shows trends both westward to Pakistan and eastward to Sikkim, then through the Chumbi gap into Xizang and thence into Yunnan. To the north of the main range, 5. H. ludlowii shows a reduction trend in size from north-western Yunnan through Xizang into Bhutan. la Siberia, Japan, Korea China, Taiwan, Vietnam 6 N. Yunnan S Japan, C & S. China Taiwan, Vietnam E. Myanmar 1 / C & SE China 2 Meghalaya 2a C. & SW China 2b SW China N. Myanmar to Nepal SW China, N. Laos N. Thailand, N. Myanmar NE India, Bhutan | S India, Sri Lanka 3 4a W. Sichuan to Nepal \ Ib Nepal to SE Xizang & NW Yunnan NW Yunnan Yunnan, Xizang, Bhutan S China & W Myanmar to S. Xizang & Kashmir 4b NE India, C. Myanmar V 4 SW China to Pakistan W. Sichuan to S. Shaanxi SW China, N. Myanmar N. Assam Fig. 7 Sections 9c-9e. Distribution of species, showing minor disjunction (I). For names of sections and species see Fig. 5 (p. 46). STUDIES IN HYPERICUM 49 Finally, clade vi originates in 6. H. daliense, a relict endemic of the Dali and Lijiang ranges of northern Yunnan. It is closely related (ancestral?) to the widespread 7. H. wightianum, of which the most similar form to H. daliense occurs in the Khasi Hills (Meghalaya). From there reduction trends are found: (a) westward to Sikkim; (b) eastward into China as far as Guanxi, Guizhou and Sichuan; (c) southward into Nagaland and Mizoram (eastern India) and, with a large disjunction, in Tamil Nadu and Sri Lanka; (d) from northern Myanmar into central Myanmar and northern Thailand. SYSTEMATIC TREATMENT 3 Sect. 3. ASCYREIA Choisy - an additional species (see p. 40). 16a. Hypericum elatoides R. Keller in Bot. Jb. 33: 549 (1904); H. Leveille in Bull. Soc. Bot. France 54: 588 ['helatoides'], 590 (1908); N. Robson in Bull. Br. Mus. nat. Hist. (Bot.) 5: 320 (1977) sub sect. Roscyna; Li Xiwen in Fl. R. P. Sinicae 50(2): 71 (1990). Type: China, Shensi [Shaanxi] settentr., Ki-san, May 1895 (fl), Scallqn in Biondi 3822 (FI!-holotype). Fig. 8A, Map 1 . H. monogynum var. franchetii Baroni [Enum. Sem. in Horto hot. Florent. anno 1898 coll.: 36 (1898?)] in Boll. Soc. Bot. Ital. 1898: 185 (1898), Diagn. PL Sin. Nov.: 35 (1898). Type: Cultivated in Hort. bot. Florentine ex China, Shaanxi, 1894 ex Giraldi, July 1895 (fl), Herb. Biondi 3809 (Fll-lectotype, selected here). H. chinense sensu R. Keller in Bot. Jb. 33: 548 (1904) pro parte quoad specim. cit. H. ascyron var. punctato-striatum R. Keller in Bot. Jb. 33: 550 (1904); H. Leveille in Bull. Soc. Bot. France 54: 592 (1908). Type: China, Shen-si [Shaanxi], presso la cima di In-kia-po, 4 June 1897 (fl), Giraldi in Biondi 3816 (Fll-lectotype, selected here). H. ascyron var. micropetalum sensu R. Keller in Bot. Jb. 33: 550 (1904) pro parte quoad specim. cit.; H. Leveille in Bull. Soc. Bot. France 54: 592 (1908). Suffrutex 0.35-0.82 m tall, with stems solitary or few, usually slender, erect from ascending or creeping, branching and rooting base, unbranched below . inflorescence. Stems shallowly 4-lined below nodes, sometimes 2-lined below or becoming almost terete, eglandular; intemodes (30-)45-105 mm, shorter than or equalling leaves. Leaves sessile or with petiole to 1.5 mm; lamina 44-110 x 18-50 mm, oblong or oblong-triangular to broadly ovate, rather paler beneath, not glaucous, plane, thinly chartaceous; apex obtuse to shortly apiculate or rounded, margin entire, base cordate- amplexicaul to truncate; venation: 4-7 pairs of main lateral veins from lower half of midrib, tertiary reticulation dense, prominent below; laminar glands very small, striiform to punctiform; intramarginal glands dense. Inflorescence (l-)5-13-flowered, from 1-2 nodes, rarely with flowering branches from one node below, the whole laxly corymbiform; pedicels (3-) 10-50 mm; bracts and bracteoles 7-17 mm long, linear-lanceolate to linear-elliptic. Flow- ers 40-80 mm in diam., stellate, with petals becoming reflexed, tardily deciduous; buds narrowly ovoid, acute. Sepals 5, free or to 0.25 united, slightly imbricate, 3-5(-7) x 1.5-3(-5) mm, ovate to triangular-ovate, obtuse to acute, entire or occasionally minutely irregularly glandular-ciliate, erect to suberect in bud and fruit; veins 5, branched; laminar glands linear (inner) or interrupted (outer); marginal glands reddish or absent. Petals 5, golden? yellow, 2540 x 7-10(-15) mm, 6-8 x sepals, oblanceolate, without apiculus, margin entire; laminar glands linear to distally striiform. Stamen fascicles 5, each with c. 45-60 stamens, longest 15-25 mm long, 0.5-0.65 x petals; anthers yellow, gland amber. Ovary incompletely 5-locular, 46(-7) x 3.5^.5 mm, broadly ovoid; styles 5, 13-17 mm, 2.3-4.5 x ovary, slender, united almost to apices; stigmas small. Capsule 9-16 x 6-11 mm, broadly to narrowly ovoid or ovoid-conic, 3-4 x sepals, evittate. Seeds dark reddish brown, 1.1- 1.5 mm, cylindric, without terminal appendage, very shallowly carinate; testa densely shallowly reticulate. 2n = ? Damp places in open grassland; 770-1000 m. China (Henan, Shaanxi, Gansu). CHINA. Henan: Neixiang Xian, Baotianman Nat. Res., Xuayao Gou, 850- 950 m, 22 May 1994 (fl), Boufford et al. 26156 (BM, GH*). Shaanxi: Danfeng Xian, Yaozhuang, Dadongqiu, 1000 m, 20 September 1958 (fr), GuoB.Z. 3873 (BM-photo, IBSC, KUN)); Tai-pei-shan [Taibaishan], 1910 (fl), Purdom 893 (A*, E, K, US); Tsincheng-hsien [Xinchengbu? Xian], Taihangshan, 820 m, 19 June 1937 (e. fr), Liou 7343 (BM - photo, PE); Lao- y-san presso Zu-lu, 6 September 1897 (fr), Giraldi in Biondi 3818 (Fl; K). Gansu: Tangye to Wushang Xian, 3 June 1956 (fl), Huang He Exped. 4465 (BM - photo, PE). H. elatoides is most closely related to H. monogynum, in particu- lar to an ovate- to oblong-leaved form from Jiangsu (e.g. Wilson 1604). It differs from Form III of//, monogynum (e.g. Wilson 2420, from west Hubei), by the suffruticose habit, the larger thinner leaves, the usually 2-noded inflorescence, the relatively short and broad sepals with a thinner, usually glandular-ciliate margin, the tardily deciduous exapiculate petals and the more numerous, tardily deciduous stamens. It is rather similar to H. prattii N. Robson (from Sichuan), but differs from it by the obtuse to rounded (not acute to apiculate) relatively narrow leaves and the small triangular-ovate, usually minutely reddish-glandular-ciliate sepals. In H. prattii the sepals are large and ovate-lanceolate to elliptic with an entire margin. Sect. 8. BUPLEUROIDES Stef. in God. Agr.-les. Fak. Univ. Sofiyall: 160(1933). Perennial herbs up to 0.75 m tall, with stems erect from branching rhizome, glabrous, without dark glands or, if present, then small, reddish or black, and marginal only. Stems incompletely 2-lined or usually terete, eglandular. Leaves opposite, perfoliate, persistent; lamina entire with venation pinnate, closed, the tertiary reticulation dense; laminar glands pale, small, punctiform, scattered; marginal gland dots irregular, small; ventral resin glands absent. Inflores- cence 4-25-flowered with branching dichasial/monochasial from 1-5 nodes, without subsidiary branches; bracts and bracteoles per- sistent. Flowers stellate or with petals reflexed, homostylous. Sepals 5, free, persistent, erect in bud and fruit, with margin entire or minutely glandular; veins 5; laminar glands pale, linear or striiform to sometimes punctiform; marginal glands reddish to black or absent. Petals 5, persistent, without apiculus, entire or usually with minute sessile, reddish to black marginal glands; laminar glands pale, numerous, striiform to punctiform. Stamen fascicles 4 (united 2+1+1+1) or usually 3 (united 2+2+1), persistent, the single ones 3 An asterisk (*) before a locality or after a herbarium symbol indicates that the specimen has not been seen by me. Type material seen by me is indicated by an exclamation mark 50 N.K.B. ROBSON each with c. 20-25? stamens; filaments united at base only; anthers yellow, gland amber; pollen type IV. Ovary with 3(4) placentae, wholly united, <-ovulate; styles 3(4), free, appressed below; stigmas small. Capsule 3(4)-valved, coriaceous, with valves narrowly longi- tudinally vittate. Seeds curved-cylindric, not or slightly carinate, without apical expansion; testa shallowly and minutely foveolate. BASIC CHROMOSOME NUMBER (x). Unknown. HABITAT. Damp places in forest; 1000-2100 m. DISTRIBUTION. North-east Turkey, Georgia. 1 . Hypericum bupleuroides Griseb. in Wiegm., Arch. Naturgesch. (Berlin) 18(1): 299, in obs. (1852); Boiss., Fl orient. 1: 809 (1867); Woronow in Kuzn., Busch & Fomin, Fl. Caucasica Critica 3, No. 9: 47 (1906); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: t. 1 f. 11 (1932), 11: 160 (1933), 12: 84 (1934), in Pflanzenareale 4: karte 2b (1933); Gorschkova in Shishkin & Bobrov, Fl. URSS 15: 246 (1949); Grossgeim, Fl. Kavk. 2nd ed. 6: 174, map 193 (1962); N. Robson in Davis, Fl. Turkey 2: 367 (1967). Type as for H. perfoliatum Ledeb. Fig. 8B, Map 2. H. perfoliatum Ledeb. in Bull. Acad. Sci. St. Petersb. 2: 314 (1837), Fl. Ross. 1: 445 (1842), non L. (1767). Type: Georgia, auf dem Hochplateau Gor-Somlia zwischen der provinz Guriel und Adschara, 1500-2100 m, 1836 (fl), von Nordmann s.n. (LE- lectotype & syntype; H!). There are two Helsinki specimens: (i) Mont Adzhar, Guriel (isolectotype, selected here) and (ii) Akhalziki [Achaltsikhe] (isosyntype). Perennial herb, 0.45-0.8 m tall, with stems solitary or few, erect from creeping and rooting base, unbranched below inflorescence. Stems incompletely 2-lined or usually terete, eglandular; internodes 50-1 20 mm, shorter to longer than leaves. Leaves perfoliate, with or without shallow sinus at points of union; lamina 70-210(280) x 37-80(-100) mm, ovate to elliptic-ovate or oblong-ovate, paler beneath, not glaucous, plane, papyraceous; apex obtuse to apiculate, margin entire, bases united in pairs; venation: 4 pairs of main lateral veins from lower third or so of midrib, tertiary reticulation dense, only main veins prominent beneath; laminar glands pale, very small, scattered; marginal glands pale, small, irregular. Inflorescence 4- 1 1 (-25)-flowered, from 1-3 nodes, without flowering branches, the whole broadly pyramidal; pedicels 1 8-25 mm; bracts and bracteoles 0.5-4 mm long, lanceolate, withering and falling, leaving scale-like base. Flowers 21-40 mm in diam., stellate, with petals becoming reflexed; buds narrowly ovoid?, rounded? Sepals free, narrowly imbricate, unequal, 2.5-4.5 x 2-2.3 mm, elliptic-oblong to ovate, rounded, entire or with minute sessile glands, erect in bud and fruit; veins 5, distally branched; laminar glands linear to striiform or distally punctiform; marginal glands reddish to black or absent. Petals 15-20 x 2.5-5? mm, c. 8 x sepals, narrowly oblanceolate, without apiculus, entire or usually with minute sessile glands; laminar glands pale, striiform to punctiform; marginal glands red- dish to black. Stamens 50-75, longest 14-17 mm long, c. 0.75-0.85 x petals; anther gland amber. Ovary (3.5?-)6-7 x 3 mm, ovoid- conic; styles 12-14 mm, 2-3 x ovary, slender; stigmas small. Capsule 9-14 x 5-8 mm, ellipsoid to ovoid, c. 3.5 x sepals, narrowly longitudinally vittate. Seeds reddish brown, 1.2-1.5 mm, cylin- dric, slightly curved, without terminal appendage, not or slightly carinate; testa minutely linear-foveolate. 2n = ? Mountain woods and damp valleys, wooded slopes; 640-2100 m. North-east Turkey (Rize, Coruh), Georgia. TURKEY. Rize: Hemsin Distr., Meydan Kobaca to Mollaveysa, 1000 m, 3 September 1952 (1. fl), Davis & Dodds D21372 (BM, E, K); Ikizdere nach Camlik, 1200 m, 13 July 1958 (fl), Huber-Morath 14974 (BASBG); gorges 1 km below Khabakhor (Lazistan), c. 1 000 m, 27 August 1 866 (fr), Balansa 89 (E, G, JE, K). oruh: inter Salaket et Khinzart, c. 1 350 m, 6 August 1911 (fl), Woronow 5670 (JE, K, S); Savval Tepe above Murgul, 1600 m (to 1900 m), 1 1 August 1957 (1. fl), Davis & Hedge D.32223 (BM, E, K). GEORGIA. Adzharskaya: basin of R. Kableani, Kishlak village, August 1948 (fl), Sochava & Naumova s.n. (LE); Batumi, Mtirala Mtn, 640 m, 14 July 1975 (1. fl), Whitmore 3 100 (BM). Georgia: Mt. Imeretia inter Kutais et Achalzich, n.d., Ruprecht s.n. (G). H. bupleuroides is an isolated Tertiary relict (Woronov, 1906: 48) of which the nearest relative is in central China (H. elatoides, sect. Ascyreia; see p. 40) and provides a morphological link between it and H. monogynum. H. xylosteifolium (Spach) N. Robson (sect. 6. C ! C Map 2 Sect. 8: 1. H. bupleuroides specimens, other records O. STUDIES IN HYPER/CUM 51 Fig. 8 A. H. elatoides: (a) habit; (b) sepal; (c) petal; (d) ovary; (e) capsule. B. H. bupleuroides: (f) habit; (g) sepal; (h) petal; (i) ovary; (j) capsule (a, f x 21 3; c-e. h-j x 2; b, g x 4). A. (a), (d) Purdom 893; (c) Wang 3694; (e) Fr. Hugh s. n. B. (f)-(i) Whitmore 3100; (j) Davis & Hedge 32223. 52 N.K.B. ROBSON Inodora), which provides another example of this disjunction be- tween central China and the south-east Black Sea region, is also a Tertiary relict with no close relatives, its nearest one being the variable H. monogynum (see Robson, 1985: 314). H, bupleuroides differs from H. elatoides in its herbaceous habit, perfoliate leaves, pseudo-trimery in the androecium and trimery in the gynoecium. In addition, the styles are only loosely appressed in the lower half, not almost completely united. Sect. 7. ROSCYNA (Spach) R. Keller in Engler & Prantl, Nat. Pflanzenfam. 3(6): 211 (1893); N. Robson in Bull. Br. Mus. not. Hist. (Bot.) 5: 319 (1977) pro parte, excl. H. elatoides R. Keller. Perennial herbs up to 2 m tall, with stems erect or sometimes basally ascending from taproot, glabrous, without dark glands; branching lateral, confined to inflorescence or from up to all nodes. Stems 42- lined when young, becoming narrowly 4-winged to terete, eglandular. Leaves opposite, decussate, sessile to subsessile, free, persistent; lamina entire with venation pinnate, closed sometimes except lower 1-2 pairs, with tertiary reticulation dense, sometimes obscure; laminar glands pale, punctiform or shortly striiform, unequal; mar- ginal gland dots dense, very small; ventral resin glands absent. Inflorescence 1-c. 35-flowered with branching dichasial/ monochasial from 1-7 nodes, often with subsidiary branches from lower nodes; bracts and bracteoles foliar or reduced, deciduous or persistent. Flowers stellate, homostylous. Sepals 5, free or united at base, persistent, spreading to deflexed in fruit, with margin entire; veins 5- 1 7; laminar glands linear and distally interrupted to striiform or wholly punctiform; intramarginal glands few or absent. Petals 5, persistent and becoming reflexed, with apiculus present, lateral, or absent; margin entire; marginal glands absent; laminar glands linear, sometimes interrupted distally, numerous, or rarely absent. Stamen fascicles 5, free (or 4?, united 2+1 + 1 + 1), persistent, each single fascicle with c. 15-30 stamens; filaments united at base only; anthers yellow, gland amber; pollen type III. Ovary with central lacuna and 5(4) axile placentae, -ovulate; styles 5(4), partly united or partly coherent or free; stigma narrowly to broadly capitate or infundibuliform. Capsule 5(4)-valved, coriaceous, with valves nar- rowly longitudinally vittate. Seeds cylindric, narrowly carinate to narrowly winged, sometimes with slight apical expansion; testa densely and shallowly linear-reticulate. BASIC CHROMOSOME NUMBER (x). 11-10? (see p. 44), 9, 8; ploidy 2. HABITAT. Moist to dry meadows or grassy or rocky slopes, in open forest or among scrub or along streamsides and river banks, also along roadsides; 0-4000 m. DISTRIBUTION. Siberia (Altai to Kamchatka and Kurile Is., Sakhalin), Mongolia, Korea, China (all provinces except Xizang and Qinghai), Vietnam (north), Taiwan, Japan; Canada (south-east), U.S.A. (north-east). 2 species (+ 2 subspecies). Key to sect. 7. Roscyna 1 Leaf apex acute to obtuse, lamina mostly lanceolate to elliptic-oblong or linear; mature stem internodes sharply 4-lined to narrowly 4-winged ( 1 . ascyron) 2 Leaf apex rounded to shallowly retuse, lamina oblong to oblong- lanceolate or oblong-ovate or triangular-oblong; mature stem internodes partially 2-lined or terete 2. przewalskii 2(1) Capsule ovoid to ovoid-pyramidal orovoid-cylindric; sepals 6-13 mm wide, ovate to lanceolate or oblong or elliptic; styles 4-15 mm, c. 0.7- 2 x ovary, almost free to up to 0.8 united; leaves c. 7-40 mm wide, base cordate to rounded 3 Capsule cylindric-ellipsoid; sepals 1 .5-7 mm wide, narrowly oblong or oblong-lanceolate; styles 2.5-3.5 mm, c. 0.5 x ovary, free; leaves 4-15 mm wide, base usually cuneate Ib. ascyron subsp. gebleri 3(2) Sepals rounded to obtuse or rarely acute; styles usually united la. ascyron subsp. ascyron Sepals acute to shortly acuminate; styles usually almost free Ic. ascyron subsp. pyramidatum 1. Hypericum ascyron L., Sp. pi.: 783 (1753) excl. syn. Moris, et Wheeler, et Ray. et loc. Oriente; Murray, Syst. veg. 1 3th ed.: 583 (1774); Lam., Encycl. 4: 147 (1796); Willd., Sp. pi. 3: 1443 (1802); Choisy, Prodr. monogr. Hyperic.: 41 (1821), in DC., Prodr. 1: 545 (1824); Sprengel, Syst. Veg. 3: 342 (1826); G. Don, Gen. Syst.: 602 (1831); Ledeb., Fl. altaic. 3: 363 (1831), Fl. ross. 1: 446 (1842); Turcz., Fl. Baic.-Dah. 1: 621 (1843); Maxim., Prim.fl. amur.: 64 (1859); Regel, Fl. ussur.: 32 (1861); Trev., Hyperic. animadv.: 8 ( 1 861 ); Miquel in Ann. Mus. Bot. Lugduno- Batavum 2: 250 (1866); Franchet & Sav., Enum. PI. Jap. 1: 55 (1874); Tanaka in Inuma, Somoku-Zusetsu, 2nd ed., 14: t. 33 (1874); Hance in / Bot., Lond. 18: 259 (1880); Maxim, in Bull. Acad. Imp. Sci. Saint-Petersbourg 27: 430 (1882), Mel. Biol. 11: 163 (1882); Franchet in Nouv. Arch. Mus. Hist. Nat., 2 e sen, 5: 207 ( 1 883), PL Da v. : 55 ( 1 884), in Bull. Soc. Bot. France 33: 436 (1886); Hemsley in /. Linn Soc. 23: 72 (1886); Franchet, PL delavay.: 112 (1889); Korschinsky inActaHort. Petrop. 12: 317 (1892); Goeschke in Gartenflora 41: 537, t. 1381 (1892); R. Keller in Bull. Herb. Boiss. 5: 638 (1897); Diels in Bot. Jb. 29: 476 (1900); Krylov., Fl. Altaya 1: 188 (1901); Komarov, Fl. Man 'chzh. 3: 40 (1905); Matsum. & Hayata, Enum. pl.formosa: 40 (1906); H. Leveille in Bull. Soc. Bot. France 53: 498 (1906), op. cit. 54: 592 (1908); Pavolini in Nuovo Giorn. Bot. Ital.,n. ser. 15: 406 (1908); Nakai, Fl. kor. 1: 95 (1909); R. Keller in Bot. Jb. 44: 48 (1909); Hayata, Icon, pl.formos. 1: 77 (1911); Matsum., Index pi. jap. 2(2): 364 (1912); H. Leveille, Fl. Kouy-Tcheou: 198 (1914); Sprague in Curtis's Bot. Mag. 140: t. 8557 (1914); Rehder in Sargent, PL wilson. 2(2): 402 (1915); House, Wildfl. New York: t. 131 A (1918-20); R. Keller in Engler & Prantl, Nat. Pflanzenfam. 2nd ed., 21: 176 (1925); Makino & Nemoto, Fl. Japan: 540 ( 1 925), 2nd ed. : 747 ( 1 93 1 ); Hulten in Kungl. Svenska Vetenskapsakad. Handi, 3rd ser. 8: 125 (1929); Hand.-Mazz., Symb. Sin. 7: 401 (1931); Komarov & Klob.-Alis., Opred. r. Dal'nevostkr. 2: 748, t. 230, f. 2 (1932); Nakai in Koryo-sikenrin no Ippan: 46 (1932); Krylov, Fl. Zap. Sib. 8: 1902 (1935); S. Suzuki in Masamune, Short fl. Formosa: 41 (1936); Kitag., Lin. fl. manshur.: 317 (1939); Nakai in J. Jap. Bot. 18: 289 (1942); Gorschkova in Fl. URSS 15: 211, t. 9 f. 2 (1949); Y. Kimura in Nakai & Honda, Novafl. jap. 10: 1 1 8, ff. 43, 44 (195 1 ); Grubov, Konsp. Fl. Mongol. Narod. Respub.: 205 (1955); Steward, Man. Vase. Pis Lower Yangtze Valley China: 258 (1958); Ohwi, Fl. Japan, Engl. ed.: 631 (1965); Lauener in Notes Roy. Bot. Card. Edinburgh 27: 1 (1966); Noda, Fl. N.-E. Prov. (Manchuria) China: 793, t. 190 f. 3 (1971); Anon, in Iconogr. Cormoph. Sinicae 2: 875, f. 3479 (1972); N. Robson in Fl. Taiwan 2: 628 ( 1 976), 2nd ed. 2: 700 ( 1 996); Y. Kimura in Asahi . . . shokobutsu, no. 64: 1509, cum tab. (1977); Liou Tchen-ngo et al. in Fl. PI. Herbac. Chinae Bor.-Or. 6: 71, t. 27 ff. 1-7 (1977); Kitag., Neo- lin.fl. manshur.: 443 (1979); Wu Chingju in Fl. Intramongolica 4: 90, t. 42 ff. 4-7 (1979); Momiyama in Satake et al., Wildfl. STUDIES IN HYPERICUM 53 Japan 2: 1 1 5, 1. 1 10 ( 1982); Grubov, Opred. Sosud. Rast. Mongolii: 181, t. 97 f. 443 (1982); Hayashi, Azegami & Hishiyama, Wild/I. Japan: 358 (1983); Anon, in Fl. in Desertis R. P. Sinicorum 2: 365, t. 130 ff. 1-4 (1987); Wang Jinwu in Fl. Hebeiensis 2: 149, 1. 1051 (1988); T. B. Lee, ///. Fl. kor. : 544 (1989); Li Xiwen in Fl. R. P. Sinicae 50(2): 43, t. 8 ff. 1-3 (1990); N. Robson in RHS Diet. Card. 2: 627 (1992), in Eur. Gdn Fl. 4: 58, ff. 9.1, 9.5, 9.9 (1995); Y. N. Lee, Fl. kor. 3rd ed.: 230, f. 677 (1998). Type: cult, in Hort. Upsaliensis ex Siberia, Herb. Linn. 943.9 (LINN- lectotype, selected here). The Linnaean references to Morison, Wheeler and Ray, and to the Orient, all apply to H. ascyron Mill., which is H. calycinutn L. Fig. 9, Maps 3 & 4. H.fl. pentagynis, caule tetragono (etc.) Gmelin, Fl. Sibir. 4: 178, t. 69(1769). Ascyrum tetragonum Moench, Meth. Bot.: 130 (1794), nom. illegit. Type as for H. ascyron L. Roscyna gmelinii Spach, Hist. not. veg. Phan. 5: 430 (1836), in Ann. Sci. Nat. Bot. II, 5: 364 (1836), [gmelini], nom. illegit. Type as for H. ascyron L. H. ascyron var. genuinum Maxim., Prim. Fl. amur:. 65 (1859); in Bull. Acad. Imp. Sci. Saint- Petersbourg 27: 430 (1882). Type as for H. ascyron L. H. ascyron var. typicum R. Keller in Bot. Jb. 58: 191 (1923). Type as for H. ascyron L. Roscyna ascyron (L.) Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 12(1951), in synon. Icones: see under infraspecific taxa (pp. 56-58). Perennial herb 0.5-1. 3(-2) mm tall, erect or sometimes ascending from shortly creeping woody base, with stems single or few, caespitose, unbranched or branched above or almost throughout. Stems 4-angled when young, becoming 4-lined or occasionally internodes 2-lined below; internodes 2-12 mm, exceeding leaves or shorter than them. Leaves sessile; lamina (30-)40-97(-120) x (4-) 7_35(^0) mm ovate-lanceolate or narrowly lanceolate or nar- rowly oblong or narrowly elliptic to oblong-linear or oblanceolate, rather paler beneath, not glaucous, plane, chartaceous; apex acute to subapiculate or obtuse (or lowermost rarely rounded), margin entire, base cuneate to cordate-amplexicaul; venation: 4-7 pairs of main laterals from lower half of midrib, with subsidiary midrib branches and dense tertiary reticulation not prominent, often obscure; laminar glands pale, dense, unequal dots or short streaks; intramarginal glands pale, small, dense. Inflorescence \-c. 35-flowered from 1-5 nodes, the whole subcorymbiform to narrowly pyramidal, some- times with flowering branches from up to 4 nodes below; pedicels 5-30 mm in fruit; bracts and bracteoles foliar but smaller and often broader, more rarely linear-lanceolate and deciduous. Flowers 30- 70(-80) mm in diam., stellate with petals spreading to reflexed; buds broadly to narrowly ovoid, rounded to subacute. Sepals (3-)5-15 x (1.5-)2-7(-10) mm, free, imbricate, subequal to unequal, the outer ones sometimes foliaceous, erect in bud and fruit, oblong to elliptic or ovate to ovate-lanceolate or obovate, rounded to obtuse or more rarely subacuminate to acute, entire; veins c. 11-17, branching and uniting distally, midrib scarcely differentiated; laminar glands lin- ear, distally interrupted to striiform; marginal glands spaced, small. Petals bright (to golden?) yellow, sometimes tinged red in bud, 14- 41 x 5-20 mm, 2-3 x sepals or relatively shorter when sepals foliaceous, obovate or oblong-obovate to oblanceolate, often some- what spathulate to subunguiculate, strongly curved to almost straight, rounded or obtuse to rarely acute or acuminate, with apiculus short and rounded or absent, margin entire; laminar glands pale, linear to distally striiform, or absent; marginal glands absent. Stamen fasci- cles 5(4?), distally red, each with c. 30 stamens, longest 9-25 mm, c. 0.4-0.67 x petals; anther reddish, gland amber. Ovary 5(4)- locular, 4-7(-9) x 3-5 mm, broadly ovoid to narrowly ovoid-pyramidal or ellipsoid; styles 5(4), 2.5-15 mm, c. 0.5-2 x ovary, free or up to 0.8 coherent or connate; stigmas broadly capitate to infundibuliform. Capsule 9-22(-30) x 5-13 mm, broadly to narrowly ovoid or ovoid-pyramidal or rarely narrowly cylindric, 2- 3 x sepals, obtuse to rounded, with numerous narrow longitudinal vittae. Seeds dark red-brown, 1-1.5 mm, cylindric, not or slightly curved, deeply carinate or narrowly winged, sometimes with slight terminal expansion; testa densely shallowly linear-reticulate. 2n = c. 22-20 (Krasnoborov et al., 1980), 18 (Kogi, 1984; Malakhova, 1990; Nishikawa, 1990; Malakhova & Markova, 1994; Stepanov, 1994; '//. getter? n = 9, Nielsen, 1924), 16 (Krogulevich, 1978). Moist to dry meadows or grassy or rocky slopes, sometimes in forest or amongst scrub, streamsides and river banks; 0-2800(-3600) m. Russian Federation (Altai to Kamchatka and Kurile Is., Sakhalin), Mongolia, Korea, Japan, Taiwan, China (all provinces except Xizang and Qinghai), Vietnam (north); Canada (Quebec, Ontario, Mani- toba?), U.S.A. (Minnesota to Vermont and south to Missouri, Illinois and Maryland). H. ascyron is a highly polymorphic species or species complex with a very wide distribution. Although several varieties or even species have been recognized, the variation appears to be almost continu- ous. Seven characters or character groups appear to vary independently or with insufficient correlation or disjunction to be useful for specific delimitation: (1) Habit/inflorescence: from (a) simple, unbranched, 3-flowered to (b) pyramidally branched with flowers on each branch or (c) widely corymbiform ('var. umbellatum 1 }. (2) Leaves: from (a) large, triangular-lanceolate, acute to (b) small, narrowly oblong to linear, acute or (c) oblong, obtuse to (upper) very rarely almost rounded or (d) rather broadly elliptic and subacute ('H. hemsleyanuni') to almost rounded. (3) Flowers: from large to small, varying approximately with (1) and (2) except for (a) an extra-large-flowered form with long, mostly united styles ('var. longistylum" in part) and (b) Man- churian plants with large flowers and small leaves. (4) Sepals: from (a) oblong or oblong-elliptic, rounded to (b) narrowly oblong, rounded (subsp. gebleri) or (c) ovate, acute (subsp. pyramidatum) or (d) obovate to foliaceous, rounded (var. macrosepalum) or (e) ovate-lanceolate. (5) Styles: from (a) medium long and up to 0.75 united to (b) long, c. 0.8 connate (see (3) above) or (c) shorter than ovary and free (mostly subsp. gebleri). (5) varies with (4) in general but not completely. Some small-flowered, small-leaved plants from south China and Japan ('forma angustifoliurn) have styles about half connate, while some large-flowered plants with broad sepals have short, free styles (Nei Mongol). (6) Capsules: from (a) large ovoid-pyramidal to (b) small ovoid- cylindric (subsp. gebleri) or (c) narrowly cylindric ('var. giraldi?). (7) In addition, the pedicels vary in length from c. 10 mm to (a) 30 mm or (b) c. 5 mm ('var. umbellatum). The most frequently distinguished taxon has been H. gebleri Ledeb., which most Russian authors have recognized, as it occurs through southern Siberia from Altai to the Korean border along with distinct larger-flowered less-branched forms and is present alone in Kamchatka, the Kuriles and Sakhalin. On the other hand, Chinese 54 N.K.B. ROBSON authors have tended to include it in H. ascyron, because intermediate forms occur in the north of that country. The North American plants were early recognized at species level (H. pvramidatum, H. macrocarpum, etc.) and Kimura (195 1) main- tained them as var. americanum partly on the basis of their rather acute sepals. Already in 1882, however, Maximowicz (1882: 163) and slightly later Coulter ( 1 886: 83) had expressed the opinion that they could not be distinguished from the Asian plants by this or any of the other differentiating characters that had been proposed. I can confirm that two of Kimura's other characters - leaf shape and styles shortly connate - do not help to define a discrete taxon. The American plants, however, all have short styles and most have somewhat acute sepals, a combination that does not occur elsewhere in the species. Kimura's other varieties were based mainly on these style length and union criteria: var. longistylum with styles up to 15 mm, 2/3 to 3/4 united, var. ascyron with styles up to 7 mm, 1/3 to 21 3 united (both with styles equal to or longer than the ovary), and var. brevistylum with styles shorter than the ovary and free. In these circumstances it seems best to distinguish the northern Asian H. gebleri and the North American H. pyramidatum as subspecies (although further investigation may show that the inter- mediate forms in the former are too numerous to maintain it even at this rank). It seems impossible to recognize the long-styled (mainly large-flowered and northern) plants and the narrow-fruited central Chinese ones (respectively vars longistylum and giraldii (~ var. hupehensis)) as distinct taxa, as both represent extremes of continu- ous variation. Likewise, the narrow-leaved small-flowered form from south China (wrongly identified by me and others as var. angustifolium Y. Kimura) merges with more typical forms. la. Hypericum ascyron subsp. ascyron Fig. 9 A, Map 3. Ascyrum sibiricum Lam. ex Poiret, Tabl. encycl. 3: 200, t. 642 (1823). Type as for H. ascyron L. ?H. salicaria Rchb., Iconogr. Bot., cent. 5: 53, t. 490, f. 183 (1827). Type: cult, ex Kazakhstan, 'ad Noor-Saisan in mont. Alt.', L[edebour] s.n. (W?-holotype). See note under subsp. gebleri. H. ascyron var. macrosepalum Ledeb., Fl. altaic. 3: 364 (1831), Fl. ross. 1: 446 (1842); Krylov, Fl Altaya: 189 (1901), Fl Zap. Sib. 8: 1903 (1935). Type: Russian Federation, W. Siberia, Altayskiy Kray, 'prope pagum Uimon ad fl. Katunja'. B[unge] (LE). Roscyna japonica Blume, Mus. bat. 2: 21 (1856). Type: Japan, Herb, von Siebold (L-holotype). H. ascyron var. [|3] longistylum Maxim., Prim.fl. amur.: 65 (1859), in Bull. Acad. Imp. Sci. Saint-Peter -sbourg 27: 430 (1882), Mel. Biol. 11: 162 (1882); Palibin, Consp.fl. Koreae 1: 44 (1898); H. Leveille in Bull. Soc. Bot. France 53: 499 ( 1 906); Nakai in J. Coll. Sci. Imp. Univ. Tokyo 26: 96 ( 1 909) [Fl. Korea], Tyosen-syokubutu 1: 159, f. 190 (1914); Matsum., Index pi jap. 2: 364 (1912); Koidz., Fl. Symb. Orient.- Asiat.: 91 (1930) pro syn. H. sagittifolii ut forma; Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 125, f. 44C (1951); Ohwi, FL Japan, Engl. ed.: 631 (1965); Liou Tchen- ngo et al. in Fl. pis. Herbac. Chinae Bor.-Or. 6: 71, t. 27 f. 8 (1977); Kitag., Neo-lin. fl. manshur.: 443 (1979); Y. N. Lee, Fl. kor. 3rd ed.: 230, f. 678 (1998). Type: Russian Federation (east Siberia), Chochzier ad Amurem non procul ab Ussuri ostio, Maximowicz s.n. (LE-holotype). H. ascyron var. micropetalum R. Keller in Bull. Herb. Boiss. 5: 638 (1897) ['forma micropetalum var. nov.'], in Bot. Jb. 33: 550 (1904) pro parte excl. specim. cit., in Bot. Jb. 58: 191 (1923); H. Leveille in Bull. Soc. Bot. France 53: 499 (1906), op. cit. 54: 592 (1908) pro parte excl. specim. cit.; Matsum., Index pi jap. 2: 364 (1912); Makino & Nemoto, Fl. Japan: 540 (1925), 2nd ed.: 747 (1931). Type: Japan, in planitie Sambingi, Faurie 1885 (Z- lectotype, selected here); prope Nuruyu, Faurie 793 (Z-syntype). H. longifolium H. Leveille in Bull Soc. Agric. Sarthe 39: 322 (1904), in Bull Soc. Bot. France. 54: 590 (1908), in Repert. Spec. Nov. Regni Veg. 6: 375 (1909), Fl. Kouy-Tcheou: 199 (1914) pro parte quoad Esquirol 1461; Rehder in J. Arnold Arbor. 15: 191 ( 1 934). Type: China, Guizhou, pres Ke-ma-tong, mont. du College, 9 August 1897 (fl & fr), Bodinier 1774 (E!-holotype). H. scallanii R. Keller in Bot. Jb. 33: 549 (1904); H. Leveille in Bull. Soc. Bot. France 54: 590 (1908), in Fl Kouy-Tcheou: 199 (1914). Type: China, Sichuan, in monte Uo-mi-san [Emei shan] prope Tcen-to-sen, September 1899 (fl), Scallan in Biondi 3808 (FI!- holotype). H. ascyron var. giraldii R. Keller in Bot. Jb. 33: 550 (1904); H. Leveille in Bull Soc. Bot. France 54: 592 (1908). Type: China, Shaanxi (all), in alto monte Huan-tou-san, July 1 889 (fl), Giraldi in Biondi 3805 (Fl l-lectotype selected here); in rupestribus montis Kian-san prope Sce-kin-tsuen, 4 August 1897 (fl), Giraldi in Biondi 3803 (FI!-syntype); Lao-y-san, Thin-kio-tsuen, July 1899 (fl), Giraldi in Biondi 3804 (FI!-syntype); in occidentem montis Ngo-san, August 1 899 (fl), Giraldi in Biondi 3806 (FI!-syntype); in monte Uo-mi-san prope Tcen-to-san, Scallan in Biondi 3807 (FI!-syntype). H. biondii R. Keller in Bot. Jb. 33: 551 (1904) pro parte quoad lectotypus. Type: China, Shaanxi, Monte di Mang-hua-san ad ovest di Si-ngan-fu e distante tre giornale, October-November 1894 (fr), Giraldi in Biondi 530 (Fll-lectotype, selected here); in monte Thae-pei-san, September [ 1 894?], Giraldi 529 (Fl-syntype). H. ascyron var. umbellatum R. Keller in Bot. Jb. 33: 550 (1904). Type: China, Shaanxi, Ki-san, 1896 (fr), Scallan in Biondi 3813 (FI!-holotype). H. ascyron [var.] macropetalum hort. ex Vilmorin & D. Bois., Fruct. Vilmor.: 24 (1904), nomen. H. kelleri H. Leveille in Bull. Soc. Bot. France 54: 499 ( 1 906) [et in clav. p. 497, qua 'Levl. & Vant.'], non Baldacci (1895). Type: Japan [without precise locality], Faurie s.n. (P-holotype). Apart from having only 4 styles, this plant (as described) would appear to be typical H. ascyron. H. yabei H. Leveille & Vaniot in Repert. Spec. Nov. Regni Veg. 5: 279 (1908) non H. Leveille & Vaniot in Bull. Soc. Bot. France 53: 501 (1906); Nakai in J. Coll. Sci. Imp. Univ. Tokyo 31: 453 (191 1). Type: Korea, Quelpaert I., August 1907, Faurie 1792 (E!- holotype). H. hemsleyanum H. Leveille & Vaniot in Bull Soc. Bot. France 54: 592 (1908), in Mem. Real. Acad. Ci. Barcelona III, 12: 553 (1916); Lauener in Notes Roy. Bot. Card. Edinburgh 27: 3 ( 1 966). Type: China, Jiangsu, Zuo-se, n.d. (fl.), d'Argy s.n. (E!-holotype; BM!-isotype). H. ascyron var. hupehensis Pamp. in Nuov. Giorn. Bot. Ital. N.S. 17: 669 (1911), op. cit. N.S. 18: 129 (1911). Type: China, Hubei, 14 syntypes from southern mountains at 600-1850 m, Silvestri 1471-1481 + 1471a, 1472a, 1479a (all Fl); Monte Cia-inen-ku presjo Sian-yang, Silvestri 1471 (Fl-lectotype, A!, photo, is selected here). This variety is very like var. giraldii, but differs in that the leaves are not or scarcely cordate, the flowers are larger (40-50 mm in diam.) and the capsule is shorter (8-10 mm wide). H. ascyron var. vilmorinii Rehder in Mitt. Deutsch. Dendrol Ges. 24: 235 (1916), in Bailey, Stand. Cyclop. Hort. 3: 1630 (1919). Type: cf. Sprague in Curtis 's Bot. Mag.: t. 8557 (1914) as H. ascyron. This appears to be a re-naming of H. ascyron var. macropetalum (1904). Fig. 9 A. H. ascyron subsp. ascyron: (a) habit (medium-styled); (b) long-styled ovary; (c) medium-styled ovary: (d) medium-styled capsule. B. H. ascyron subsp. pyramidatum; (e) capsule. C: H. ascyron subsp. gebleri: (f) leaf; (g) capsule. D. H. concinnum: (h) habit; (i) capsule (a, f, g, h x 1 ; rest x 3). A. Bisset 3465, except (b) Chen 199. B. Burgess 343. C. Redrawn from Ledebour, Ic.fl. ross.: t. 487. D. (h) Krautters. n.; (i) Meebold 9798. 56 N.K.B. ROBSON '-' ^vtV.r C fl^^Ji % Map 3 Sect. 7: la. //. ascyron subsp. ascyron ; Ib. //. ascyron subsp. gebleri O specimens, other records A. H. ascyron var. a damii R. Keller in Bot. Jb.: 58: 191 (1923). Type: Japan or Korea, Warburg s.n. (Bf-holotype). H. sagittifolium Koidz., Fl. symb. orient. -asiat.: 91 (1930). Type as for H. yabei H. Leveille & Vaniot (1908 non 1906). '//. ascyron f. longistylum Maxim.' is also cited. H. ascyron forma vilmorinii (Rehder) Rehder, Bibl. cult, trees : 462 (1949). H. ascyron var. ascyron f. angustifolium Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 125 (1951). Type: Japan, Honshu, Prov. Kai [Yamanashi], Yamanaka, 29 July 1934, Y. Kimura 3486 (TI- holotype). Icones: Rchb., Iconogr. Bot., cent. 5: t. 490, f. 183 (1827); Sprague in Curtis's Bot. Mag.: t. 8557 (1914); Liou Tchen-ngo et al. in Fl. PL Herbac. Chinae Bor.-Or. 6: 71, t. 27 ff. 1-7 (1977). Plant 0.5-1. 3m tall. Leaves 40-120 x (5-)7-40 mm, ovate-lanceo- late or rarely narrowly elliptic to linear-lanceolate or narrowly oblong, base usually cordate-amplexicaul. Flowers 45-80 mm in diam. Sepals 6-10 mm wide, ovate or oblong to elliptic or lanceolate or obovate, rounded to obtuse or rarely apiculate to acute. Styles 4 15 mm, almost free to up to 0.8 united. Capsule ovoid to ovoid-pyramidal or ovoid-cylindric. Distribution of the species (see p. 53), except for north-west and north-east China (Xinjiang, north Manchuria), Sakhalin, Kamchatka, the Kurile Is. and North America. RUSSIAN FEDERATION. Ob: ad ripas fl. Ob prope Bogorodskoje, 29 June 1892 (fl), 30 August 1892 (fr), Krylov 1918 (H). Irtysh: (Gorschkova, 1949). Altai: Altai, pre-1893 (fl), Thomas in Herb. Boiss. (G); Gorno- Altaiskaya Avt. Obi., Elekmonarskiy raion, okrestnosti sela Chemal u ruchya, 29 August? 1961, Velikanov s.n. (H). Angara-Sayan: Irkutzk, 1874 (fl), Augustinowicz s.n. (K, LE*); Transbaikalia, mentis Arschan, 12-16 July 1911 (fl), Ahuger s.n. (H); Reg. transbaical., ad pagum Verchne-Ubukunsk (juxta flumen Selenga), 500-1000 m, 20 July 1900 (fl), Ehrenberg s.n. (H). Dauria: Nerchinsk, Schilkafluss, bei Monastyr, 1892 (fl & fr), Karo 220 (E, H); Juzhn. Zabaykalskiy, Khentai-Gukonskoye nagorye, Bas. r. Gukoi, 18 August 1960 (fl), Maksimova s.n. (K, LE*). Zeya-Bureya: Zejskaja Frisian am Zeaflusse, July 1 899 (fl), Karo 377 (BM, E, G*, H, JE). Uda: Khabarovsk!* krai, Ul'chskii raion 1 km E. of Susanino, road to Anninskie Vody, 40 m, 27 August 1976 (fr), Alanko (H). Ussuri: Prov. Amurensis, fluvium Amur vallis Uril, in pratis inter Uril et Yrjasnaja, 5 July 1 895 (fl), Komarov 1090 (BM, K); Prov. Amur, in monte Hehzier prope stationem ferrosam Korfowskaja, circa a 40 km procul ab oppido Chabarowsk, 3 August 1913, Enander s.n. (E). MONGOLIA. Khentei, Khangai, Mongol-Daurian and Great Khingan regions (Grubanov, 1996). CHINA. Nei Mongol: NingchengXian,CunjingouGongshe, 21 July 1981 (fl), Li S.X. et al. 4438 (IFP). Heilongjiang: Khalka Gol, 27 July 1902 (fl), Campbell s.n. (BM); Daishan, 20 July 1956 (fl), Zhongde Exped. 7614 (PE). Jilin: Districtus Omoso, Trajectus Tien-guan-czai-lin, 12 August 1896 (fl & fr), Komarov 1091 (BM); Zhangjiadian, 26 July 1958 (fl), Wang C.S. et al. 1512 (IBSC). Liaoning: Dairen [Talien], June-August 1910 (fl), Stuart s.n. (BM); Huanren, 23 August 1964 (fr), Cui & Zhu 186 (PE). Hebei: Tche-ly, montagnes du Fou-Ping, August 1910 (fr), Chanel 542 (BM, E); Tong Ling, 7 July 1 935 (fl), Liu K 1 1 606 (IBSC). Shanxi: Qin Xian, Wulongchuan, 1 6 July 1959 (fl), Chen Y.L. 899 (PE); Lishan Xian?, Muchang Gou, 1550 m, 29 June 1955 (fl), Huanghe Exped. 2288 (PE). Shaanxi: Shensi Central, voyage a Cheuhaiyao, 1100m, 13 August 1916 (fr),Licent 2438 (BM,K);Liuba Xian, Miaotaizi, 1 500 m, 6 September 1977 (fr), Wang & Shi 308 (PE). Gansu: Lien hoa shan, Ha Kai valley, 2700 m, 1 4-20 July 1 925 (fl), Rock 1 2772 (E, GH, K); HeshuiXian,TaibaiZhen, 1250m, 17 July 1954, Huanghe Exped. 606 (IBSC, PE). Ningxia: Jingyuan Xian, Danan Chuan, Nantai Gou, 2300 m, 28 July \964(f[),JingyuanExped. 36(PE).Henan:LushiXian, 1 180m, 28 September 1958 (fr), Fu J.Q. 1446 (IBSC); Lushih-hsien, Hsuingerhling, 12 July 1935 (fl), Liou 4712 (K). Shandong: Fei Hsien, Meng Shan, 700m, 29 July 1936 (fr), Cheo & Yen 24 1 (BM); Lao Shan, 600 m, 1 July 1 930 (fl), Chiao C. Y. 2658 (GH, IBSC, NY); behind Fu Shan, 1 50 m, 7 July 1 923 (fl), Sha F.H. 1 69 (PE). STUDIES IN HYPER1CUM Jiangsu: Nanking, 2 1 July 1 922 (fl), Yu P.Lin Herb. Univ. Nanking 2999 (E); Yixing Xian, Longchi Shan, 25 June 1 962 (f\),MaoS.H. etal. 1 1 (KUH, NAS). Anhui: Yang chia ping, July 1905 (fl), Schindler 39 (BM, E, K); Taiping, Longyuan, 23 June 1959 (fl), Anhui Exped. 600 (IBSC). Zhejiang: Ningpo, 1 874 (fl), Forbes 1 08 1 (BM); Tianmu Shan, 1 5 July 1 957 (fl ), He X. Y. 25408 (IBSC). Fujian: Diongloh, Sin Sai, 24 July 1925 (fl), Cheng RE. 2407 (BO). Jiangxi. Lu Shan, n.d. (fr), Hsiung Y.K. 6928 (IBSC); Xunwu Xian, Jingxi Gonshe,700m, 13 August 1962(fr), YueJ.S.elal. 1824 (RUN). Guangdong: Yangshan Xian, S. of Linchow, July-September 1 932 (fr), Tsui T.M. 580 (GH, K,NY,PE);Chaoan Xian, 25 April \973(fr),LuoX.R. 1297 (IBSC). Hainan: Chahar, Hsiao- wu-tai-shan Exped., 6 July 1936 (fl), Wu & Yang 36810 (PE). Guangxi: Nandan Xian, Mangchang, Jiuwei Shan, 2500 m, 3 July 1937 (fl), Wang C. 40964 (A, IBSC); Gui Xian, Tan Tang, 4 October 1958 (st), Zhong S.Q. 21098 (KUN). Hunan: prope urbem Wukang, 350-1400 m, 4 June-9 August \9\8,Handel-Mazzetti 1 2270 (E);Qianyang Xian, 28 June 1954(fl), LeeC. 7:2487 (IBSC, PE).Hubei:Hsing Shan, June 1901 (fl), W/7so/i2200(E, K); Wuchang, Luojiashan, 6 September 1 932, Chung H.H.91 77 (A); Lichuan Xian, 1500 m, 24 September 1951, Dai & Qian 1541 (PE). Sichuan: Jiuzhaigou, NW of Songpan, 3000 m, 1 September 1 986 (fl), Lancaster 1 602 (BM); Mu-li hsien, 2450 m, 19 August 1978 (fr), Zhao C.S. et al. 7574 (SZ). Guizhou: Yanhe Xian, Daping, 1000 m, 22 July 1957 (fl), N. Guizhou (Qianbei) Exped. 2416 (PE); Ya-Tze-Ho, Tsingchen, 5 July 1935 (fl), Teng S. W. 762 (IBSC). Yunnan: Mo-so-yan, pres de Lan Kong, 5 August 1 883 (fl & fr),Delavay 1 11 (P);Pingbian Xian, 1300m, 20 July 1 934 (fl),7kn//. 7:6 1036 (A, IBSC, NAS, PE); Chungtien [Zhongdian], Haba, 2600 m, 25 November 1937 (fr), Yu, T.T 14964 (KUN). Qinghai: no precise locality, 1901 (fl), Zimmermann 229 (US). VIETNAM. Indo-China, July 1924 (fl), Petelot s.n. (BO). TAIWAN. Hsinchu: Sintiku, 18 May 1923 (fl), Simada 1209C (TAI). KOREA. North: 80 km from Seoul, Wusan and Pyongyang [3 localities], 1 892 (fl), Veitch 56 (BM). South: Chemulpo, July 1 884 (fl), Carles 1 05 (BM, E, K); Kang Wong Prov., Inje Distr., Sorak-san Nat. Park, road to Paekdamsa Temple, 450 m, 28 September 1989, Kirkham, Flanagan & Boyce KFBX 26 (K); in Chinnampo, August 1906 (fl), Faurie 635 (E); Port Chusan [Ullungdo I.], June 1859 (fl), M//W928 (K); Quelpaert [Cheju], in herbidis Hoakin, August 1909 (fl), Taquet 1678 (E, K). JAPAN. Hokkaido. Oshima: Yezo, in planitie Hakodate, August 1905 (fl), Faurie 6902 (BM); Hakodate, 1861 (fl & e. fr), Maximowicz Her II s.n. (BM, K). Shirabeshi: Mt. Moiwa, August 1912 (fl & fr), Sakamura in Kudo 1254 (TAI). Iburi: Tomokomai Forest, 31 August 1914 (fl & fr), Yoshlmi in Kudo 121 (TAI). Ishikari: Sapporo, 29 July 1991 (e. fr), Tokobuchi s.n. (K). Kushiro: Me-akan dake, 2 August 1929 (fl), Tanaka 164 (BM); Nita Teshikaga-Cho, 5 August 1974 (fl), Furuse 6660 (K). Honshu. Tochigi: Nikko-Chuzugi in Shimotsuko, 7 September 1952 (fr), Kubota NSM 695 (BM, E, H, K). Gunma: Mt. Akagi, 11 July 1949 (fl), Tanaka s.n. (K); Chusenji[-ko], August 1 887 (1. fl), Bisset4269 (BM, E). Nagano: Karuizawa, 1 050 m, 3 August 1912 (fl), Fox s.n. (BM). Tokyo: Komaba, 26 October 1 878 (fr), Bisset 1472 (E); Musashi, Akabane, 10 August 1910 Sakurai s.n. (H). Kanagawa: Lagami, Mt. Hakone, August 1906, Yokohama Nursery Co. s.n. (E). Shizuoka: Prov. Kai, Minamitsuru-gun, Mt. Fuji, 1933 (fl & e. fr), Makino 7697 (CAS, E). Mie: Ise, Nago, August 1907 (fl & fr), Yokohama Nursery Co. s.n. (E). Nara: Mt. Izawa, 8 September 1941 (fr), Seki in HTU 77587 (TAI). Shikoku. Tokoshima: Kumatani, 1914 (fl), Takayanagi in Sasaki HTU 76870 (TAI). Kyushu. Nagasaki: Nagasaki, Kundshovan, 1 863 (fr), Maximowicz Iter II s.n. (K). CULTIVATED. Specimens seen from England ( 1 763->), Scotland ( 1 827- 1 898), Ireland ( 1 968), France ( 1 84 1 ), Belgium ( 1 979), Sweden ( 1 8th C) and Finland (191 1-1978). lb. Hypericum ascyron subsp. gebleri (Ledeb.) N. Robson, stat. nov. Type as for H. gebleri Ledeb. Fig. 9C, Map 3. Hypericum gebleri Ledeb., Fl. altaic. 3: 364 (1831), Icon, pi: 487 (1834), Fl ross. 1: 446 (1842); Maxim.; Primit. fl. amur.: 461 ( 1 859), in Bull. Acad. Imp. Sci. Saint-Petersbourg 27: 43 1 ( 1 882), Mel. Biol. 11: 163 (1882); Krylov, FL Altaya: 189 (1901); Komarov, Fl. Manshur. 3: 42 (1905); H. Leveille in Bull. Soc. Bot. France 54: 593 (1908); Nakai, Tyosen-syokubutu 1: 158, f. 189 (1914), Fl. Mt. Paik-tu son: 67 (1917); Miyabe & Miyake, Fl. 57 Saghalin: 78 (1915); Kudo in J. Agric. Hokkaido Univ. 12: 47 (1923); ? in Kitabahuto-Syokobutu-Tyosyo: 179, t. 17, f. 85 (1924); Makino & Nemoto, Fl. Jap.: 541 (1925); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 176 (1925); Komarov, Fl. pal. Kamch. 2: 303 (1929); Hulten in Kungl. Svenska Vetenskapsakad Handl., 3rd ser. 8: 1 26, map 554 (1929); Komarov & Kleb.-Alis., Opred. r. Dal'nevost kr. 2: 748 (1932); Krylov, Fl. Zap. Sib. 8: 1903 (1935); Sugawara, Ill.fl. Saghalien 3: 1301, t. 598 (1940), repr. (1975); Gorschkova in Fl. URSS 15: 213 (1949); Grubov, Konsp. Fl. Mongol. Narod. Respub.: 205 (1955), Opred. Sosu. Rast. Mongolii: 181 (1982); Pavlov et al., Fl. Khazakhstana 6: 161 (1963); Noda, Fl. N.-E. Prov. (Manchu- ria) of China: 194(1911); Liou Tchen-ngo et al. in Fl. PL Herbac. Chinae Bor.-Or. 6: 73, t. 27, ff. 9-12 (1977). Type: Kazakhstan, Altai, 'Habitat ad montium latera juxta fl. Buchtarma ex adverse Sinensium praesidii Dschingis-tei', July-August (fl), Ledebour s.n. (LE-holotype). Roscyna gebleri (Ledeb.) Spach, Hist. nat. veg. Phan. 5: 430 ( 1 836), in Annls Sci. Nat. Bot. II, 5: 64 (1836). H. ascyron var. brevistylum Maxim., Primit. fl. amur.: 65 (1859); H. Leveille in Bull. Soc. Bot. France. 53: 498 (1906); Kitag., Neo- lin.fl. manshur.: 443 ( 1 979). Type: Russian Federation, E. Siberia, prope Dshai, 50 circa stadia a pago Kitsi, Maack s.n. (LE- syntype); infra montes Burejae, Maack s.n. (LE-syntype). Maximowicz (loc cit. above) stated that this plant was H. gebleri Ledeb. unless the narrowly ovate-oblong leaves made it different; by 1882 (Bull. Acad. Imp. Sci. Saint-Petersbourg 27: 431) he had decided to include it in that species. H. sachalinense H. Leveille in Repert. Spec. Nov. Regni Veg. 6: 330 (1909); Gorschkova in Fl. URSS 15: 257 (1949). Type: Russia, E. Siberia, Sakhalin I., Vladimirov, August 1908 (fl), Faurie 518 (Ei-lectotype, selected here; BM!, W!-syntypes); Vladimirov, September 1908 (fr), Faurie 519 (BM!, E!, W!-syntypes); Saghalien, September 1908 (fr), Faurie s.n. (E!-syntype). Icones: Ledeb., Icon. fl. ross. 5: t. 487 (1834); Suguwara, Ill.fl. Saghalien 3: 1300, t. 598 (1940). Plant 0.5-1 m tall. Leaves 30-60 x 4-15 mm, narrowly lanceolate or narrowly elliptic-lanceolate to oblong-linear or oblanceolate, base usually cuneate. Flowers 45-50 mm in diam. Sepals 1.5-7 mm wide, narrowly oblong or oblong-lanceolate, rounded or usually obtuse to acute. Styles c. 2.5-3.5 mm, about half as long as ovary, free. Capsule cylindric-ellipsoid. Kazakhstan, Russia (Altai, Angara-Sayan, Dauria, Zeya-Bureya, Uda, Ussuri, Sakhalin, northern Kurile Is., Kamchatka), Mongolia, China (extreme west Xinjiang, Heilongjiang, Jilin?), Korea (north). KAZAKHSTAN. Altai: Songaria chin., ad lacum Saisang-Nor, 1838 (fl), [Fischer] s.n. (K, LE*). RUSSIAN FEDERATION. Angara-Sayan and Dauria (Gorschkova, 1949). Zeya-Bureya: Amur, 1871 (fl), Augustinowicz s.n. (K, LE*). Uda: Bureinskiy Khrebet, infra montes Burejae, Maack s.n. (LE*). Ussuri: (Gorschkova, 1949). Sakhalin: Vladimirov, August 1908 (fl), Faurie 518 (BM, E, W). Kamchatka: Kamtchatka australis, Savoiko, 27 July 1928 (fl), Eyerdam s.n. (E), Malaise s.n. (K); Kimitina R., 50 km NW from Mashura (Hulten, 1929). Kurile Islands: no locality cited (Gorschkova, 1949). MONGOLIA. Khentei (Grubanov, 1996). CHINA. Heilongjiang, Jilin? (Komarov, 1950). KOREA. North: via ab oppido Musang ad oppidum Kapsan, Vallis Tadin-don, 22 June 1897 (fl), Komarov 1091 (K). When making a survey of Chinese specimens in 1993, 1 had not then decided to recognize subspecies in H. ascyron. I do not, therefore, have detailed records of the occurrence of subsp. gebleri in the north of China. Reichenbach's illustration of H. salicaria (see synonymy of 58 N.K.B. ROBSON Map 4 Sect. 7: Ic. H. ascyron subsp. pyramidatum specimens, O other records. subsp. ascyron) shows a plant with narrow leaves basally rounded and small flowers with narrow but unequal foliaceous sepals, short free styles and a pyramidally ovoid capsule. If this apparent interme- diate form between subsp. ascyron and subsp. gebleri has arisen independently of the Chinese intermediates, then subsp. gebleri would be diphyletic in origin. The epithet gebleri would then go with the Altai plant. Ic. Hypericum ascyron subsp. pyramidatum (Aiton) N. Robson, comb, et stat. nov. Type as for H. pyramidatum Aiton. Fig. 9B, Map 4. H. bartramium Miller, Card. Diet. 8th ed., no. 10 (1768). Type: Cult, ex America in Hort. Chelsea (BM!-holotype). H. pyramidatum Aiton, Hort. kew. 3: 103 (1789), op. cit., 2nd ed. 4: 422 ( 1 8 1 2); Willd, Sp. pi. 3: 1444 ( 1 802); Vent., Jard. Malmaison 2: t. 1 18 (1804); Pursh, Fl. Amer. sept.: 374(1814); Choisy, Prodr. monogr. Hyperic.: 41 (1821), in DC., Prodr. 1. 544 (1824); Sprengel, Syst. veg. 3: 342 (1826); Torrey & Gray, Fl. N. Am. 1: 158 (1838); Gleason, New Britton & Brown III. Fl. 2: 541 (1952); Steyerm., Fl. Missouri: 1058 + map (1963); Utech & Iltis in Trans. Wisconsin Acad. Sci. 58: 337, map 5, f. 1 (1970); Mohlenbr., ///. FL Illinois, Hollies to Loasas: 29, t. 10 (1978); Scoggan, Fl. Canada 3: 1097 (1978); Swink & Wilhelm, Pis of Chicago region, rev. ed.: 390 (1979); Kaul in Great Plains Fl. Ass., FL Great Plains: 239 (1986). Type: Cult, in Hort. kew., n.d. (fl & fr), [Alton!] (BM!-holotype). H. amplexicaule Lam., Encycl. 4: 147 (1797), non Gilib. (1782). Type: Cult. Paris (Jardin des plantes) ex N. America (P-LAM- holotype). H. ascyroides Willd., Sp. pi. 3: 1443 (1802); Pursh, Fl. Amer. sept.: 374 (1814); Nuttall, Gen. N. Am.: 16 (1818); Bigelow, Fl. Bost. 2nd ed.: 279 (1824); Hooker, Fl. bor.-amer. 1: 109 (1840). Type: Habitat in Pensylvania, Muhlenberg s.n. (B-WILLD-holotype, microfiche!). H. macrocarpum Michx., Fl. bor-amer. 2: 82 ( 1 803); Sprengel, Syst. veg. 3: 342 ( 1 826). Type: Canada, circa Montreal, Michaux s.n. (P-holotype). H. ocymoides Loddiges, Cat. PL Conrad Loddiges & Sons, Hack- ney, 15th ed.: 71 (1830), nomen. Roscyna americana Spach, Hist. nat. veg. Phan. 5: 431 (1836), in Ann. Sci. Nat. Bot. II, 5: 364 (1836). Type as for//, pyramidatum Aiton H. ascyron sensu Coulter in Bot. Gaz. 11: 83 (1886); Jones & Fuller, Vase. Pis Illinois: 324, map 795 (1955); Gillett & Robson in Publs Bot., Natl. Mm. Nat. Sci. Canada, no. 11: 23, t. 9, map 7 (1981); Magee & Ahles, Fl. Northeast : 737 (1999) et auct. Amer. plur. Hypericum ascyron van americanum (Spach) Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 124 (1951). Icones: Vent., Jard. Malm. 2: t. 118 (1804); Mohlenbr., ///. Fl. Illinois, Hollies to Loasas: 29, t. 10 (1978). Plant 0.5-2 m tall. Leaves 40-80 x 18-33 mm, ovate-lanceolate to lanceolate or oblong, base cordate to rounded. Flowers 40-70 mm in diam. Sepals 8-13 mm wide, ovate to lanceolate or oblong-elliptic, acute to shortly acuminate. Styles 3-7 mm, 0.8-1.5 x ovary, almost free or rarely up to 0.8 united. Capsule ovoid. Canada (Quebec, Ontario, Manitoba?), U.S.A. (north-east, from Minnesota to Massachusetts and south to Missouri, Ohio and Mary- land). CANADA. Quebec: Montreal, July 1 866 (fl), Goldie s.n. (K); lie au Corbeau, 26 May 1 859 (fl), Bourgeau s.n. (K); Deux-Montagnes, Saint-Columban, 30 August 1936, Marie-Victorin 46649 (CM). Ontario: London, 17 September 1883 (fl), 5 September 1885 (fr), Burgess 343 (BM); L. Huron, Goderich, 19 August 1901, Macoun in Herb. Geol. Surv. Canada 34076 (K); Rice Lake Plain, July 1866 (fl), Macoun s.n. (K); Lambton Co., 8 km S. of Watford, 17 July 1951 (fl), Mulligan 848 (H). U.S.A. Connecticut: (Magee & Ahles, 1999). Illinois: Beardstown, 1842 (fl), Geyers.n. (K); Dixon, pre 2 April 1 866 (fl), Rolfe s.n. (BM); Augusta, n.d. (fl), Mead s.n. (K). Indiana: (Crovello, Keller & Kartesz, 1983). Iowa: Decora, 20 July 1 884 (fl), Holway s.n. (H). Massachusetts: Boston, n.d. (fl), B.D.G. s.n. (K). Maine: (Magee & Ahles, 1999). Maryland: (Brown & Brown, 1984). Michigan: White Pigeon, n.d. (fl), Carey s.n. (K); Flint [address], August 1876 (1. fl), Clarke 270 (K); Dickinson Co., 6.4 km N. of Norway, 6 August 1961,/fertry s.n. (CM). Minnesota: Minneapolis, 22 July STUDIES IN HYP ERIC UM 59 1991 [71 891 1 (fl), Sandberg 613 (H); Little Falls, n.d. (1. fl), Vasey s.n. (K); Duluth, junction ofSaively Boulevard and Glen Lake, 1942, Beers.n. (CM*). Missouri: Watson, 1 1 July 1 894 (fl), Bush 37 (K); Atherton, 27 June 1 898 (fl), Bush 19 (K). New Hampshire: (Magee & Ahles, 1999). New York: Buffalo, n.d. (fl), Clinton s.n. (BM); near Troy, n.d. (fl), Torrey s.n. (G-DC, K); upper Susquehanna, along Cheming R., 12 July 1894 (fl & fr), Lucy & Elmira 1171 (H). Ohio: Portage Co., Mantua township, 15 July 1922, Webb s.n. (CM); Wayne Co., near Woostar, 20 June 1910, Hopkins s.n. (CM*). Pennsylvania: Philadelphia, December 1 886 (fl), Herb. Corstorphine s.n. (BM); Bethlehem, banks of Lehigh R., 1 October 1849 (fr), Anon. s.n. (K); Lawrence Co., Slippery Rock Creek above Wurtemburg, 25 July 1900 (fl & fr), Shafer s.n. (CM); Lackawanna Co., Susquehanna R. at Falling Springs, 14 July 1937, Glowenke6\\ (CM). Vermont: (Magee & Ahles, 1999). Wisconsin: PalkCo., August 1 900 (fl), Baker s.n. (JE); Wyoming, n.d. (fl), Vuran s.n. (H). The characteristic features of the American plants, namely acute to acuminate sepals and shortly connate styles, are certainly not con- stant enough to allow recognition as a species. However, these characters rarely occur together in Asia, where almost free styles are not uncommon but acute sepals appear to be confined to Japan and acuminate sepals are apparently absent. It would seem appropriate, therefore, to give the American plants subspecific rank. W.G. Dore suggested that the sporadic distribution of//, ascyron in Canada, which seems to coincide well with the location of earlier native American Indian camp sites, indicates that this plant was probably distributed by aborigines (Gillett & Robson, 1981: 25), though the reason for this association is not clear. 2. Hypericum przewalskii Maxim, in Bull. Acad. Imp. Sci. Saint- Petersbourg 27: 431 (1882), Mel. Biol. 11: 164 (1882), Fl. tangut. 1: 99, t. 18 ff. 1-12 (1889); Forbes & Hemsley in J. Linn. Soc. 23: 74 (1886); H. Leveille in Bull. Soc. Bot. France 54: 593 (1908); Anon., Iconogr. Cormoph. Sinicae 2: 880, f. 3489 (1972); Li Xiwen in Fl. R. P. Sinicae 50(2): 45, t. 1 2 ff. 4-6 ( 1 990). Type: China, Gansu, parte altae alpina occidentali, ad fl. Yussun- Chatyma, 1880, Przewalski s.n. (LE-holotype). Fig. 10, Map 5. H. obtusifolium R. Keller in Bot. Jb. 33: 551 (1904); H. Leveille in Bull. Soc. Bot. France 54: 591 (1908); non Makino (1903). Type: China, Shaanxi, in monte Huan-tou-san, July 1889 (e. fr), Giraldi in Biondi 3824 (Fl l-lectotype, selected here); in rupestribus mentis Kian-san, prope Sce-kiu-tsuen che dista circa 65 km, 4 August 1897 (e. fr.), Giraldi in Biondi 3825 (FI!-syntype). H. chinense van minutum R. Keller in Bot. Jb. 33: 548 (1904 ); H. Leveille in Bull. Soc. Bot. France 54: 590 (1908). Type: China, Shaanxi, Thae-pei-san, 15-20 July 1897 (fl), Giraldi in Biondi 3837 (FI!-holotype; K!-isotype). H. pedunculatum R. Keller in Bot. Jb. 33: 549 (1904); H. Leveille in Bull. Soc. Bot. France 33: 590 (1908); Li Xiwen in Fl. R. P. Sinicae 50(2): 71 (1990). Type: China, Shaanxi, Tsil-lin-san, 10 July 1900 (fl), Giraldi in Biondi 7135 (FI!-holotype). H. biondii R. Keller in Bot. Jb. 33: 551 (1904); H. Leveille in Bull. Soc. Bot. France 54: 591 (1908) pro parte quoad spec. cit. Biondi 529. Type: China, Shaanxi, in monte Thae-pei-san, August 1899 (e. fr), Giraldi in Biondi 529 (Fll-lectotype, selected here). The other syntype (Biondi 530) is H. ascyron. H. macrosepalum Rehder in Sargent, PL Wilson. 3: 451 (1917); Li Xiwen in Fl. R. P. Sinicae 50(2): 43 (1990). Type: China, Sichuan, Ta-p'ao-shan, NE of Tachien-lu [Kangding], 3300 m, 5 July 1908 (fl), Wilson 2426 (A!-holotype). N.B. Sargent cites the locality as Tai-pei-shan, but the label of the type specimen has it as above. Icon: Anon, in Iconogr. Cormoph. Sinicae 2: 880, f. 3489 (1972); Li Xiwen in Fl. R. P. Sinicae 50(2): 58, t. 12 ff. 1-3 (1990). Perennial herb (0.17-)0.3-0.55 m tall, erect or sometimes ascend- ing from creeping and rooting base, with stems few to numerous, simple or usually branched below inflorescence or throughout. Stems incompletely 4-lined or 2-lined when young, nearly always soon becoming terete; internodes 25-80 mm, shorter than leaves except upper (slightly longer) or rarely all longer. Leaves sessile; Map 5 Sect. 7: 2. H. przewalskii, type ; '//. pedunculatum' specimens, other record D; 'H. macwsepalum A. 60 N.K.B. ROBSON Fig. 10 H. przewalskii: A. 'pedunculatuni 1 : (a) habit; (b) sepal; (c) petal; (d) ovary. B. przewalskii, type: (e) habit; (f) sepal. C. 'macrosepalum': (g) habit; (h) sepal; (i) ovary (a, e, g x 2/3; rest x 3). A. Wilson 582. B. G.H. Yang 58812. C. Pratt 541. STUDIES IN HYPERICUM 61 lamina 20-65(-80) x 10-32 mm, broadly oblong or oblong-ovate to narrowly oblong or oblong-lanceolate (reducing in size down stem), rather paler beneath, not glaucous, plane, chartaceous; apex rounded to shallowly retuse, margin entire, base cordate-amplexicaul; vena- tion: 4-6 pairs of main laterals from lower half of midrib, with subsidiary midrib branches and densely reticulate tertiary venation not or scarcely prominent beneath; laminar glands pale, dense, unequal dots or short streaks; intramarginal glands pale, dense, small. Inflorescence 1-7-flowered from 1-3 nodes, corymbiform, sometimes with flowering branches from up to 5 nodes below, the whole then narrowly pyramidal to cylindric; pedicels 12-22 mm (- 45 mm in fruit); bracts and bracteoles foliose, ovate, persistent. Flowers 20-40 mm in diam., stellate; buds narrowly ovoid to cylindric, obtuse to rounded. Sepals 8-10 x 2-4 mm, sometimes enlarging up to 15 x 6.5(-8) mm in fruit, free or basally united, imbricate, subequal or unequal or rarely foliaceous, erect in bud, spreading to deflexed in fruit, narrowly or broadly oblong or elliptic to lanceolate or ovate-lanceolate or triangular-lanceolate, rounded to obtuse or rarely shortly mucronate, entire but margin undulate; veins (5)7-9, branching distally and reticulating, midrib scarcely differentiated; laminar glands rather sparse, punctiform, distal and submarginal, also a few basal, linear; marginal glands dense, small. Petals bright yellow, not tinged red in bud, 12-18x4-8 mm, 1.5-1.8 x sepals, oblong-oblanceolate, slightly curved, rounded, without apiculus, margin entire; laminar glands pale, linear, sometimes interrupted distally; marginal glands absent. Stamen fascicles 5, each with 15-30 stamens, longest 10-24 mm, c. 0.85-1 x petals; anther gland amber. Ovary 5(6)-locular, 4-9 x 4-6 mm, ovoid; styles 5(6), 4.5-13 mm, 1-1.8 x ovary, 0.5-0.9 united or appressed (when short); stigmas narrowly capitate. Capsule 14-21 x (5)7-14 mm, broadly to narrowly ovoid or subcylindric, 1-2.1 x sepals, obtuse, with numerous narrow longitudinal vittae. Seeds dark red- dish-brown to greyish-brown, 1.3-1.5 mm, cylindric, not curved, shallowly carinate, without terminal expansion; testa densely shallowly linear-reticulate. 2n = ? Mountain slopes, river bank thickets, meadows and roadsides; 2150-3400(-4000) m. China (Gansu, Shaanxi, Henan, west Hubei, Sichuan, north Yunnan, Qinghai). CHINA. Gansu: T'ao R. basin, meadows of Toyuku, near Choni, 2550 m, July 1 925 (fl), Rock 1 2909 (E, K, PE); Wushan Xian, Laojunshan, 1 600 m, 30 May 1956 (fr), Huanghe Exped. 4241 (PE). Shaanxi: Monte Hua-tzo-pin, 20 June 1894 (fl), Giraldi in Biondi 533 (FI); Tai-pei-shan, 1910 (fl), Purdom s.n. (K); Zhashui Xian, Laolin Gongshe, 2200 m, 14 June 1973 (f\),HouX.X. 1888 (IBSC). Henan: Luanchuan Xian, 1959 (fr), Anon, in Herb. Beijing Inst. Bot. 20661 (IBSC). Hubei: Fang Hsien uplands, August 1907 (fl), Wilson 582 (BM, E, K, NY); Badong Xian, Shennongjia, Niudongwan, 2100 m, 23 July 1957 (fl), Fu G.X. & Zhang Z.S. 1033 (IBSC, NAS). Sichuan: Kangding Xian (Tatsien-lu), 2700-2870 mm, 25 September 1928 (fr), Fang 3545 (E, GH, K, US); Zhegu Shan, 3300 m, 19 August 1957 (fr), Zhang Z. Y. & Zhou H.F. 23864 (IBSC); Li Xian, Dabanzhao, 3650 m, 28 June 1957 (fr), He D.P. 44559 (SZ). Yunnan: Lichiang Range, 1933 (fr), McLaren's Collec- tors s.n. (BM). Qinghai: Yellow R. banks, SE of Radja, 3300 m, June 1926 (fl), Rock 1 42 1 3 (E, GH, K, NY); Minhe Xian, Niuxinshan Xian, Tangeryuan Xiang, 2740 m, 9 August 1958 (fr), Zhu Z.Y. et al. 260 (KUN); Huangyuan Xian, Guayin Forest Distr., 1 1 July 1958 (fl), Tsoong P.C. 8890 (PE). H. przewalskii is closely related to H. ascyron, differing from it essentially by the obtuse to retuse leaf apex and usually 2-lined mature stem internodes. The remarks about variation in style length in H. ascyron (p. 53) would seem to apply also to H. przewalskii. Where they are short, they seem to be almost free and appressed rather than united. Two variants of//, przewalskii have been given specific rank, but neither would appear to merit it. '//. pedunculatum" (Fig. 10A) has longer narrower leaves and is more branched than typical //. przewalskii and, on the basis of the Wilson and Giraldi specimens, appeared at first to be distinct. A study of more abundant material from Chinese herbaria, however, revealed a complete intergradation between the two extremes. '//. pedunculatum' is the south-eastern- most population of H. przewalskii, in western Hubei and adjacent Shaanxi, and the one nearest morphologically to H. ascyron. 'H. macrosepalum' (Fig. IOC) seems to be aberrant in having more persistently 4-angled internodes, but is otherwise typical of forms of//, przewalskii with a one-flowered inflorescence and foliar sepals in fruit from Sichuan and Shaanxi. [For Sect. 9. Hypericum see Part 4 (2, 3).] Sect. 9a. CONCINNA N. Robson in Bull. Br. Mus. not. Hist. (Bot.) 8: 173(1981). Hypericum sect. Androsaemum subsect Pseudoandrosaemum R. Keller in Engler & Prantl, Nat. Pflanzenfam. 3(6): 22 1 ( 1 893); op. cit.,2nded.,21: 1 77 (1925) pro parte, quoad//, concinnum Benth. Hypericum sect. 9. Hypericum sensu N. Robson in Bull. Br. Mus. nat. Hist. (Bot.) 5: 320 (1977) pro parte, quoad H. concinnum. Perennial herbs or sometimes suffrutices up to 0.45 m tall, with stems erect or sometimes ascending from woody taproot or caudex, glabrous, with dark glands; branching lateral, from lower nodes. Stems 4-lined when young, becoming 2-lined to terete, eglandular. Leaves opposite, decussate, sessile to shortly petiolate, free, tardily deciduous, often conduplicate, sometimes falcate; lamina entire with venation pinnate, closed, with tertiary reticulation obscure; laminar glands pale and sometimes black, punctiform, equal; mar- ginal gland dots dark, dense, small; ventral resin glands absent. Inflorescence \-c. 17-flowered with branching mostly or wholly monochasial, terminal, sometimes with subsidiary branches from 1- 2 nodes below; bracts and bracteoles reduced, persistent. Flowers stellate, homostylous. Sepals 5, free, persistent, erect in fruit, with margin entire to eroded-denticulate; veins 7-11; laminar glands pale, linear; marginal or submarginal glands few, dark or absent. Petals 5, persistent, with apiculus absent or almost so; margin entire or undulate-crenate; marginal glands absent or few, dark; laminar glands pale, linear to striiform. Stamen fascicles 5, united 2+2+1 (i.e. '3'), persistent, with stamens totalling 40-c. 100; central 4 filaments united above the base, the rest at the base; anthers yellow, gland amber; pollen type X. Ovary with central column and 3 axile placentae, <-ovulate; styles 3, divergent; stigmas narrowly capitate. Capsule 3-valved, subcoriaceous?, with valves narrowly longitudi- nally vittate. Seeds cylindric, not carinate or winged or terminally expanded; testa minutely shallowly pitted. BASIC CHROMOSOME NUMBER (X). 8, ploidy 2. HABITAT. Dry slopes and ridges; 390-900 m. DISTRIBUTION. U.S.A. (northern California) 1 species. 1. Hypericum concinnum Benth., PI. Hartw.: 300 ('1848', i.e. 1849?); Coulter in Bot. Gaz. 11: 108 (1886), in A. Gray, Syn.fl. N. Amer. 1: 289 (1897); Greene, Fl. francisc.: 112 (1891); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 211 (1893), op. cit. 2nd ed. 21: 177 (1925); McMinn, Man. Calif, shrubs: 351, f. 403 (1939); Abrams, Ill.fl. Pacific States 3: 1 16, f. 3235 (1951); Jepson, Man. Fl. Pis Calif.: 638, f. 631 (1951); Munz & Keck, Calif. FL: 192, f. 27 (1959), Suppl. (Munz): 18 (1968); Rickett, Wild Fls of U.S. 5: 160, t. 52 (1971?); Kozloff & Beidleman, Pis San Francisco Bay Region: 168, t. 32 (1994); N. Robson in 62 N.K.B. ROBSON Map 6 Sect. 9a: 1. H. concinnum specimens, other record O. Cullen et al., Eur. Gdn Fl. 4: 58 ( 1 995). Type: U.S.A., California, 'in valle Sacramento', Hartweg 1670 (K!-holotype; BM!, G!, GH!, W!). Fig. 9D, Map 6. H. bracteatum Kellogg in Proc. Calif. Acad. nat. Sciences , 2nd ed. 1: 67 (1873), non Buch.-Ham. ex D. Don (1825) (nom. illegit.). Type: U.S.A., California, Yuba Co., Marysville (fl), E. W. Garvitt s.n. (CAS-holotype). H. seleri R. Keller in Bot. Jb. 58: 192 (1923). Types: U.S.A., California, Marin Co., 'macchia am Abhang des Mount Tamalpais', C. & E. Selers.n. (Bf-indicated lectotype); [Amador Co.?] 'im Tal des San Stanislaus River', A. Stiibel s.n. (Bf- syntype). Icon: Rickett, Wild Fls of U.S. 5: 160, t. 52 (1971). Perennial herb or sometimes suffruticose, 0.1 5-0. 3 3 (-0.45) m tall, bushy, erect or more rarely ascending from woody caudex, rarely rooting at the base, branching at most nodes, with branches strict. Stems 4-lined and ancipitous at first, eventually terete, eglandular; internodes 3-10 mm, shorter than leaves. Leaves sessile to shortly petiolate (to 0.5 mm), tardily deciduous; lamina 13-32 x 1.5-8 mm, narrowly elliptic or narrowly oblong to linear (l:b = 4-10), concolorous, glaucescent?, often conduplicate and sometimes falcate, subcoriaceous; apex acute to subacute, margin entire, base cuneate; venation: 2-3(4) pairs of main laterals from lower 2/5 of midrib, with tertiary reticulation obscure and slightly prominent or invis- ible; laminar glands punctiform, small, pale and sometimes a few black; intramarginal glands black, spaced (10-12). Inflorescence 1- c. 17-flowered, terminal, often with flowering branches from 1-2 nodes below, the whole then cylindric; pedicels 1-3.5 mm; upper leaf pairs bracteose; bracteoles lanceolate to linear-elliptic, entire. Flowers 20-35(^0) mm in diam., stellate or reflexed; buds ovoid- cylindric to ovoid, acute. Sepals unequal, imbricate, 6-9 x 2-3 mm, broadly to narrowly ovate, acute to acuminate, entire or eroded- denticulate; veins 9-11, branching; laminar glands pale, linear to striiform, and rarely 1-2 black, punctiform; marginal glands black, few or absent. Petals golden yellow, not red-tinged in bud, (10-) 12- 15 x 4-5 mm, c. 1 .7-2 x sepals, obovate to oblong-obovate, apiculus absent or almost so, margin entire or undulate-crenate; laminar glands pale, linear to striiform; marginal glands black, numerous, sessile or impressed. Stamens 40-80(-100?), '3 '-fascicled, longest 8-10(-12) mm, c. 0.8 x petals; anther gland amber. Ovary 3-locular, 3_4 x 1 .8-2 mm, narrowly ovoid; styles 3, 6-9(-12) mm, 2-2.7(-3) x ovary, free, divergent; stigma narrowly capitate. Capsule 6-9 x 4- 4.5 mm, narrowly ovoid, about equalling sepals; valves longitudinally vittate. Seeds dark brown, c. 1 .5 mm long, cylindric, not carinate or appendiculate; testa minutely and shallowly pitted. 2n = 16 (Raven, Kyhos&Hill, 1965). Dry slopes and ridges, Yellow Pine (Pinus ponderosa) forest, chaparral; 390-600(-900) m. U.S.A. (California - Sierra Nevada from Mariposa Co. north to Shasta Co., Coastal Ranges from Marin Co. to Mendocino Co.). U.S.A. California: Marin Co., Mt Tamalpais, 600 m, March 1930 (1. fr), Meebold9798 (K); San Mateo Co.?, near San Francisco, 1866 (fl), Kellogg STUDIES IN HYPER1CUM 63 s.n. (BM, G, W); Napa Co., junction of State highways 37 and 128, 30 May 1957 (fl), Alava 989 (W); Mendocino Co., Coastal R.[ange], Eel R., 540-600 m, June 1894, Purpus 1148 (K); Amador Co., New York Falls, 600 m, May 1891 (fl), Hansen 32 (BM, G, JE, K); Butte Co., Paradise, 27 May 1928 (fl), Heller 14560 (G); *Yuba Co., Marysville, n.d.? (fl), Garvitt s.n. (CAS). CULTIVATED. U.S.A. California: Rancho Santa Ana Bot. Gdn [seed ex Marin Co., Mt Tamalpais, Matt Davis Trail, 390 m T coll. Munz], 22 June 1 955 (fl), Balls 10608 (BM). H. concinnum is the only derivative of sect. Roscyna that is di- rectly related to H. ascyron subsp. gebleri. Unlike nearly all the other derivatives (in sects 9, 9b-d), it has relatively large imbri- cate sepals and an amber connective gland, characters which, along with its long narrow, usually conduplicate leaves and bushy habit, prompted its removal from its original position in sect. Hypericum (Robson, 1977) to a separate section (Robson, 1981). Its isolated geographical location gives added support to this sys- tematic decision; together these facts suggest that it or its ancestor might even have become distinct from H. ascyron in Siberia, before crossing to North America. In any case, it does not seem to be directly related to the eastern North American H. ascyron subsp. pyramidatum. [For Sect. 9b. Graveolentia, see Part 4(3).] Sect. 9c. SAMPSONIA N. Robson sect, nov., section! 9d. Elodeoidibus affinis sed foliis perfoliatis, capsulae valvis glandulis succiniis ovoideis vel plusminusve elongatis instructis, inter alia differt. A sectioni 8. Bupleuroides foliis lanceolatis vel oblongis vel oblanceolatis (haud ovatis vel oblongo-ovatis), sepalis oblongis semper integris, stylis e baso decurvatis, capsula vittis glandiformibus, inter alia differt. Hypericum sect. 9. Hypericum sensu N. Robson in Bull. Br. Mus. nat. Hist. (Bot.) 5: 320 (1977) pro parte, quoad H. sampsonii. Perennial herbs or rarely suffrutices up to 0.8 m tall, with stems erect or basally decumbent from sometimes woody base, glabrous, with dark glands; branching lateral, from upper or most nodes. Stems terete, eglandular. Leaves opposite, perfoliate, united, persistent; lamina entire with venation pinnate, closed, with tertiary reticula- tion rather lax; laminar glands pale or very rarely some black, punctiform, unequal; intramarginal gland dots dark; ventral resin glands absent. Inflorescence c. 1 2^40-flowered with branching mostly dichasial, from 2 nodes, with subsidiary branches from up to 6 nodes below; uppermost bract pair and bracteoles reduced, decidu- ous, other bracts foliar, persistent. Flowers substellate with cupuliform base, homostylous. Sepals 5, free, persistent, erect in fruit, with margin entire; laminar glands pale and sometimes dark, striiform to punctiform; intramarginal to marginal glands dark. Petals 5, persistent, erect after flowering, without apiculus; margin entire or with prominent glands 4 ; marginal glands dark, immersed or prominent; laminar glands pale, shortly striiform to punctiform, and very rarely also dark, punctiform. Stamen fascicles 5, united 2+2+1 (i.e. '3'), persistent, with stamens totalling 15^*2; filaments basally united; anther gland dark; pollen type X. Ovary with 3 completely or incompletely axile placentae, each -ovulate; styles 3, free, outcurving from base; stigmas broadly to narrowly capitate. Capsule 3-valved, chartaceous?, with valves bearing punctiform or elongate vesiculate amber glands. Seeds cylindric, not carinate or appendiculate; testa finely ribbed-scalariform. BASIC CHROMOSOME NUMBER (x). Unknown. HABITAT. Thickets, streamsides, grasslands and marginal areas; 1 10-1700 m. DISTRIBUTION. Southern Japan, Taiwan, central and south China, north Vietnam, central Myanmar, India (Meghalaya). 2 species. Key to sect. 9c. Sampsonia 1 Common leaf base expanded; stamens c. 0.5 x petals; styles c. 2mm, c. 0.65 x ovary; ovary 3-locular 1 . sampsonii Common leaf base narrowed; stamens c. 0.9 x petals; styles c. 1mm, c. 0.35 x ovary; ovary 1-locular 2. assamicum 1 . Hypericum sampsonii Hance in J. Bot. Lond. 3: 378 ( 1 865), op. cit. 8: 275 (1870) ['sampsoni']; Maxim, in Bull. Acad. Imp. Sci. Saint-Petersbourg 12: 60 (1867); Franchet & Sav., Enum. PI. Jap. 1: 55 (1874), op. cit. 2: 298 (1875); Maxim, in Bull. Acad. Imp. Sci. Saint-Petersbourg 27: 431 (1882), Mel. Biol. 11: 165 (1882); Forbes & Hemsley in /. Linn. Soc. Lond. 23: 74 (1886); Henry, List PL Formos.: 19 (1896); Matsum. & Hayata, Enum. pl.formosan: 43 ( 1 906); H. Leveille in Bull. Soc. Bot. France 53: 500 (1906), op. cit. 54: 593 (1908); Gagnepain in Lecomte, Fl. Indo-Chine, 1: 285 (1909); Hayata, Icon, pl.formosan. 1: 80 (1911); Dunn & Tutcher in Bull. Misc. Inf. Kew, Addit. ser. 10: 4 1 (1912); Matsum., Indexpl.jap. 2: 369(1912);MakinoinIimuna, Somoku-Dzusetsu 14: t. 30 (1912); Makino & Nemoto, Fl. Jap.: 545 (1925), op. cit., 2nd ed.: 752 (1931); R. Keller in Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 180 (1925); Sasaki, List PI. Formosa: 295 (1928); Hand.-Mazz., Symb. Sin. 7: 404 (1931); S. Suzuki in Masamune, Short fl. Formosa: 141 (1936); Y. Kimura in Bot. Mag. (Tokyo) 54: 86, f. 7 (1940), in Nakai & Honda, Nov. fl. jap. 10: 127, ff. 45^6 (1951); Ohwi, Fl. Japan: 780 (1953), Engl. ed.: 631 (1965); Steward, Man. Vase. Pis Lower Yangtze Valley China: 259 (1958); Lauener in Notes Roy. Bot. Card. Edinburgh 27: 5 (1966); Hatusima, Fl. Ryukyus: 416 (1971); Anon. In Iconogr. Cormoph. Sinicorum 2: 880, f. 3490 (1972); N. Robson in Fl. Taiwan 2: 643, t. 433 (1976), 2nd ed. 2: 710, t. 334 (1996); Momiyama in Satake et al.., Wildfl. Japan 2: 1 15, t. 1 10 f. 3 (1982); Li Xiwen in Fl. R. P. Sinicae 50(2) 60, t. 8 ff. 4-7 (1990). Type: China, Guangdong, near Lukpo, c. '100 mill. Pass.' W. of Canton [Guangzhou], June 1865 (fl & fr), Sampson s.n. (BM!-holotype). Fig. 11 A, Map 7. H. electrocarpum Maxim, in Bull. Acad. Imp. Sci. Saint-Petersbourg 12: 62 ( 1 867), Mel. Biol. 6: 26 1 ( 1 867); Hemsley in J. Bot., Lond. 5: 207 (1876); R. Keller in Bot. Jb. 33: 554 (1904), op. cit. 44: 49 (1909), in Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 180 (1925). Type: Japan, Kyushyu, circa Nagasaki, in fruticetis montium, pluribus locis, 1 863 (fl & fr), Maximowicz s.n. (LE- holotype; BM!, K!). H. electrocarpum forma parvifolium R. Keller in Bot. Jb. 44: 49 (1909) [parvifolia]. Type: Japan, Kyushu, Fukuoka, in insula Oshima [Tai-kiu], July 1900 (fr), Faurie 3872 (Z-holotype; BM!). H. esquirolii H Leveille in Repert. Sp. Nov. Regni Veg. 6: 330 ( 1 909), Fl. Kouy-Tcheou: 198 (1914). Type: China, Guizhou, Kweichow, 6 July 1905, EsquirolSll (E-holotype). 'Kimura ( 195 1) uses 'Petals with fringe of hairs' (i.e. glandular hairs) and Momiyama (1982) uses 'Petals with raised fissures' (raised veins?) to separate H. sampso, of the Japanese sect. Hypericum, as well as the fruit character. 64 N.K.B. ROBSON Fig. 11 A. H. sampsonii: (a) habit; (b) sepal; (c) petal; (d) capsule. B. H. assamicum: (e) habit; (f) sepal; (g) petal; (h) capsule (a, e x 2/3; rest x 6). A. Fan 39, except (d) Faurie 175. B. Simon 180. STUDIES IN HYPERICUM 65 Map 7 Sect. 9c: 1. H. sampsonii specimens, other record O; 2. H. assamicum H. oshimaense R. Keller in Bot. Jb. 58: 194 (1923). Type as for H. electrocarpum forma parvifolium. Icon: Y. Kimura in Nakai & Honda, Nova fl. Jap. 10: 128, f. 45 (1951); Li Xiwen in Fl. R. P. Sinicae 50(2): 46, t. 8 ff. 4-7 (1990). Perennial herb 0.2-0.8 m tall, erect from decumbent rooting base, with stems single or few, branched above or almost throughout, with branches curved-ascending. Stems terete, eglandular; internodes 20-85 mm, exceeding leaves or shorter than them. Leaves in perfoliate pairs; lamina (20-)25-70(-80) x (7) 10-35 mm, broadly or narrowly lanceolate to oblong or oblanceolate, paler beneath, not glaucous, plane, chartaceous; apex obtuse to rounded, margin entire, common base somewhat expanded, rounded; venation: 4-5 pairs of main laterals from lower half to third of midrib, branching and uniting near margin, with tertiary reticulation not prominent, rather lax; laminar glands all pale to mostly black, punctiform, dense; intramarginal glands black, dense. Inflorescence c. 20-40-flowered from 2 nodes, corymbiform, with flowering branches from up to 6 nodes below and uppermost pair sometimes overtopping terminal inflorescence, the whole corymbiform to subpyramidal or cylindric; pedicels 2-3 mm; uppermost bract pair and bracteoles linear-lanceo- late to linear, deciduous, other bracts foliar, persistent, all entire. Flowers 6-10(-15) mm in diam., substellate with cupuliform base; buds ovoid, obtuse. Sepals 5, 3-7(-10) x 1-3 mm, free, unequal, erect in bud and fruit, oblong to oblong-spathulate or linear-oblong, rounded, entire; veins 5(3), branching outward distally; laminar glands numerous, pale and rarely black, striiform to punctiform; intramarginal glands irregular, black, rarely absent. Petals 5, bright yellow, not red-tinged in bud, 4-8(-13) x 1.5^(-7) mm, elliptic- oblong, rounded, margin entire or subentire; laminar glands pale (very rarely a few black), shortly striiform to punctiform; marginal glands black, sessile to subsessile. Stamens 30-42, longest (2-)3-4 mm, c. 0.5 x petals; anther gland black. Ovary 3-locular, 2.5-3 x 1.5-2 mm, ovoid to narrowly pyramidal; styles 3, c. 2 mm, c. 0.65 x ovary, free, outcurving; stigmas broadly capitate. Capsule 6-9 x 4- 5 mm, broadly ovoid to broadly or narrowly ovoid-pyramidal, exceeding sepals; valves with scattered ovoid to elongate amber vesicular glands. Seeds orange-brown, c. 1 mm long, cylindric, not carinate or appendiculate; testa finely ribbed-scalariform. 2n = ? Thickets, streamsides, grassy places, roadsides and cultivated mar- gins; 110-1 700m. China (eastern Gansu to Jiangsu southward, including Hong Kong), Taiwan, Japan (Kyushu, south Honshu), Vietnam (extreme north), Myanmar (Shan States). CHINA. Shaanxi: Ziyang Xian, 700 m, 10 August 1959 (fr), Li P.Y. 6959 (KUN). Gansu: Wen Xian, Fanba Gongshe, Heiyingou, 820 m, 19 August 1976 (fr), Yang J.X. et al. 3778 (IBSC). Henan: Shangchen Xian, Fushan Xiang, Yuzidian, 110m, 16 April 1959 (st). Anon, in Herb. Beijing Inst. Bot. 12 (PE). Jiangsu: Bau Hwa Shan [Baohua Shan], 450 m, 7 June 1922 (fl), Merrill 1 1486 (GH, K, NY), ibid.. Steward 2120 (K); Yixing Xian, Longchi Shan, 25 June 1962 (fr), Mao S.H. 106 (NAS). Anhui: Chien Shan Hsien, Tien Chu Shan, 200 m, 1 1 June 1936 (fl), Fan & Li 39 (BM); Taiping Xian, Qidu Gongshe, Longguang Dadui, 400 m, 19 June 1959 (fl & fr), Anhui Exped. 822 (IBSC). Zhejiang: Hangzhou, 30 May 1957 (fl), Chang S.K850 (PE); Tianmu Shan. 1 September 1984 (fl), He X.K29123 (IBSC). Fujian: Kushan, near Foochow, 500 m, 10 August 1905 (fr), Chung 3738 (SING); Dehua Xian, 700 m, 4 July 1959 (fr), Huang S.M. 190743 (IBSC). Jiangxi: Kiennan Distr., Sai Hang Cheung, near Tung Lei village, 1-29 August 1934 (fr), Lau 4102 (BM, GH); Guixi Xian, Lengshui Xiang, 350 m, 21 July 1958 (fr), Nie & Lai 3685 (KUN). Guangdong: Yang Shan, S. of Linchow, July- September 1932, Tsui 429 (K);Fengchuan Xian, Sike Xiang, 31 May 1958(fl 6 fr), Wand S. 164186 (KUN). Guangxi: Ch'uan Distr., Pai-yun-an and vicinity, 3 June 1937 (fl & fr), Tsang W.T. 27599 (US): Lingui Xian, Liangfeng,Yanshan, lOApril 1 950 (R\TsoongC.H. 808 129 (IBSC). Hunan: inter urbe Hsinhwa et Pantjing, inter vicos Dawan et Gwantjiling, 200-400 m, 30-31 May 1918 (fl), Hand.-Mazz. 11981 (K); Heng Shan, Fangguang Temple. 450 m, 5 June 1943 (fl), Chun S.H. 3510 (IBSC). Hubei: Ichang, 1885? (fl & fr), Henry 585 (BM, K, US); Zhuxi Xian. 890 m, 13 September 1 959 (fr). Li P. Y. 1 1 1 35 (KUN). Sichuan: Emei Xian, Emei Shan, 1 300 m, 10 66 N.K.B. ROBSON July 1941, Feng W. P. 17249(A,IBSC);LeiboXian,Xining214Chang. 1500 m, 13 July 1983 (fir), Cao & He 120957 (SZ); Tianqan Xian, 910 m, 10 June 1982 (fl), Peng D.Y. 45433 (CDBI). Guizhou: au-dessus de Chong kai, 25 July 1911 (fr), Esquirol 3030 (BM); Anlong Xian, Shuangjiang Gongshe, 900 m, Anshun Exped. 258 (KUN). Yunnan: Nou-yon-se, 27 May 1904 (fl & fr), Cavalerie 2030 (K): Suijiang Xian, 850 m, 29 May 1973 (fl & fr), Sun B.S. etal. 5 15 (KUN). TAIWAN. Taipei: in littore Tamsui, 22 May 1903 (fr), Faurie 175 (BM,W); Dandan, Kiirun, 17 May 1930 (fl & fr), Suzuki 4469 (TAI). Taoyuan: Nankan, Toen, 5 May 1929 (fl & fr). Kudo 575 (TAI). JAPAN. Kyushu: circa Nagasaki, June 1899 (fl), Faurie 3084 (BM); Todoroki valley, 10 May 1933 (st), 16 June 1935 (e. fl), Greatrex s.n. (K). VIETNAM. Bac Phan [Tonkin]: prov. de Hoa-binh, a Kim-boi, April 1926 (fl), Colani in Petelot s.n. (K); Tu-Phap, May 1 888 (e. fr), Balansa 3774 (K, P*). MYANMAR. S. Shan States: Mong Noi, Kengtawng, banks of Salween R. below flood level, 210 m, 7 March 1911 (fl), Robertson 253 (K). CULTIVATED. England: Kew, ex China, Jianxi, 14 July 1983 (1. fl), K. Ace. 467-82-04934 (K). Hance (1865) and apparently all succeeding authors placed this species in sect. 13. Drosocarpium, on the basis of its gland-dotted capsule valves, despite the large morphological and geographical differences between it and all the species in that mainly south-east European section. Specialisations aside, H. sampsonii is morpho- logically nearest to H. przewalskii (sect. Roscyna)', the perfoliate leaf pairs and glandular-punctate capsule valves would be anoma- lous in sect. Hypericum. It seems appropriate, therefore, to place this species and its very close relative, H. assamicum, in a separate section directly derived from H. przewalskii. 2. H. assamicum S.N. Biswas in Webbia 25: 671 (1971), inSharma & Sanjappa, Fl. India 3: 52 (1993). Type: India, Meghalaya, Sylhet, Nowgong, March 1848 (fl & fr), Simons s.n. (CAL- holotype and isotype). The Kew and/or Bogor specimens cited below are very likely part of the same Simons collection. Fig. 1 IB, Map 7. H. sampsonii sensu Dyer in Hook, f., Fl. Brit. India 1: 255 (1875) pro parte excl. typum. Icon: Biswas in Webbia 25: 672 (1971). Perennial or sometimes suffruticose herb 0.2-0.3(-0.4) m tall, erect, with stems often many from stout woody base. Stems terete; internodes 2-6.5 mm, shorter than to exceeding leaves. Leaves in perfoliate pairs; lamina 15-50 x 6-15 mm, oblong to oblanceolate, often suddenly somewhat narrowed below middle, glaucous be- neath, plane, thinly chartaceous; apex obtuse to rounded, margin entire or rarely 'glandular-crenulate', base broadly cuneate to rounded, common base narrowed; venation: 2-3(4?) pairs of main laterals usually from lower third to half of midrib, branching and uniting near margin, with tertiary reticulation not prominent; laminar glands black, punctiform, dense; intramarginal glands black, dense. Inflorescence c. 12-18-flowered from 2 nodes, corymbiform, with flowering branches from up to 2 nodes below, the whole then corymbiform to subpyramidal; pedicels 1-10 mm; uppermost bracts and bracteoles linear-subulate, other bracts foliar. Flowers c. 12 mm in diam., apparently stellate; buds ovoid-ellipsoid. Sepals 5, 5-8.5 x 1-2 mm, free, unequal, erect in bud and fruit, 3 larger oblanceolate - spathulate, 7-8.5 x 2 mm, 2 smaller oblanceolate to narrowly oblong, 5-5.5 x 1 mm, obtuse to acute, entire or with subapical marginal glands prominent; veins 3, midrib branched; laminar glands rather few, pale and sometimes black, punctiform; intramarginal to marginal glands irregularly spaced, black. Petals 5, 'yellowish', not tinged red in bud, c. 5 x 1 .7 mm, oblanceolate-spathulate, subobtuse or obtuse, margin entire? or with marginal glands prominent; laminar glands pale, striiform to punctiform, and a few black, punctiform; intramarginal to marginal glands black, immersed to sessile. Stamens c. 15, longest 4-5 mm, equalling petals or almost so; anther gland black. Ovary 1-locular with intrusive placentae, c. 2.7 x 2 mm, ovoid; styles 3, c. 1 mm, c. 0.35 x ovary, free, erect; stigmas capitellate. Capsule 5-6 x c. 4 mm, cylindric-subglobose, about equalling sepals, valves with scattered punctiform amber vesicular glands. Seeds reddish brown, c. 0.8 mm long, cylindric, not carinate or appendiculate; testa finely ribbed-scalariform. 'Jungles'; lowland. India (Meghalaya). INDIA. Meghalaya: Nowgong, 27 March 1885 (fl), Clarke 37671 (K); Nowgong, n.d. (fl & fr), Simons 180 (K); no loc., n.d. (fr), Simons s.n. (BO); Nowgong, March 1848, Simons s.n (CAL*); Mopung, July 1855, Anon. s.n. (CAL*); Nowgong (Bramaputra plains), Kurz s.n. (CAL*). H. assamicum is clearly closely related to H. sampsonii; the question is whether or not it is distinct. With two possible exceptions all its characters are derivative relative to those of H. sampsonii, and so it can be seen as a development from the south-eastward distributional 'arm' of//, sampsonii through Yunnan to south-eastern Myanmar on the Salween River. This 'outstation' of//, sampsonii is c. 850 km from the only locality of//, assamicum, in the Bramaputra plains of Assam. H. assamicum can be distinguished from H. sampsonii by the narrowed (not expanded) common leaf base, the longer stamens, the unilocular ovary with relatively and absolutely shorter styles and possibly the more clumped habit with woody base. Of these characters, only the narrowed common leaf base and longer stamens are possibly not derivative. Relative to H. sampsonii, it is therefore probably a neo-endemic. Sect. 9d. ELODEOIDA N. Robson, sect, nov., sectioni 9. Hyperico similis, sed caulis teretibus eglandulosis, bracteis et bracteolis et interdum foliis superioribus glanduloso-marginatis et auriculis glandulosis instructis, differt; a sectioni 9e. Monanthema inflorescentia 1-c. 50-floribus, petalis margine integris vel glanduloso-ciliatis post anthesin erectis, foliis glandulis laminaribus maioribus densis, differt. Hypericum sect. Hypericum sensu N. Robson in Bull. Br. Mus. nat. Hist. (Bot.) 5: 320 (1977) pro parte quoad H. seniawinii et //. petiolulatum. Hypericum sect. Adenosepalum sensu N. Robson in Bull. Br. Mus. nat. Hist. (Bot.) 5: 335 (1977) pro parte quoad H. elodeoides et //. napaulense. Perennial herbs up to 1m tall, with stems erect to prostrate, creeping and sometimes branching at base, glabrous, with dark (black or rarely red) glands on leaves, sepals (rarely absent), petals and anthers; branching lateral, from various nodes. Stems (mature) terete or rarely (when slender) 2-lined, eglandular. Leaves opposite, dec- ussate, sessile or up to 10 mm pseudopetiolate, free, persistent; lamina entire or sometimes wholly or proximally gland-fringed, sometimes with gland-fringed auricles, with venation pinnate, closed and tertiary reticulation dense to lax; laminar glands punctiform, pale and/or black, relatively large (cf. sect. 9e), dense; marginal glands dark, dense to sparse; ventral resin glands absent. Inflorescence 1-c. 50- flowered, with branching dichasial and/or monochasial, from l-4(-6) nodes, often with subsidiary branches below; bracts and bracteoles entire to gland-fringed and often glandular-auriculate, reduced. Flowers stellate or rarely infundibuliform, homostylous. Sepals 5, free, persistent, erect or rarely spreading in fruit, with margin entire to glandular-ciliate; veins 3-5; laminar glands pale and/or dark, linear to punctiform; marginal and/or intramarginal STUDIES IN HYPERICUM 67 glands dark. Petals 5, persistent, erect after flowering, with or Petal laminar glands all or mostly black; leaves sessile, triangular-ovate without apiculus, margin entire, with or rarely without immersed or to ovate-oblong (l:b = 2-3.2) 4b. H. elodeoides subsp. wardii sessile dark glands, or glandular-ciliate; laminar glands pale and/or 3(2) Leaves sessile or with broad petio|e to , mm base subcordate . dark, linear to punctiform, or rarely absent. Stamen fascicles 5, amplexicaul to cuneate; sepals with marginal glands in a regular row; united 2+2+1 (i.e. '3'), persistent, with stamens totalling 9-c. 60; styles 2.5- 10 mm, c. 1.5-3 x ovary 1. H. seniawinii filaments basally united; anther gland dark; pollen type X. Ovary . , ,. N 1*1-11 c Leaves with petiole 1-1 Omm, base usually cuneate to angustate; sepals with 3(4) completely axile placentae, each o-few-ovulate; styles . - , *, with few irregularly spaced marginal glands or none; styles 1-2.5 mm, 3(4), free, divergent from discrete bases; stigmas narrowly capitate c 7 _, 3 x ovary (2 H p etiolu | atum) 4 to claviform. Capsule 3(4)-valved, chartaceous to papyraceous, with valves longitudinally vitiate. Seeds cylindric, not (rarely 4 < 3 > Styles 1.5-2.2 mm, 1-3 x ovary; leaves broadest at or below middle; scarcely) carinate or appendiculate; testa scalariform-reticulate to inflorescence from 2-3 nodes; stems erect to decumbent ... . . ,. ,, . ^ 2a. H. petiolulatum subsp. yunnanense scalanform or foveolate. BASIC CHROMOSOME NUMBER (x). 8; ploidy 2, 4. St >i les M " 5 mm ' f; ; 7 x ovar y ; lea , ves ^ roadest a < or al ^ ve middle; inflorescence usually from apical node only; stems decumbent to pros- HABITAT. Forests, forest margins, slopes, grasslands, roadsides, trate 2b. H. petiolulatum subsp. petiolulatum and sometimes damp meadows and ricefields; (1 18-)500-3600 m. 5(1) Leaves lanceolate to oblong-lanceolate or linear, 10-60 mm long, DISTRIBUTION. East and south-east Asia from China (Shaanxi, sometimes glandular-ciliate; styles (3- )4- 18 mm long, 2-7 x ovary .. Henan and Anhui southward) to north Vietnam, and west along the 6 Himalaya to Kashmir. Leaves broadly OV ate-oblong or broadly elliptic to suborbicular, 6-14 5 species (+ 2 subspecies). mm long* entire; styles c. 3 mm long, 1.5 x ovary 5. H. kingdom! Key to sect. 9d. Elodeoida 6(5) Styles 3-7 x ovary, 6-18 mm; pedicels 0.5-3(-6) mm; petal laminar glands pale; leaves obtuse to rounded 3. H. hengshanense 1 Sepals entire or occasionally with few glandular cilia; bracts without or with short glandular auricles; leaves all entire 2 Styles c. 2 x ovary, (3)4-8 mm; pedicels 3-12 mm; petal laminar glands all or mostly black; leaves usually acute Sepals regularly or rarely irregularly glandular-ciliate; bracts and some- 4a H elodeoides subsp. elodeoides times at least upper leaves glandular-ciliate and with glandular-ciliate auricles 5 j Hypericum seniawinii Maxim, in Bull. Acad. Sci. Imp. Saint- Id) Petal laminar glands pale; leaves pseudopetiolate or, if sessile, then Petersbourg 27: 434 (1882), Mel. Biol. 11: 169 (1882); H. narrowly oblong to oblong-lanceolate (l:b = 3-4) 3 Leveille in Bull. Soc. Bot. France 54: 595 (1908); Hand.-Mazz., Map 8 Sect. 9d: 1 . H. seniawinii specimens, other record O. Fig. 12 A. H. seniawinii: (a) habit; (b) sepal; (c) petal; (d) ovary; (e) capsule; (f) habit (narrow-leaved form). B. H. petiolulatum subsp. yunnanense: (g) habit; (h) sepal; (i) petal: (j) ovary; (k) capsule. C. H. petiolulatum subsp. petiolulatum: (1) habit; (m) petal; (n) ovary; (o) capsule (a, f , g x 2/3; rest x 6). A. H.H. Chung 2879, except (f) Ford 16. B. Maire 292, except (g) Maire 698. C. Kingdon Ward 13270, except (1) Stainton, Sykes & Williams 3374. STUDIES IN HYPERICUM 69 Symb. Sin. 7: 402 (1931); Lauener in Notes Roy. Bot. Card. Edin- burgh 27: 5 (1966); Li Xiwen in FL R. P. Sinicae 50(2): 66, t. 13 ff. 1-6(1 990). Type: China, E. China australior, n.d. (fl), Seniawin s.n. in Herb. Fischer (LE-holotype). Handel-Mazzetti (1931) noted that the type could not be found in Leningrad. Fig. 12A, Map 8. H. lateriflorum H. Leveille in Bull. Soc. Agric. Sarthe 39: 322 (1904), in Bull. Soc. Bot. France 54: 595 (1908), in Repert. Spec. Nov. Regni Veg. 6: 375 (1909), Fl. Kouy-Tcheou: 198 (1914). Type: China, Guizhou, environs de Kouy-yang [Guiyang], a Tchang-chao-se, 29 August 1899, Bodinier2108 (E-holotype). H. elodeoides sensu Hand.-Mazz., Symb. Sin. 7: 403 (1931) pro parte quoad spec. cit. (Wang 447). Icon: Li Xiwen in Fl. R. P. Sinicae 50(2): 63, t. 13 ff. 1-6 (1990). Perennial herb (0.15-)0. 3-0.6 m tall, erect from creeping, rooting and sometimes branching base, with stems usually single, some- times branched above, with branches virgate to curved-ascending. Stems terete, eglandular; internodes 20-55 mm, shorter than leaves or usually exceeding them. Leaves sessile or with broad 'petiole' to 1 mm; lamina (15-)20-50 x 5-13 mm, oblong to oblong-lanceolate, paler and rarely minutely papillose (Guangdong) beneath, not glau- cous, plane, chartaceous; apex obtuse to rounded, margin entire, base subcordate-amplexicaul to broadly or narrowly cuneate; vena- tion: 3(4) pairs of main laterals from lower third of midrib, with tertiary reticulation not prominent, dense; laminar glands pale, dense, rather large; intramarginal glands all black or the occasional one pale, dense. Inflorescence c. 5-50-flowered, from 1-3 nodes, densely subcorymbose to broadly pyramidal, with flowering branches from up to 9 nodes below (often with a 'gap' of sterile nodes), the whole then cylindric; pedicels 1-2 mm; bracts and bracteoles ovate to linear-lanceolate, entire or occasionally with basal glandular cilia or rudimentary auricles, persistent. Flowers 9-15(-c. 20) mm in diam., infundibulariform to stellate or recurved; buds ellipsoid, acute to obtuse. Sepals 5, 2.5-5 x 1-2 mm, subequal to equal, erect in bud and fruit, oblong-lanceolate, acute, entire or subentire; veins 5(3), unbranched or outer pairs partly united; laminar glands pale, linear to striiform; marginal glands all black or rarely a few pale, in regular or interrupted row, sessile or slightly prominent. Petals 5, bright? yellow, not tinged red in bud, 7-10 x 2-3 mm, 2-3 x sepals, narrowly oblong to narrowly oblanceolate-elliptic, margin subentire, laminar glands pale, striiform to punctiform or absent; marginal glands black, distal, sessile or more proximal immersed. Stamens 24-c. 50, '3 '-fascicled, longest 5-12 mm, slightly shorter than petals; anther gland black. Ovary 3-locular, c. 1.5-3 x 0.7-1.3 mm, narrowly ovoid; styles 3, (2.5-)4-10 mm, c. 1.6-3 x ovary, free, divaricate-incurved; stigmas narrowly capitate. Capsule c. 5-6 x 4- 5 mm, ovoid, exceeding sepals; valves with dense longitudinal vittae. Seeds yellowish brown, c. 0.5 mm long, scarcely carinate; testa finely foveolate. 2n = ? Slopes, grasslands and roadsides; (118-)500-1600(-2000) m. South-east China (Henan, Anhui and Zhejiang south to Guangxi and Guangdong). CHINA. Henan: Kikungshan, 9 August 1925 (fl), Steward 2728 (K). Anhui: Chiu Hwa [Jiuhua] Shan, 500 m, 30 July 1934 (fl & fr), Fan & Li 14 (NAS); Huang Shan, 4 August 1935 (fl), Liou & Tsoong 10992 (PE). Zhejiang: Tien- mu-shan, 2 October 1947 (fr), Y.W. Law 1 183 (K, PE); Longquan Xian, 1 100 m, 26 July 1930 (fl), Shan R..H. 5415 (KUN). Fujian: Yenping, Cha-ping, 730 m, 2 August 1924 (fr), H.H. Chung in Herb. Univ. Amoy 2879 (BM, K); Shanghang Xian, 1400 m, 20 October 1932 (fr), Ling Y. 4063 (PE); Jiangxi: Lu Shan, Xianrendong, 118 m, 12 July 1955 (fl), Li B.C. 085 (IBSC); Yongxiu Xian, near Sa-tiu-hong, 800 m, 2 1 August 1932 (fl), Tsiang Y. 10623 (IBSC, NY); Hong San, 920 m, June-July 1936 (fl), Gressitt 1651 (BM); prope oppidum Ningdu ad margines agrorum montis Wuhwa-schan, c. 800 m, 21-23 July 1921 (fl), Wang T.H. 447. Guangdong: Loh-Fan Shan, near Siu Liu waterfall, c. 1050m, August 1920(fl), Whiting s.n. (K);Renhua Xian, Changjiang Xiang, 1670 m, 26 August 1958 (fl), Tang L. 7211 (KUN). Guangxi: Rong Xian, Tiantangshan, Laoyachong to Tiesuoling, 880-1 160 m, 30 June 1956 (fl), Chun S.//.9713 (IBSC, KUN); Longshen Xian, Dadi Xiang, 2 July 1955 (fl), Guangfulingu Exped. 00614 (IBSC). Hunan: in monte Yun-schan prope urbem Wukang, 1200 m, 9 July 1918 (fl), Handel- Mazzetti 2543 (K); Chengbu Xian, Dongtouwan, 1 July 1959 (fl), Tarn P.C. 63643 (IBSC). Hubei: Tongshan Xian, 650 m, 1 1 November 1974 (fr), Dai L. Y. et al. 2520 (WH); no precise locality, 1 885- 1 888 (fl), Henry 7352 (GH). Sichuan: [Nanchuan Xian, 1200 m, 6 July 1957 (fl), Tsiung & Zhou 91838 (PE). This is probably an error for H. petiolulatum subsp. yunnanense}. Guizhou: environs de Kouy-Yang [Giuyang], a Tchong-tchao-se, 29 August 1 899, Bodinier 2708 (E*). H. seniawinii and the very closely related H. hengshanense appear to be south-eastern derivatives of the H. pedunculatum form of H. przewalskii. H. seniawinii differs from H. hengshanense in lacking glandular auricles on the leaves but sometimes it bears them on the bracts and bracteoles. The leaves are usually relatively narrow compared to those of//, hengshanense but in both species the leaves become markedly narrower towards the south of their respective ranges. Wang 447 (Jiangxi) is intermediate in having glandular- ciliate bracts and sepals and glandular-auriculate bracts, but the leaves all lack glandular auricles and all except the uppermost are shortly and broadly petiolate. For differences between it and H. petiolulatum, see the latter (p. 70). 2. Hypericum petiolulatum Hook. f. & Thomson ex Dyer in Hook, f., Fl. Brit. India 1: 255 (1874); H. Leveille in Bull. Soc. Bot. France 54: 594 (1908) pro parte quoad typum; N. Robson in Blumea 20: 261 (1972) excl. locc. Sumatra et Sabah, in Hara & Williams, Enum. Fl. Pis Nepal 2: 62 (1979); Robson & Long in Grierson & Long, Fl. Bhutan 1(2): 378 (1984); Li Xiwen in Fl. Xizangica 3: 279, t. 115 ff. 5-9 (1986), in Fl. R. P. Sinicae 50(2): 64, t. 14 f. 1-4 (1990); Biswas in Sharma & Sanjappa, Fl. India 3: 75 (1993), in Fl. West Bengal 1: 265 (1997); Mukherjee & Chaudhri in J. Econ. Taxon. Bot. 20: 125 (1996). Type: India, Sikkim, 2700 m, 1 August 1849 (fl), Hooker s.n. (K!-holotype). Fig.l2B&C, Map 9. H. thomsonii R. Keller in Bot. Jb. 33: 552 (1904). Type as for H. petiolulatum Hook. f. & Thomson non H. petiolatum Walter. Note that the specimens cited by Keller belong to H. subcordatum (R. Keller) N. Robson; see comments by Handel-Mazzetti (1931: 402). H. petiolatum sensu R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed., 21: 179 (1925), orth. mut. vice H. petiolulatum. Icones: see under subspecies. Perennial (or sometimes annual!) herb 0.1-0.5 m tall, erect or ascending to procumbent or prostrate from creeping and rooting base, with stems much branched, the branches curved-ascending to spreading or straggling, all or mostly flowering. Stems terete, slen- der, eglandular; internodes much longer than leaves. Leaves with petiole l-7(-10) mm; lamina 5-35(-52) x 3-10(-17) mm, oblong or lanceolate-elliptic to obovate or suborbicular, paler or glaucous beneath, chartaceous to submembranous; apex rounded to rarely obtuse, margin entire, base cuneate to angustate or more rarely rounded to subcordate; venation: 3 pairs of main laterals from lower third of midrib, with tertiary reticulation fine, rather dense, not prominent; laminar glands pale, rather large, fairly dense, usually prominent, rarely also 1-2 black; marginal glands black, dense especially distally. Inflorescence ( l-)5-28-flowered, from 1 -2 nodes, usually with long (l)3-7-flowered branches from up to 5 nodes 70 N.K.B. ROBSON Map 9 Sect. 9d: 2a. H. petiolulatum subsp. yunnanense ; 2b. H. petiolulatum subsp. petiolulatum O specimens, other record A. below, the whole narrowly to broadly pyramidal; pedicels 4-1 3 mm; bracts and bracteoles linear (or lower bracts foliar), entire or more rarely with scattered black marginal glands, sessile or on short cilia, and sometimes glandular auricles. Flowers 5-8 mm in diam., stellate; buds ellipsoid, obtuse. Sepals 5, 2.3-3. 2(-3.4) x 0.5-0.9 mm, equal to unequal, erect in bud, spreading in fruit, very narrowly oblong- lanceolate or rarely narrowly elliptic-oblong to linear, acute or rarely subacute, entire or occasionally sparsely glandular-ciliate (especially towards base); veins 3, unbranched; laminar glands pale or rarely black, linear to punctiform, variable in size and number; marginal or submarginal glands black or reddish, few (sometimes only apical) or absent. Petals 5, bright? yellow, not red-tinged in bud, 3-5.5 x 1-1.2 mm, c. 1.5 x sepals, narrowly oblong, acute, margin entire, laminar glands pale, 1-2, punctiform, or absent; marginal glands black, 1-2 near apex and occasionally elsewhere, sessile or immersed. Stamens (9-) 17-22, '3 '-fascicled, longest (2.5- )3^t.5 mm; anther gland black. Ovary 3(4)-locular, 1 .2-2 x ( 1-) 1 .5-2 mm, broadly to narrowly ellipsoid; styles 3(4), (0.5-)l-2.5 mm, free, outcurving; stigmas claviform. Capsule 3.5^- x 3.5 mm, broadly ovoid to orbicular, exceeding sepals; valves longitudinally vittate. Seeds yellow-brown, 0.5-0.6 mm, not carinate; testa densely but shallowly scalariform-reticulate. 2n = ? Mountain slopes, thickets and grasslands, stream banks, cliffs, roadsides, forest margins; (250-) 1200-3 100 m. China (west and south-west), Vietnam (north), Myanmar (north), India (Arunachal Pradesh, Benghal, Sikkim), Bhutan, Nepal (for records from the Philippines and Sabah see H. taihezanense, Part 4(3)). H. petiolulatum is closely related to H. seniawinii, of which it appears to be an upland derivative. It has two subspecies: subsp. yunnanense is confined to China and adjacent north Vietnam, whereas subsp. petiolulatum is distributed from Yunnan, Sichuan and Xizang along the Himalayan range to Nepal. H. petiolulatum subsp. yunnanense has been confused with smaller-flowered forms of//, seniawinii with narrow and/or cuneate- based leaves, but can be distinguished from them by the shorter styles and smaller, broader capsule and usually by the absence or paucity of marginal sepal glands (not in a continuous row). The gap in distribution between these taxa is small, but the above-mentioned differential characters do appear to hold. 2a. Hypericum petiolulatum subsp. yunnanense (Franchet) N. Robson in Blumea 20: 262 (1973); Li Xiwen in Fl R. P. Sinicae 50(2): 66 (1990). Type as for H. yunnanense Franchet. Fig. 12B,Map9. H. yunnanense Franchet in Bull. Soc. Bot. France 33: 437 (1886), PL delavay. : 103 ( 1 889); H. Leveille in Bull. Soc. Bot. France 54: 594 (1908). Types: China, Yunnan, in pratis humidis ad Song-pin, supra Tapin-tze, 18 August 1884 (fl & fr), Delavay 1943 (P!- lectotype, Robson 1973). The other two collections cited by Franchet belong to H. monanthemum. H. mairei H. Leveille in Repert. Spec. Nov. Regni Veg. 11: 298 (1912); Lauener in Notes Roy. Bot. Card. Edinburgh 27: 4 ( 1 966); non H. Leveille (1915). Type: China, Yunnan, Tong-Chuan, 2600 m, June 1910, Maire in Herb. Bonati 7492 (E!-holotype). H. pseudopetiolatum var. grandiflorum Pampanini in Nuovo Giorn. Bot. Ital. N.S. 17: 672 (1910). Type: China, Hubei, Jen-kai-kou, 1500 m, 10 July 1905 (fl), Silvestri 1492 (Fl-holotype, BM!-photo). H. centiflorum H. Leveille in Bull. Geogr. Bot. 25: 23 (1915). Type: China, Yunnan, plaine de Tong-Tchouan, 2500 m, August 19 1 2 (1. fl & fr), Maire s.n. (E-lectotype, Robson 1973; BM!); plaine et vallons a Tcha-Ho, 2600 m, July 1912 (fl & fr), Maire s.n. (E- syntype; BM!). Icones: none. For figures labelled as subsp. yunnanense by Li Xiwen (1973, 1990 - above) see subsp. petiolulatum. Stems erect to decumbent, rooting at base, branches curved-ascend- ing. Leaf lamina 15^0 x 6-16 mm, lanceolate or oblong to oblong-lanceolate (broadest at or below middle). Inflorescence from 2-3 nodes. Ovary 1.5-2.2 mm long; styles 1.5-2.2 mm, 1-1.3 x ovary. STUDIES IN HYPER1CUM 71 Grassy slopes, roadsides, cliffs, forest margins and grasslands; (350-) 820-3 100m. South China (Shaanxi and Henan south to Yunnan and Guangxi), north Vietnam. CHINA. Shaanxi: Zhouzhi Xian, Chenhe Xiang, 1570 m, 10 August 1958 (fr), Zhang X.M. 298 (IBSC); Yang Xian, 1700 m, 24 July 1952 (fl & fr), Fu K. T. 5182 (PE). Henan: Jigong Shan, 9 August 1925 (fl & fr), Steward 9835 (US). Fujian: Changting Xian, Sidu Gongshe, Louzi Ba, 500 m, September 1993, Meihuashan Exped. 35 (IBSC). Jiangxi: Fenyi Xian, Dagangshan, 350 m, 28 August 1985 (fl), Yao K. 9260 (A, CAS, NY). Guangxi: Huaping Forestry Area, 870 m, 14 August 1962 (fl), Yuan S.F. 5656 (IBSC); Lingyun Xian, 28 July 1937 (fl), Lau S.K. 28791 (A). Hunan: Sinning Hsien, Ma-Ling Tung, 600 m, 15 September 1935 (fl & fr), Fan & Li 492 (A, BM, W); Dongkou Xian, outskirts of Dongkou Zheng, 330 m, 1910 (fl & fr), Maire 1425 (NY); Yunshan to Yangjiapai, 4 September 1950 (fr), Chang H.T. 4448 (IBSC). Hubei: Badong Xian, Shennongjia, 2300 m, 23 July 1957 (fl), Fu & Zhang 1020 (PE); Shenlungkai, 1 850 m, 1 976 (fl & fr), Chow K.S. 76034 (A, BM, K, NY). Sichuan: Dujangyan Mun. (Guan Xian), Niangziling Mtn, upstream from Longxi on Longxi R., 1400 m, 4 September 1988 (fl), Boufford & Bartholomew 24593 (A*, BM, CAS*, PE*); Nan cuang hien, 1450 m, 15 September 1957(fr), Xung & Li 93496 (SZ); Emei Shan, 1700 m, 26 September 1935 (fl & fr), Tu T.H. 509 (IBSC, PE). Guizhou: Songtao Xian, vicinity of Lengjiaba near confluence of rivers Xiaohe and Dune, NE side of Fanjing Shan, 820-1120 m, 10 October 1986 (fr), Sino-American Guizhou Bot. Exped. 236 1 (A); near Leigongshan Muchang, 24 August 1 959 (fl & fr), S. Guizhou (Qiannan) Exped. 3535 (KUN). Yunnan: N. of Mengtze, 2400 m, n.d.(fl), Henry 10274 (K); Dali, Cangshan, 7 August 1922 (fl & fr), Rock 641 3 (A, NY, US); Binchuan Xian, Jizu Shan, 28 September 1940 (fr), Ching R.C. 24944 (KUN). VIETNAM. North: Tonkin, Chapa, c. 1500 m, August 1939 (fr), Petelot 2295 (GH). 2b. Hypericum petiolulatum subsp. petiolulatum Fig. 12C, Map 9. H. petiolulatum var. orbiculatum Franchet in Bull. Soc. Bot. France 33: 437 (1886), PL delavay.: 103 (1889); H. Leveille in Bull. Soc Bot. Fr. 54: 594 (1908). Type: China, Yunnan, ad juga montis Koua-la-po (Hokin) in humidis, 3000 m, 4 August 1885 (fl), Delavay 1942 (P!-holotype). H. humifusum subsp. orbiculatum S.N. Biswas in Bull. Bot. Surv. India 29: 53, f. 2 (1989), in Sharma & Sanjappa, Fl. India 3: 67, f. 24 (1993), in Fl. West Bengal 1: 264 (1997). Type: Nepal, near Tagat, 2550 m, 5 July 1954 (fl), Stainton, Sykes & Williams 3374 (CAL-holotype; BM!). Icones: Li Xiwen in F I. Xizangica 3: 280, 1. 1 1 5 ff. 5-9 (1986); in Fl. R. P. Sinicae 50(2): 65, t. 14 ff. 1-4 (1990); S.N. Biswas in Bull. Bot. Surv. India 29: 53, f. 2 (1989). Stems decumbent to prostrate, branches diffuse. Leaf lamina 5-25 x 4-1 1 (- 1 5) mm, oblanceolate to elliptic or orbicular (broadest at or above middle). Inflorescence usually from one node. Ovary 1.5-2 mm long; styles 1-1.5 mm, 0.65-0.75 x ovary. Mountain slope thickets and grassland; 2100-3000 m. China (Yunnan, Sichuan, Xizang), Myanmar (north), India (Arunachal Pradesh, Benghal, Sikkim), Bhutan, Nepal. CHINA. Sichuan: Mu li [Muli] hien, 2200 m, 30 August 1978 (fl & fr), Zhou C.S. et al. 8212 (SZ). Sichuan/Yunnan: Yangpi (Yungpei?) pass, 2490 m, 19 August 1922 (fl & fr), J.W.&J. Gregory s.n. (BM). Yunnan: Col de Kou-la- po (Hokin), 3000 m, 4 August 1885 (fl), Delavay 1962 (P). Xizang: Nyalam Xian, 2350 m, 16 August 1972 (fl & fr), Tibet Medicinal Exped. 1236 (PE); Cona Xian, 3000 m, 26 August 1975 (fl & fr), Qingzang Exped. 75-1594 (KUN, PE). MYANMAR. Nam Tamai valley, 28 N, 97 400 E, 1 200 m, 1 6 September 1937 (fr), Kingdon-Ward 13270 (BM); Nam Tamai valley (Adung Wang), 28 150 N, 97 350 E, 2100 m, 29 September 1937 (fr), Kingdon-Ward 13314 (BM). INDIA. Arunachal Pradesh: Delei valley, 28 150 N, 86 350 E, 3000 m, 23 August 1928 (fr), Kingdon-Ward 851 5 (K). Benghal: Darjeeling, Mukerjee et al. 645 (CAL*, DD*). Sikkim: Lachen, 2700 m, 1 August 1849 (fl), Hooker s.n. (K). BHUTAN. Tobrang, Trashi Yangsi Chu, 2100 m, 7 July 1949 (fl & fr), Ludlow, Sherriff & Hicks 20858 (BM, E*); Chenkaphug, E. of Thimpu, c. 3000 m, 20 July 1979 (fl), Grierson & Long 2799 (BM, E*). NEPAL. East: Milka Bhanjzang, 2550 m, 3 July 1969 (fl), Williams 1 100 (BM). Central: above Siklis (S. of Annapurna), 28 070 N, 84 060 E, 29 August 1976, Troth 999 (BM, US*). West: near Tagat, 2550 m, 5 July 1954 (fl), Stainton, Sykes & Williams 3374 (BM, CAL*). Map 10 Sect. 9d: 3. H. hengshanense A; Sect. 9d: 4a. H. elodeoides subsp. elodeoides : 4b. H. elodeoides subsp. wardii O, Sect. 9d: 5. H. kingdonii A. 72 N.K.B. ROBSON 3. Hypericum hengshanense W.T. Wang in Bull. Bat. Lab. N. E. Forest. Inst., Harbin, no. 5: 27 (1979). Li Xiwen in Fl. R. P. Sinicae 50(2): 59 (1990). Type: China, Hunan, Hengshan, 820 m, 20 July 1948 (fl), Y. Liu 109 (PE-holotype, BM!-photo; NAS!). Map 10. H. hengshanense var. xinlinense Z.Y. Li in Bull. Bot. Res., Harbin 8: 129, t. 2 (1988). Type: China, Hunan, Mons Ziyunshan, 1 100 m, 13 September 1984 (fl & fr), Ziyunshan Exped. 1789 (PE- holotype). Icones: W.T. Wang in Bull. Bot. Lab. N.E. Forest Inst., Harbin, no. 5: 27 (1979); Z.Y. Li in Bull. Bot. Res. Harbin 8: 132, t. 2 (1988). Perennial herb 0.62-1 m tall, erect from creeping and rooting base, with stems single?, shortly branched above, with branches virgate. Stems terete, eglandular; internodes c. 30-45 mm, shorter than leaves. Leaves sessile; lamina (15-)30-60 x (3-)7-16 mm, oblong- lanceolate to narrowly oblong-elliptic, paler beneath, not glaucous, plane, chartaceous; apex obtuse to rounded, margin entire or ('var. xinlinense^} black-glandular-ciliate to -denticulate, base slightly oblique-cuneate or (uppermost) rounded, upper pair(s) sometimes with black-glandular-fimbriate auricles; venation: 2(3) pairs of main laterals from lower quarter to third of midrib, with tertiary reticula- tion dense; laminar glands pale, scattered, large; intramarginal glands black, dense. Inflorescence 5-c. 18-flowered, from (1)2-3 nodes, subcorymbiform to broadly pyramidal; pedicels 0.5-3(-6) mm; bracts and bracteoles linear-lanceolate to linear, with margin and auricles black-glandular-fimbriate. Flowers 1 5-25 mm in diam., stellate; buds ellipsoid, obtuse. Sepals 5, 5-8 x 1.5-2 mm, equal, erect in bud and fruit, oblong-lanceolate to linear-oblong, acute, glandular-ciliate; veins 3, unbranched; laminar glands pale, linear to striiform; intramarginal glands few, distal, black, or absent; mar- ginal glands (to 16 on each side) on cilia, black. Petals 5, bright? yellow, not tinged red in bud, 9-15 x 2.5-3 mm, c. 2 x sepals, narrowly oblong, margin entire?, laminar glands pale, striiform to punctiform, marginal glands black, punctiform. Stamens , '3'- fascicled, longest 8-15 mm, 0.6-0.75 x petals; anther gland black. Ovary 3-locular, 2-2.5 x c. 1-1.5 mm, ovoid; styles 3, 6-18 mm, 3- 7 x ovary, divaricate-incurved, 'long-inserted' (Li Xiwen 1990); stigmas narrowly capitate. Capsule 5.5-6(-9?) x 3-3. 5(^?) mm, ovoid, exceeding sepals; valves with dense longitudinal vittae. Seeds not seen. 2n = ? Slopes, thickets and roadsides; 600-1 100 m. South China (Jiangxi, Guangdong, northern Guangxi, Hunan). CHINA. Jiangxi: sine loc., 600 m, 16 July 1964 (fl), Yang & Yao 1140 (NAS); Guangchang, Kuang Fang, Yashi Shan, 17 October 1962(fr), YueJ.S. 2523 (IBSC). Guangdong: Lianshan Xian, Shangshuai Xiang, 980 m, 1 1 July 1 958 (fl), Tarn P.C. 58761 (KUN). Guangxi: Waitsup Distr., Tong Shan, near Sap-luk Po village, 16 September 1933, Tsang 22802 (W); Yuangupo He, Linfan Shan, 30 August 1958 (1. fl), Li Y.KAOl 127 (IBSC). Hunan: Heng Shan, 1-6 June 1943 (st), Chen Z.D. 217 (IBSC); see also holotype and that of var. xinlinense. H. hengshanense is closely related to H. seniawinii, differing from it by the glandular-auriculate leaves and bracts, the glandular-ciliate sepals and the longer styles. Its very restricted distribution is wholly within that of H. seniawinii, and it is morphologically intermediate between that species and H. elodeoides. See discussion under 1 . H. seniawinii (p. 69). The Guangxi populations have smaller, relatively narrow leaves and smaller flowers. 4. Hypericum elodeoides Choisy in DC., Prodr. 1: 551 (1824); Dyer in Hook, f., Fl. Brit. India 1: 255 (1874); Franchet in Bull. Soc. Bot. France 33: 438 (1886), PL delavay.: 104 (1889); H. Leveille in Bull. Soc. Bot. France 54: 594 (1908) [helodeoides]; Burkill in Rec. Bot. Surv. India 4: 99 (1910); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 179 (1925); Banerji in J. Bombay Nat. Hist. Soc. 51: 774 (1953), op. cit. 55: 251 (1958), in Rec. Bot. Surv. India 19(2): 27 (1966); Y Kimura in Kara, Fl. E. HimaL: 209 (1966), op. cit. 2: 81 (1971); N. Robson in Nasir & Ali, Fl. W. Pakistan 32 (Guttiferae): 10 (1973), in J. Jap. Bot. 52: 285, excl. f. 3 (1977), in Hara & Williams, Enum. PL Nepal 2: 61 (1979); Robson & Long in Grierson & Long, Fl. Bhutan 1(2): 377 (1984); Li Xiwen in FL Xizangica 3: 279, t. 115 ff. 1-4 (1986), in FL R. P. Sinicae 50(2): 59, t. 1 1 ff. 3-7 (1990); Biswas in Sharma & Sanjappa, Fl. India 3: 56, f. 19 (1993), in FL Bengal 1: 262 (1997); Mukherjee & Chaudhuri in J. Econ. Taxon. Bot. 20: 125 (1996). Type: Nepal, no precise locality, 1821 (fl), Wallich 4812A (G-DC!-holotype; BM!, K!, SING!). Fig. 13A&B, Map 10. H. napaulense Choisy in DC., Prodr. 1: 552 ( 1 824). Type: Nepal, no precise locality, 1821 (fl), Wallich s.n. (G-DC! -holotype). H. nervosum D. Don, Prodr. Fl. Nepal.: 219 (1825), nom. illegit. superfl. Type as for H. elodeoides. H. pallens D. Don, Prodr. FL Nepal.: 219 (1825), nom. illegit. superfl., non Banks & Solander ( 1 794). Type as for H. elodeoides. H. adenophorum Wall. [Numer. List: 170, no. 4812] ex Dyer in Hook, f., Fl. Brit. India 1: 256 (1874) in synon. Icones: Li Xiwen in Fl.Xizangica 3: 280, 1. 115, ff. 1-4 (1986); in F/. R. P. Sinicae 50(2): 56, t. 1 1 ff. 3-7 (1990). Perennial herb 0. 1 5-0.5(-0.73) m tall, erect (sometimes from creep- ing and rooting base), with stems caespitose, unbranched or rarely branched above, with branches virgate Stems terete, eglandular; internodes 5-35 mm, usually shorter than leaves. Leaves sessile; lamina 10-50 x (2-)4-12(-17) mm, lanceolate or more rarely ovate- lanceolate to oblong-lanceolate or oblong-elliptic to linear, paler or glaucous beneath, plane or margins recurved, chartaceous; apex acute to subacute or more rarely obtuse to rounded, margin entire or (upper) glandular-ciliate towards base, base cordate-amplexicaul to rounded, the upper usually with glandular-ciliate auricles; venation: (2)3 pairs of main laterals from lower third to fifth of midrib, usually prominent beneath, with tertiary reticulation lax; laminar glands pale, dense, large, punctiform or shortly striiform; intramarginal glands black, sparse. Inflorescence (l-)5-c. 30-flowered, from 1(2- 4) nodes, corymbiform to cylindric, very rarely with flowering branches from 1-2 nodes below; pedicels 3-12 mm; bracts and bracteoles ovate-lanceolate to oblong-lanceolate or linear, with margin and auricles black-glandular-ciliate or very rarely subentire to entire without auricles. Flowers 10-20 mm in diam., stellate; buds ellipsoid. Sepals 5, 5-9 x l-1.5(-3) mm, equal, erect in bud and fruit, narrowly elliptic-lanceolate to narrowly oblong-lanceolate, acute, glandular-ciliate; veins 5, unbranched, prominent; laminar glands pale or black, linear to striiform; marginal glands on cilia and a few sessile, black. Petals 5, golden yellow, not tinged red in bud, 7-15 x 3-4 mm, oblanceolate to obovate-oblong, margin entire, laminar glands black and sometimes a few pale, linear to puncti- form, dense to sparse, marginal glands black, distal, few, sessile. Stamens c. 60, '3'-fascicled, longest 8-11 mm, c. 0.75 x petals; anther gland black. Ovary 3-locular, 2-4 x 1.3-1.7 mm, narrowly ovoid; styles 3, (3-)4-8 mm, c. 2 x ovary, divaricate, curved- ascending or straight; stigmas narrowly capitate. Capsule 5-8 x 4-5.5 mm, ovoid, about equalling sepals; valves longitudinally vitiate. Seeds yellow-brown, 0.5-0.6 mm, not carinate; testa densely scalariform-reticulate. 2n = 16 (n = 8; Sugiura, 1944), 32 (n = 16, Sandhu&Mann, 1989). STUDIES IN HYPERICUM 73 Fig. 13 A. H. elodeoides subsp. elodoides: (a) habit; (b) sepal; (c) petal; (d) capsule. B. H. elodeoides subsp. wardii: (e) sepal. C. H. wightianum: (f) habit; (g) leaf; (h) sepal; (i) petal; (j) capsule (a, f x 2/3; d, g, j x 4; b, rest x 6). A. Polunin, Sykes & Williams 448. B. Kingdon Ward 22780. C. Forrest 4298. 74 N.K.B. ROBSON Forests, forest margins and clearings, thickets, damp meadows and rice fields, grassy slopes and tracksides; (750?-) 1050-3600 m. South China (Guangxi, Sichuan, Yunnan, Xizang; also recorded from Guizhou, Guangdong, Hunan, Hubei, Jiangxi and Fujian), Myanmar (Mt. Victoria), India (Manipur, Meghalaya, Arunachal Pradesh, Benghal, Sikkim, Uttar Pradesh, Himachal Pradesh, Kash- mir), Bhutan, Nepal. H. elodeoides is closely related to H. hengshanense, but has shorter leaves and styles and is in general smaller. The main reduction trend in this species, in which the acute leaves gradually become linear, runs mainly east to west along the Himalayan range to Kashmir, omitting north Myanmar and most of Arunachal Pradesh. In a separate, southward trend through Manipur into western Myanmar, however, the leaves remain relatively broad (though smaller) and all become entire and rounded at the apex, and the sepals, bracts and bracteoles also become subentire or entire. These plants have been given subspecific rank as subsp. wardii. 4a. Hypericum elodeoides subsp. elodeoides, Biswas in Sharma & Sanjappa, Fl. India 3: 58 (1993). Fig. 13 A, Map 10. Leaves usually acute to subacute, prominently nerved beneath, with conspicuous pale laminar glands. Bracts and upper leaves glandu- lar-auriculate. Sepals and bracts with glandular-ciliate margin Distribution of the species (see above) except Manipur and Myanmar. CHINA. [See also general distribution] Guangxi: Damiaoshan Xian, Yuanbaoshan, 21 October 1958 (fr), Chun S.H. 16863 (KUN). Sichuan: S. Wushan, n.d. (fl), Wilson 1 389 (NY); Yuexi Xian, 3200m, 31 August 1976(fl 6 fr), Sichuan Plant Exped. 14230 (CDBI). Yunnan: au pied de Tsiang chan, au dessus de Ta-li, 2500 m, 25 September 1884, Delavay 189 (P); Menghwa [Weishan], Weipaoshan, 2300 m, 8 September 1933 (fl & fr), Tsiang 11386 (IBSC); Luoci Xian, Jiawoping, 2400 m, 23 October 1964 (fr), Qiu B.Y. 596341 (KUN). Xizang: W. of Yatung, 3000 m, 3 August 1936 (fl), Spencer Chapman 331 (K); Nyalam Xian, 2300 m, 13 August 1972 (fl), Xizang Exped. 108 1 (PE); Cona Xian, 2700 m, 4 October 1974 (fr), Qingzang Exped. 74-2773 (KUN, PE). INDIA. Meghalaya: Khasia, Lailankote, 1 650 m, 28 November 1 87 1 (fr), Clarke 14795 (BM); Arunachal Pradesh: Dirong Dzong, 1500-1800 m, 4 August \938(f\),Kingdon-Ward 1 4043 (BM). Benghal: *Darjeeling,Lebong CartRoad, \900m,MukherjeeA. 1 303 (CAL). Sikkim: Pemiongchi, 2 100m, 7 October 1875 (fr), Clarke 25094 (K); Changkyepyakop, 4200 m, 25 November 1911 (fr), Ribu & Remee 5718 (BM). Uttar Pradesh: Kumaon, Almora, 1500 m, n.d. (fl), Strachey & Winterbottom 2 (BM, GH, K); Tehri- Garhwal, below Mussoorie, 1950 m, September 1898, Gamble 27205 (K). Himachal Pradesh: Simla, Nugkunder, 2700 m, July 1885 (fl), Collett 763 (K); Dalhousie, 2100 m, 1 1 September 1874 (fl & fr), Clarke 22203 (BM). Kashmir: Basaoli [Basoli], 1 800 m, 26 September 1 876, Clarke 3 1 607 (BM); Budrawan [Bhadrawan], 1800 m, 23 September 1876 (fr), Clarke 31503 (K). BHUTAN. Central: Bumthang distr., Dhur, near Bumthang, 3000 m, 23 July 1949, Ludlow, Sherriff & Hicks 19503 (BM); Thimpu distr., Bele-dato Paro, 3600-2250 m, 9 July 1938, Gould965 (K). North: Upper Mo Chu distr., Gasa Dzong, 2800 m, 14 September 1 984 (fr), Sinclair & Long 4987 (E*, K). NEPAL. East: Tinjure Danda, 2700 m, 7 September 1967 (fl & fr), Williams & Stainton 8408 (BM); Chauki, 2500 m, 17 August 1972 (fl), Dobremez 1 5 1 (BM, GR*). Central: Godavari, Kathmandu Valley, 1 590 m, 20 August 1965 (fl). Schilling 591 (K); Annapurna Himal, Siti Khola, 2100 m, 5 August 1954 (fl), Stainton, Sykes & Williams 67 1 1 (BM). West: Bartadi, 1200m, 29 July 1953(fl),7>*i 135a(BM); Kaure/Sallyana, N. slope, 1560 m, 14 August 1969 (fl), Flatt 83 (BM). The Yunnan specimens sometimes have rounded leaves, but they are much longer than those of subsp. wardii. One specimen from Bhutan (Chenkaphung, E. of Thimpu, c. 3000 m, 20 July 1979 (fl & fr), Grierson & Long 2798 (BM, E*)) is morphologically intermediate between H. elodeoides and H. himalaicum and would appear to be of hybrid origin. 4b. Hypericum elodeoides subsp. wardii N. Robson in J. Jap. Bot. 52: 286 (1977); Biswas in Sharma & Sanjappa, FL India 3: 58 (1993). Type: Burma, Mt. Victoria, 2475-2700 m, 27 October 1956 (fl & fr), Kingdon Ward 22780 (BM!-holotype). Fig. 13B, Map 10. Leaves obtuse to rounded, not prominently nerved beneath, usually with obscure pale laminar glands. Bracts and upper leaves not glandular-auriculate. Sepals and bracts entire or subentire. India (Manipur), Myanmar (Chin). INDIA. Manipur: Sirhoi, 2400 m, 27 September 1948 (1. fl), Kingdon Ward 181 13 (A, BM, NY*). MYANMAR. Chin: Mt. Victoria, Esakan, 2100 m, 5 September 1956 (fl & e. fr), Kingdon Ward 22667 (BM). The Manipur specimen is somewhat intermediate between the two subspecies in leaf characters and in sometimes (Kingdon Ward 181 13 in A) having single glandular cilia on the sepals. 5. Hypericum kingdonii N. Robson, stat. et nom. nov. Type as for H. wightianum subsp. axillare N. Robson. Map 10. H. wightianum subsp. axillare N. Robson in J. Jap. Bot.: 287, ff. 3- 4 (1977); Li Xiwen. in Fl. Xizangica 3: 278, t. 113 ff. 1-2 (1986), mFl. R. P. Sm/ctf 50(2): 57 (1990); Biswas in Sharma & Sanjappa, Fl. India 3: 80 (1993). Type: China, Xizang, Rima [Zayii], 1800 m, 27 August 1950 (fr), Kingdon Ward 29161 (BM!-holotype). Icones: Li Xiwen in Fl. Xizangica 3: 277, t. 1 13 ff. 1-2 (1986). Perennial herb 0.15-0.48 m tall, suberect to decumbent from short rooting base, with stems usually branched from upper to nearly all nodes. Stems terete, eglandular; internodes 10-25 mm, longer than leaves. Leaves sessile, lower soon deciduous; lamina 6-14 x 3-7 mm, broadly ovate-oblong or broadly elliptic to suborbicular, pale or glaucous beneath, recurved, subchartaceous, apex rounded, mar- gin entire, base shallowly cordate to rounded; venation: 3-4 pairs of main laterals from lower 2/5 of midrib, tertiary reticulation lax or apparently absent; laminar glands pale, punctiform, small, dense; intramarginal glands black, dense. Inflorescence 5-25-flowered from 1-2 nodes, usually with flowering branches from up to 6 nodes below and often also towards base of stem, the whole narrowly pyramidal to subcylindric; pedicels 1-2 mm; bracts and bracteoles narrowly elliptic or linear-lanceolate, black-glandular-denticulate to -fimbriate or subentire, with intercalary sessile black glands and well developed black-gland-fringed auricles, persistent. Flowers c. 8-14 mm in diam., stellate; buds ellipsoid, obtuse. Sepals 5, 4.5- 6 x (1-) 1.5-2 mm, equal, erect in bud and fruit, lanceolate to oblong-linear, acute to subacuminate, irregularly glandular-denticu- late to -fimbriate or subentire; veins 3-5, branching; laminar glands pale, linear to punctiform and often a few black, punctiform; mar- ginal or intramarginal glands black, irregular, often between glandular cilia or fimbriae. Petals 5, golden? yellow, not tinged red in bud, 6- 8 x ? mm, c. 1 .3-1 .5 x sepals, narrowly ovate, acute, margin entire; laminar glands black, striiform to punctiform, sparse to rather dense; marginal glands distal and in apiculus, black. Stamens c. 20-25, '3'- fascicled, longest 5-6 mm, c. 0.7-0.85 x petals; anther gland black. Ovary 3-locular, c. 2 x 1 mm, ellipsoid; styles 3, c. 3 mm, 1.5 x ovary, divergent; stigmas narrow. Capsule 5-7 x 3.5^.5 mm, narrowly ovoid to ellipsoid, c. 1.1 x sepals; valves with numerous lingitudinal vittae. Seeds yellow-brown, c. 0.5 mm, not carinate; testa finely scalariform. 2n = ? Rice paddy bunds, grassy slopes; 1200-2700 m. STUDIES IN HYPER/CUM 75 BASIC CHROMOSOME NUMBER (x). Unknown. HABITAT. Usually damp or wet, e.g. forest and woodland clearings, meadows, rocky or grassy slopes, marshes, bogs; 2100-4800 m. DISTRIBUTION. South-west China (south Shaanxi, Sichuan, Yunnan, Xizang), Laos (north), Thailand (north), Myanmar (north), north India (Nagaland to Kashmir), Bhutan, Nepal, Pakistan, south India (Tamil Nadu), Sri Lanka. 7 species (+ 1 subspecies). Key to sect. 9e. Monanthema China (Xizang, Yunnan), Myanmar (Kachin), India (Arunachal Pradesh). CHINA. Xizang: Bomi Xian, Tongmi Cun, 2000 m, 22 July 1 965 (st), Chang & Lang 850 (PE); Zayu Xian, 1960 m, 29 July 1973 (fl & fr), Qingzang Exped. 73-932 (PE). Yunnan: Kiukiang Valley, Sochieh, 1700 m, 25 July 1938 (fl), YuT.T. 19392 (PE). MYANMAR. Kachin: Sources of the Irrawaddy, Adung Valley, 2100- 2400 m, 31 March 1931 (fr), Kingdon Ward 9352 (BM); North Triangle, Uring Bum above Ahkail, 2700 m, 8 November 1953 (fr), Kingdon Ward 21570 (BM). INDIA. Arunachal Pradesh: Delei Valley, 2100-2400 m, 15 August 1928 (fl), Kingdon Ward 8535 (K). My original association of this taxon with H. wightianum was clearly wrong, as is shown by the dense, relatively large leaf laminar 1 Sepal margin glandular-ciliate to entire, not glandular-denticulate; ovary glands (characteristic of sect. Elodeoida, not of sect. Monanthema) and capsule 3-locular; sepals never with glands between glandular and the larger flowers, which agree with those of H. wightianum emergences 2 essentially only in having sessile marginal glands in the sepals Sepal margin glandular-denticulate or ovary and capsule 1-locular between the (irregular) glandular cilia. These sessile glands, how- sepals often with sessile or intramarginal glands between glandular ever, are sometimes absent, resulting in an entire margin, which is emergences 7 unknown in H. wightianum. H. kingdonii is much better treated as a northern alpine relative of//, dodeoides subsp. wardii. 2(1) 2^5 VOld ( t sub ^ ose - styles 1.5--3 5(-4) mm; uppermost leaf pair foliose, entire or with few basal stalked glands 3 Sect. 9e. MONANTHEMA N. Robson, sect, nov., section! 9. Capsule ellipsoid or, if ovoid-ellipsoid to broadly ovoid, then styles 4- Hyperico similibus, sed caulis maturis teretibus vel ubi tenuibus 5 mm ; uppermost leaf pair bracteose 5 2-4(6)-lineatis, eglandulosis, bracteis bracteolisque glanduloso- 3(2) Leaves (at least uppermost) sessile; sepals with pale laminar landular cihatis et aunculatis glandulosis instructis, inflorescentia lines or, if lines black, then margin glandular-ciliate; ovary ovoid to 1-12-floribus, sepalis petalisque 5(4), petalis margine integris vel subglobose ( 1 . H. monanthemum) 4 cilio glandulifero apicali unico instructis post anthesin deflexis vel patulis corrugatis, gynoecio (2)3(4)-meris, differt. LeaVCS *" sh ?, ly pet ' late; sepals a11 with black linear laminar S lands < entire; ovary ellipsoid 2. H. subcordatum Hypericum sect. Adenosepalum sensu N. Robson in Bull. Br. Mus. nat. Hist. (Bot.) 5: 335 (1977) pro parte quoad H. fllicaule. 4(3) * epals a " d P ? als 5; * epals , subequal ; f '-"d^Hate to subentire; flowers 12-25 mm in diam.; leaves 10-35 mm long Perennial herbs up to 0.4 m tall, with stems erect to prostrate, la. H. monanthemum subsp. monanthemum creeping and branching at base, glabrous, with dark (black or rarely Sepals and petals 4; sepals in unequa , pairs often fo , iose emire; red) glands on leaves, sepals, petals and anthers; branching lateral, floors 6-12 mm in diam.; leaves mostly 5-10 mm long from various nodes. Stems (mature) terete or (when slender) 2^(6)- lb. H. monanthemum subsp. filicaule lined, eglandular or (4. H. himalaicum but very rarely) with few reddish glands on lines. Leaves opposite, decussate, sessile or to 3 5(2) St y' es4 -5 mm; capsule broadly ovoid; all leaves sessile lamina oblong , .. , t f . ' . . . to triangular-ovate 3. H. tngonum mm pseudopetiolate, free, persistent; lamina entire or occasionally dark-glandular-ciliate, without or occasionally with gland-fringed Styles 2-3 mm; capsule ellipsoid; some or all leaves petiolate, lamina auricles, with venation pinnate, closed and tertiary reticulation triangular-ovate to elliptic or oblanceolate to obovate 6 dense or very rarely rather dense to lax; laminar glands pale and/or 6(5) Stem (when mature) nearly always with long spreading branches from black, punctiform to striiform, dense to almost absent, relatively up to 5 nodes; styles equalling or shorter than ovary; sepals usually with small; ventral resin glands absent. Inflorescence l-15(-c. 50)- black laminar glands 4. H. himalaicum flowered, with branching dichasial to monochasial from 1-2 nodes, . . . Stem unbrancned below apical node or with a few short suberect sometimes with subsidiary branches from up to 4 nodes below; branches; styles usually longer than ovary; sepals usually without black bracts and bracteoles gland-fringed and glandular-auriculate or laminar glands 5. H. ludlowii entire and then usually foliar. Flowers stellate, homostylous. Sepals 5(4), free, persistent, erect in fruit, with margin glandular-ciliate to 7(1) Ova .7 and capsule e ' lipsoid to cylindric-ellipsoid; leaves all entire. entire; veins 5-7(9), laminar glands pale and/or dark, linear to ^ Sepals lance ' ate t0 narr wly oblo "g- elli P' ic < glandular-den- ' ticulate 6. H. dahense punctiform; marginal or submargmal glands black or reddish. Petals 5(4), persistent, deflexed to spreading, crumpling after flowering, Ovary and capsule subglobose to globose; leaves (at least upper) often with or without apiculus, margin entire or with apical glandular glandular-ciliate, lower often pseudopetiolate; sepals broadly oblong or cilium; laminar glands pale and/or dark, linear to punctiform, and * broadl y elli P tic ' glandular-ciliate to glandular-laciniate ... usually 1^1 sessile or subapical (immersed) dark glands. Stamen fascicles 5, united 2+2+1 (i.e. '3') or rarely 2+1+1+1 (i.e. '4'), 1. Hypericum monanthemum Hook. f. & Thorns, ex Dyer in persistent, with stamens totalling 10-45, filaments basally united; Hook, f., Fl. Brit. India 1: 256 (1874); Franchet in Bull. Soc. Bot. anther gland dark (black or rarely reddish); pollen type X. Ovary France 33: 438 (1886); H. Leveille in Bull. Soc. Bot. France 54: with (4)3(2) completely axile (or rarely incompletely axile to pan- 594 (1904); N. Robson in Hara & Williams, Enum. Fl. Pis Nepal etal) placentae, each -ovulate; styles (4)3(2), free, divergent from 2: 62 (1979); Li Xiwen in Fl. Xizangica 3: 276, excl. t. 1 14 ff. 3- discrete bases; stigmas narrow. Ca/ww/e (4)3(2)-valved, chartaceous 6 (1986), in Fl. R. P. Sinicae 50 (2): 54, t. 10 ff. 8-13 (1990); to papyraceous, with valves longitudinally vittate. Seeds cylindric, Biswas in Sharma & Sanjappa, Fl. India, 3: 69 (1993); Mukherjee not carinate or appendiculate; testa scalariform-reticulate to & Chaudhri in 7. Econ. Taxon. Bot. 20: 126 (1996). Type: Sikkim, scalariform or foveolate. Lachen, 3300-3600 m, 12 July 1849 (fl & fr), J.D. Hooker s.n. 76 N.K.B. ROBSON m Fig. 14 A. H. monanthemum subsp. monanthemum: (a) habit; (b) sepal; (c) petal; (d) capsule. B. H. monanthemum subsp.filicaule: (e) habit; (f) sepal; (g) petal; (h) capsule; (i) small form, see note. C. H. subcordatum: (j) habit; (k) sepal; (1) petal; (m) capsule (young) (a, e, i, j x 2/3; rest x 6). A. (a) Ludlow, Sherriff& Hicks 20395, (b, c) Forrest 6136, (d) Kingdon Ward 9913A. B. (e-h) Polunin 1121, (i) Kingdon Ward 9882. STUDIES IN HYPERICUM 77 Map 11 Sect. 9e: la. H. monanthemum subsp. monanthemum ; Ib. H. monanthemum subsp. filicaule O specimens, other records A; 2. H. subcordatum . (K!-lectotype, selected here; GH!); Latong, 3450 m, 13 July 1849 (fl & fr), J.D. Hooker s.n. (K!-syntype). Fig. 14A&B, Map 11. Perennial herb (0.05)0.1-0.4 m tall or long, erect or decumbent to prostrate from creeping, rooting and branching base, with stems scattered or clustered or often carpeting, often slender, unbranched above or rarely with one pair of ascending branches below inflores- cence. Stems terete or 2(4)-lined, eglandular; internodes 5-50 mm, longer than leaves. Leaves sessile (uppermost or very rarely all) or to 1 mm petiolate, lower smaller, erect to appressed, soon deciduous; lamina 4-25(-35) x 2-15(-25) mm, broadly ovate or circular to broadly oblong or broadly elliptic or obovate-spathulate, paler or slightly glaucous beneath, plane, subchartaceous to submembraneous; apex rounded or retuse to obtuse, margin entire, plane, base rounded to truncate or subcordate or (lower) cuneate; venation 3(4) pairs of main laterals from lower third of midrib, with tertiary reticulation dense; laminar glands punctiform, all pale and very small or some or all black; intramarginal glands black and dense or reddish to pale and rather sparse. Inflorescence l-5(-7)- flowered, from 1(2) nodes, subumbelliform or bifurcate; pedicels 0.5-2.5 mm (-5 mm in central flower); bracts foliaceous, the pair usually wider than inflorescence, entire; bracteoles narrowly ovate to lanceolate, black-glandular-ciliate and -auriculate, persistent. Flowers (6-) 10-25 mm in diam., stellate to reflexed; buds narrowly ovoid, obtuse. Sepals 5, subequal or 4 in unequal pairs, erect in bud and fruit, 2.5-7 x 1-3 mm, oblong or elliptic to narrowly ovate or linear-lanceolate, obtuse (rarely acute) to rounded, glandular-ciliate to entire; veins 5-7, branching and often reticulating towards mar- gin; laminar glands all pale or parts or all black, linear; marginal glands black, on cilia or sessile, or reddish to pale, sessile or intramarginal. Petals 5-4, golden yellow, not tinged red in bud, (3) 5-15 x 1^.5 mm, 1-2 x sepals, lanceolate-oblong to narrowly ovate, obtuse to acute, margin entire, laminar glands few, pale or black, linear to striiform, or absent, apiculus gland reddish, other marginal glands absent. Stamens 10-45, '3' -fascicled, longest c. 4- 9 mm, 0.6-0.8 x petals; anther gland black. Ovary (4?)3(2?)-locular 2-5 x 1.5-2.5 mm, ovoid to subglobose; styles (4)3(2), 1.5-3.5(-4) mm, 0.5-1 x ovary; stigmas narrow. Capsule (5-)6-8 x (3-)4-6 mm, broadly ovoid-ellipsoid to ellipsoid; valves with numerous longitudinal vittae. Seeds yellowish brown, 0.7-0.8 mm, cylindric, acute at both ends; testa shallowly foveolate. 2n = ? Clearings in forests and bamboo forests, thickets, grassy or stony slopes and streamsides, rock crevices; 2270-4400(^4800) m. Himalayan Range from west Sichuan and Yunnan to Nepal. Joseph Hooker collected two similar plants in Sikkim, one with a 5- merous perianth and the other with a 4-merous perianth. Thistleton Dyer ( 1 874) placed the former in Hypericum (H. monanthemum) and the latter along with other 4-merous Hypericum relatives in the Linnaean genus Ascyrum (A. filicaule Dyer), even though all previ- ously described Ascyrum species had been shrubs or herbaceous derivatives from eastern North America and the Caribbean. In addition, A. filicaule has black anther glands, whereas all the Ameri- can plants are completely destitute of black glands. The Himalayan plant would therefore seem to have developed 4-mery independently. A detailed study has revealed that not only is A. filicaule a Hypericum, but also that it is only with difficulty separable from H. monanthemum. This latter 'species' displays a western reduction trend, from China to Nepal, while the reduction trend in A. filicaule goes in the reverse direction, from Nepal and Sikkim to Yunnan and Myanmar. In Nepal, Sikkim and Bhutan there is an area of morpho- logical overlap, where some specimens have perianth states intermediate between 5-mery and 4-mery. It therefore seems appro- priate to treat these taxa as subspecies. la. Hypericum monanthemum subsp. monanthemum Fig. 14A, Map 11. H. monanthemum var. nigropunctatum Franchet, PI. delavay.: 103 (1889) ['nigro-punctatum']. Type: China, Yunnan, in locis umbrosis montis Tsang-chan, supra Tali, 4000 m, 3 August 1 884 (fl), Delavay 1944 (P!-holotype; K!-isotype). H. yunnanense Franchet in Bull. Soc. Bot. France. 54: 437 (1886) pro parte quoad specc. Delavay 93 et 1941. 78 N.K.B. ROBSON H. mairei H. Leveille in Bull. Acad. Int. Geogr. Bot. 25: 23 (1915) non H. Leveille (1912). Type: China, Yunnan, paturages du plateau de Ta-Hai, 3200 m, July 1912, Maire s.n. (E-holotype). H. bachii H. Leveille, Cat. PI. Yun-nan: 131(1916). Type as for H. mairei H. Leveille (1915) non H. Leveille (1912). Icon: Li Xiwen in Fl. R. P. Sinicae 50(2): 53, t. 10 ff. 8-13 (1990). Plant erect. Stems terete. Leaves sessile or subsessile, 10-25(-35) x 8-15(-25) mm, never scale-like, subchartaceous, sometimes slightly glaucous beneath, base rounded to broadly cuneate, intramarginal glands black. Inflorescence l-5(-7)-flowered, pedicels 1-2.5 mm. Flowers 12-25 mm in diam. Sepals 5, subequal, 4-7 x 1.5-3 mm, margin glandular-ciliate to subentire. Petals 5, 8-15 x 2.5^.5 mm, c. 2 x sepals. Stamens 20-45. Styles c. 0.6-1 x ovary. Clearings in forests and bamboo forests, grassy slopes, streamsides; 2270-c. 4000 m. China (west Sichuan, Yunnan, south-east Xizang), Myanmar (Kachin), Bhutan, India (Benghal, Sikkim), Nepal. CHINA. Sichuan: Emei Shan, 3300 m, 25 July 1931 (fl & fr), Tang & Wang 23435 (GH, KUN, NAS); Ebian Xian, 2600 m, 25 July 1976 (fl), Sichuan PL Exped. 13188 (CDBI). Yunnan: Dali Xian, Diancang Shan, 2900-3200 m, 19 July 1984 (fl & e. fr), Sino-Amer. Bot. Exped. 1 166 (A, BM, CAS, KUN, US); Lichiang Range, E. flank (27 300 N), 3300 m, July 1910 (fl), Forrest 6136 (BM, E*, K); Chih-tze-lo Hsien. Pi-lo Shan, 4000 m, 27 August 1934 (fr), Tsai H.T. 58243 (A, IBSC, KUN, PE). Xizang: Zayu Xian, 3600 m, 13 August 1973 (fl), Qingzang Exped. 73-1083 (KUN, PE); Kongbo Prov., Tumbatse, Rong Chu, 3480 m, 2 July 1938 (fl), Ludlow, Sherriff & Taylor 5093 (BM); Yatung, 27 510 N, 88 350 E, 1897 (fl & fr), Hobson s.n. (K). MYANMAR. Kachin: Nam Tamai valley ( Adung Wang-Gamlang Wang), 3000-3600 m, 1 October 1937 (fl), Kingdon Ward 13344 (BM); Adung valley, sources of the Irrawaddy, 3600 m, 8 August 1931 (fl & fr), Kingdon Ward 9163 (BM). BHUTAN. North: Me La, 3900 m, 23 June 1949 (fl), Ludlow, Sherriff & Hicks 20395 (BM, E*); Upper Pho Chu (east branch), Leji, 3600 m, 28 June 1949 (fl), Ludlow, Sherriff & Hicks 16671 (BM, E*). INDIA. Benghal: *Tonglu, 3000 m, Smith & Cave 2537 (Lloyd Bot. Gdn, Darjeeling). Sikkim: Theumthang [Thaunthang], 4050 m, 2 August 1913 (fl), Cooper 408 (BM, E*); Lachen, 3300-3600 m, 12 July 1849 (fl & fr), Hooker s.n. (GH, K). NEPAL. East: Topke Gola, 3450 m, 4 July 1971 (fl), Beer 8271 (BM). The Cooper (Sikkim) and Beer (Nepal) specimens are intermediate between subspp. monanthemum andfilicaule. Ib. Hypericum monanthemum subsp. filicaule (Dyer) N. Robson, comb, et stat. nov. Type as for Ascyrum filicaule Dyer. Fig. 14B, Map 11. Ascyrum filicaule Dyer in Hook, f., Fl. Brit. India 1: 252 (1874); Li Xiwen in Acta Bot. Yunnan. 3: 329, ff. 1.4, 1.5 (1981). Type: India, Sikkim, Lachoong Valley, 3600 m, 3 September 1849 (fl), J. D. Hooker s.n. (K!-holotype). Hypericum filicaule (Dyer) N. Robson in Bull. Br. Mus. nat. Hist. (Bot). 5: 305 (1977); Li Xiwen in Fl. R. P. Sinicae 50(2): 52, t. 10 ff. 1-7 (1990); Biswas in Sharma & Sanjappa, FL India, 3: 58 (1993). Icon: Li Xiwen in FL R. P. Sinicae 50(2): 53, t. 10 ff. 1-7 (1990). Plant erect to decumbent or prostrate and mat-forming, with stems unbranched above or (in Yunnan) branched from upper 2-3 nodes. Stems terete or narrowly 2-lined. Leaves to 1 mm petiolate, (4 )5- 10(-15) x 2-8(-H) mm, broadly elliptic, lower gradually smaller, sometimes becoming minute and scale-like, thinly chartaceous to membranous, paler beneath, base cuneate to rounded-attenuate, intramarginal glands reddish or pale. Inflorescence 1 -flowered, pedicels (terminal) 0.5-1.5 mm. Flowers 6-12 mm in diam. Sepals 4, sometimes foliar, outer 4-12 x 1.5-7 mm, inner 3-7 x 1-3 mm, obtuse, margin entire. Petals 4, 3-8 x 1-2.7 mm, 0.75 -1 .6 x sepals. Stamens 10-20. Styles 0.5-0.8 x ovary. In rock crevices and on grassy slopes; 3000-4800 m. China (north-west Yunnan, south-east Xizang), India (Arunachal Pradesh, Sikkim), Nepal and possibly Myanmar (see note below). CHINA. Yunnan: Gongshan Xian, Longpanla, 3500 m, October 1935 (st), Wang C.W. 67066 (KUN); loc. not stated, 1917-1919 (st), Forrest 14279 (BM, E*). Xizang: Bomi Xian, 4000 m, 15 August 1965 (fl), Ying & Hong 651 164 (PE); ? Xian, Chickchor, 3600 m, 5 July 1935 (fl), Kingdon Ward 1 1899 (BM); Kongbo Prov., Lusha Chu, 3600 m, 8 June 1938 (fl), Ludlow, Sherriff & Toy/or 4713 (BM). INDIA. Arunachal Pradesh: Senge Dzong, 3600-3900 m, 30 May 1935 (fl), Kingdon Ward 1 1561 (BM); Bhutan frontier, Orka La, 3900 m, 12 June 1 938 (fl), Kingdon Ward 1 3728 (BM). Sikkim: Lachen, 3600 m, 20 June 1 849 (fl), Hookers.n. (K); Lachoong, 3600m, 3 September 1 849 (fl), Hookers.n. (K). NEPAL. Central: Rambrong, Lamjung Himal, 4050 m, 7 July 1954, Stainton, Sykes & Williams 62 1 6 (BM); Khola Kharka, c. 4050 m, 1 7- 1 9 July 1949(fl), Polunin 1121 (BM). Subsp. filicaule almost certainly occurs in Bhutan. NOTE. An extremely reduced form of subsp. filicaule was col- lected by Kingdon Ward in the same region of north Myanmar as that in which he found plants of subsp. monanthemum. I am reluctant to describe it as a distinct taxon in the absence of other similar material from Myanmar or adjacent Yunnan or Xizang, as it differs essentially from some Chinese populations (e.g. Feng 6550) only in size of parts and occurs in a region very close to the Xizang part of the range of subsp. filicaule: Herb 'forming mossy clumps on granite boulders', with stems prostrate to ascending, branching distally. Stems filamentous, eglandular, narrowly 4-lined, becoming 2-sided; internodes 1 .5-3 mm, upper exceeding leaves, lower shorter than them. Leaves all with petioles 0.2-0.3 mm; lamina broadly ovate to orbicular, upper 2^1- x 1.5-3 mm, decreasing in size towards base, subappressed, outcurving, paler beneath; apex rounded, margin entire, base cuneate to angustate; veins: 3 pairs of laterals from lower third, tertiary reticulation lax; laminar glands pale rather sparse; marginal glands very small, pale, spaced. Inflorescence 1 -flowered. Flowers c. 8 mm in diam., stellate; buds narrowly ellipsoid. Sepals 4, unequal, 2 x 1 .2 (outer) or 0.6 (inner) mm, elliptic to oblanceolate, entire, rounded; laminar glands pale, punctiform to striiform; marginal glands absent. Petals 4, 'yellow', 5-5. 5 x 1.5-2 mm, c. 2.5 x sepals, oblong, almost entire; laminar glands absent, marginal gland solitary, terminal, on short cilium, reddish. Stamens 12, '3'-fascicled with 4 in each fascicle, longest 2.5-3 mm, c. 0.5 x petals; anther gland black. Ovary 1-locular, c. 1.5 x 1 mm, narrowly ovoid; styles 3, 0.9-1.1 mm, c. 0.65 x ovary; stigmas purple, narrowly capitate. Capsule 2.5-3.5 x 2-2.5 mm, ellipsoid; valves finely vittate. Seeds yellow- brown, c. 0.5 mm, cylindric; testa shallowly finely scalariform. MYANMAR. Kachin: Adung Valley, sources of the Irrawaddy, 3900-4200 m, 31 March 1931 (fl & fr), Kingdon Mm* 9882 (BM). 2. Hypericum subcordatum (R. Keller) N. Robson, comb, et stat. nov. Type as for H. thomsonii var. subcordatum R. Keller. Fig. 14C, Map 11. H. thomsonii var. subcordatum R. Keller in Bot. Jb. 33: 553 (1904). Type: China, Shaanxi, Huan-tou-san, July 1899 (fl & fr), Giraldi 540 (Fl-holotype). H. thomsonii R. Keller in Bot. Jb. 33: 552 (1904) pro parte excl. typum. STUDIES IN HYPERICUM 79 H. mororanense R. Keller in Bot. Jb. 33: 552 (1904) pro parte quoad spec. Sin. cit. H. petiolulatum var. subcordatum (R. Keller) H. Leveille in Bull. Sac. Bot. France 54: 594 (1908). Perennial herb 0. 1 -0. 1 7 m tall, erect from creeping and rooting base with stems unbranched or branched from upper 1 (2) nodes. Stems slender, eglandular; internodes 2-lined above, terete below, 30-40 mm, longer than leaves. Leaves all petiolate, with petiole 1-1 .5 mm; lamina 10-16 x 8-12 mm, broadly elliptic or oblong-elliptic to obovate, paler (or subglaucous?) beneath; apex rounded, margin entire, base cuneate to truncate (or uppermost subcordate); veins: 3 pairs of laterals from lower third or lower, tertiary reticulation rather dense, obscure; laminar glands dense, small, pale or sometimes a few black, scattered; marginal glands dense or subirregular, black. Inflorescence (l-)3-5-flowered, occasionally with branches from 1(2) nodes below, the whole corymbiform or bifurcating to subcylindric; pedicels 2-2.5 mm; bracts and bracteoles foliar, entire. Flowers c. (8-)10-12 mm in diam., stellate; buds ellipsoid? Sepals 5, subequal, 3-^.5 x 0.8-1.2 mm, oblong-lanceolate to lanceolate, acute, margin entire; veins 5, unbranched; laminar glands linear to (distally) punctiform, black; marginal glands few, sessile or submar- ginal, all or most distal, black. Petals 5, golden? yellow, 4.5-6 x 1-1.5 mm, 1.35-1.5 x sepals, narrowly elliptic, entire, laminar glands few, black, linear to striiform, marginal glands few, black, subapical or absent. Stamens 17-20, '3'-fascicled, longest 4-5.5 mm, c. 0.9 x petals; anther gland black. Ovary 3(4)-locular c. 2 x 1 mm, ellipsoid; styles 3(4), c. 1.7-2 mm, c. 0.85-1 x ovary, narrowly or scarcely capitate. Capsule 4.8-6 x 3.5^1- mm, ovoid to subglobose; valves densely longitudinally vittate. Seeds yellow-brown, 0.6 mm, cylindric; testa shallowly finely scalariform. 2n = ? No habitat details known; 1800-2850 m. China (Shaanxi, Sichuan). CHINA. Shaanxi (south): Cho-toc Miao, Hu Shien, n.d. (fl), Fr Hugh [Scallan] s.n. (BM); MtThae-pei-san, August 1899 (fr), FrHugh s.n. (BM); Taibai Shan, Lotosze, 1800 m, 21 September 1932 (fr), Hao K. S. 4303 (PE); Taibai [Thae-pei] Shan, Pingan Si to Fangyang Si, 11 September 1937 (fr), Liou & Tsoong 8 1 7 (PE); Taibai Shan, Fangyang Si to Mingqing Si, 2850 m, 1 1 August 1 977 (fl & fr), Guo & Hu 525 (IBSC); Huan-tou-san, July 1 899 (fl & fr), FrHugh s.n. (BM); Mt Miao-Wang-san, 1 898 (fr), FrHugh s.n. (BM); Wu Shan, summit, 1 July 1903 (fl), Wilson 3259 (K). Sichuan: Emei Shan, 2300 m, 24 June 1 940 (fl), Ma W. W. 2606 (KUN, US); Ebian Shan, Shouping Shan, 25 August 1939 (fr), Shang (Sun?) S.S. 1063 (A, KUN, US); Hongya Xian, Wawu Shan, 1 July 1939 (fl), Yao C. W. 3978 (PE); Nanchuan Xian, Jin Shan, 1980 m, 1 July 1957 (fr), Tsuing & Zhou 92391 (IBSC); Leibo Xian, 2400 m, 23 June 1959 (fl), Kuan C.T. 7635 (IBSC); Wuxi Xian, Shuanggou Xiang, 2600 m, 23 July 1958 (fl & fr), Yang G.H. 58939 (PE). H. subcordatum is very closely related to H. monanthemum subsp. monanthemum and continues the northward trend of that taxon. Indeed, it could be regarded as a small delicate 'form' of H. monanthemum var. nigropunctatum (i.e the form of the type subspe- cies with black-gland-lined sepals) except that the leaves are all petiolate and the sepals acute and entire, characters that do not occur in combination in H. monanthemum itself. These differences, to- gether with the occurrence of both taxa on Emei Shan, where they remain distinct, would appear to justify the separation of H. subcordatum as a species. 3. Hypericum trigonum Hand.-Mazz., Symb. Sin. 7: 403, t. 8 f. 6 (1931). Type: China, Yunnan (NW), 'am Passe Akelo zwischen Djinscha-djiang ("Yangtse") und Landsang-djiang (Mekong), 27 19', am Wege von Djitsung nach Kakatang', 2900-3100 m, 30 August 1915 (fl & fr), Handel-Mazzetti 7920 (W!-holotype; US!-isotype). Fig. 15 A, Map 12. H. monanthemum sensu Li Xiwen in Fl. R. P. Sinicae 50(2): 276 (1990) pro parte excl. typum. Icon: Hand.-Mazz., Symb. Sin. 7: 403, t. 8 f. 6 (1931). Perennial herb 0.25-0.4 m tall, erect from short creeping and rooting base, with stems unbranched below inflorescence or with slender ascending or spreading branches from up to c. 5 upper nodes. Stems terete, eglandular; internodes 20-43 mm, longer than or equalling leaves. Leaves sessile, lower not smaller but ascending Map 12 Sect. 9e: 3. H. trigonum A; 5. H. ludlowii ; 6. H. daliense O. 80 N.K.B. ROBSON Fig. 15 A. H. trigonum: (a) habit; (b) sepal; (c) petal; (d) capsule (young). B. H. himalaicum: (e) habit; (f) sepal; (g) petal; (h) capsule. C. H. ludlowii: (i) habit; (j) habit, small form; (k) sepal; (1) petal; (m) capsule (a, e, i, j x 2/3; rest x 6). A. Kingdon Ward 9913. B. Stainton, Sykes & Williams 1666. C. (i, k, 1 ) Ludlow, Sheriff & Taylor 5461, (j) Ludlow, Sheriff & Hicks 19502, (m) Ludlow & Sherriff 14214. STUDIES IN HYPERICUM 81 and soon deciduous; lamina 20-30 x 9-15 mm, oblong or oblong- ovate to triangular-ovate, paler beneath, not glaucous, plane, subchartaceous; apex rounded to obtuse, margin entire, base broadly cuneate to shallowly cordate; venation: 4 pairs of main laterals from lower half of midrib, with tertiary reticulation dense; laminar glands punctiform, all pale and very small or all black; intramarginal glands black, dense. Inflorescence 4-c. 15-flowered from terminal node, with flowering branches from up to 4 nodes below, the whole corymbiform to cylindric; pedicels l-2.5(-5, central flower) mm; bracts narrowly lanceolate to linear, black-glandular-ciliate and -auriculate (cilia long), persistent. Flowers 15-c. 25 mm in diam., stellate or reflexed; buds cylindric, rounded. Sepals 5, equal, erect in bud and fruit, 6-7 x 1 .5-2 mm, narrowly oblong to elliptic-oblong or lanceolate, acute, glandular-ciliate; veins 5, branching and some- times reticulating towards margin; laminar glands all pale or parts black, linear or interrupted; marginal glands black, on cilia. Petals 5, golden yellow, not tinged red in bud, 9-1 1 x 2.5-4.5 mm, c. 1 .5(-2?) x sepals, narrowly oblong-elliptic to lanceolate-elliptic, acute (i.e. with prominent apiculus), margin entire, laminar glands few, pale, linear and sometimes black, 1-3, subapical; marginal gland solitary, in apiculus, black. Stamens c. 25-35, '3 '-fascicled, longest 7-9 mm, c. 0.75-0.8 x petals; anther gland black. Ovary 3-locular?, c. 4-5 x 1 .8-2 mm, narrowly ovoid-ellipsoid; styles 3, 4-5 mm, about equal- ling ovary; stigmas narrowly capitate. Capsule 7-8 x c. 4.5 mm, broadly ellipsoid, c. 1-1.5 x sepals; valves with numerous longitu- dinal vittae. Seeds not seen. 2n = ? Marshes and wet meadows?; 2600-3650 m. China (north-west Yunnan). CHINA. Yunnan: Jugo-shan Pass, 3600 m, 30 June 1922 (fl), J.W. & C.J. Gregory s.n. (BM); Litiping between Likiang and Weihsi, 8 October 1939 (fr), Ching R.C. 22069 (A, KUN); Weixi Xian, Yezhi, 3650 m, 6 October 1934 (fr), Tsai H.T. 59699 (A, IBSC, KUN, NAS, PE); Gongshan Xian, 9 October 1940 (fr), Feng K.M. 8308 (KUN); Mekong-Salwin Divide, Sila, 3400 m, 25 September 1928 (fl & fr), YuT.T. 22728 (A). The H. trigonum group (Spp. 3-5), with the closely allied H. monanthemum group (Spp. 1-2), would appear to be related to H. przewalskii and to be derived from it along with, but independently of, the H. seniawinii group (sect. Elodeoidd) (see p. 46). Although members of both groups have penetrated westward along the Himalayan Range, the H. seniawinii group is basically south-eastern and upland and the montane species are derivative. The H. trigonum group, on the other hand, is basically south-western and montane. Both groups contain variable taxa that are sometimes difficult to separate specifically, but H. trigonum and its relatives are particu- larly complex in this regard. The group comprises H. trigonum (north-west Yunnan), H. himalaicum (Pakistan to north-west Yunnan and adjacent Sichuan) and H. ludlowii (Yunnan, south-east Xizang and Bhutan). H. trigonum is basic to the group, being most similar to the long-styled form of//, monanthemum, and the two other taxa are apparently derived from it. The problem is to decide whether there are three species or one species with three subspecies or something in between. The decision is complicated by a population in extreme northern Myanmar and adjacent Arunachal Pradesh that is interme- diate in some respects between H. trigonum and H. himalaicum. Short of making four subspecies, which would not, I think, be a satisfactory reflection of the situation, it seems best to recognize three species and place the intermediate population in H. himalaicum, despite the long distance between its range (Arunachal Pradesh- Myanmar frontier) and that of the most primitive form of H. himalaicum proper (central Nepal). 4. Hypericum himalaicum N. Robson in J. Jap. Bot. 52: 287, ff . 3- 4 (1977), in Hara & Williams, Enum. Fl. Pis Nepal 2: 61 (1979); Robson & Long in Grierson & Long, Fl. Bhutan 1(2): 377, f. 30e-g (1984); Li Xiwen in Fl. Xizangica 3: 278 (1986), in Fl. R. P. Sinicae 50(2): 56 (1990) pro parte uterque quoad typum; Biswas in Sharma & Sanjappa, Fl. India, 3: 64 (1993); Mukherjee 6 Chaudhuri in J. Econ. Taxon. Bot. 20: 125 (1996). Type: Nepal, Gossain Than, n.d. (fl), Wallich 4814 (K-W!-holotype; BM!, K!-isotypes). Fig. 15B, Map 13. H. pollens D. Don, Prodr.fl. nepal. : 2 1 9 ( 1 825), nom. illegit. superfl. pro parte excl. typum, non H. pollens Banks & Solander (1794). H. setosum Wall. [Num. List: no. 4814 (1831), nomen] ex Dyer in Hook, f., FL Brit. India 1: 256 (1874) in syn., non H. setosum L. (1753). H. napaulense sensu Dyer in Hook, f., Fl. Brit. India 1: 256 (1874) pro parte excl. typum; Y. Kimura in Kihara, Fauna & Fl. Nepal Himal: 278 (1955); Anon, in Bull. Dept. Med. PI., Thapathali 1: 7 ( 1 967); N. Robson in Nasir & Ali, Fl. W. Pakistan 32:11, f. 2A- F (1973) pro parte excl. syn. H. monanthemum; pro parte omnes excl. typum. H. monanthemum var. brachypetalum Franchet, PI. delavay.: 104 (1889). Type: China, Yunnan, in pratis ad collum Yen-tze-hay, 3200 m, 17 June 1887 (fl), Delavay 5180 (Pl-lectotype, selected here; K!-isolectotype); Yunnan, in collibus prope Pien-kio, 11 November 1887 (fl), Delavay s.n. (P'.-syntype). H. humifusum sensu Y. Kimura in Hara, Fl. E. Himal: 209 (1966). Icones: N. Robson in Fl. W. Pakistan 32: 11, f. 2A-F (1973), in Fl. Bhutan 1(2): 377, f. 30 e-g (1984). Perennial herb 0.05-0.35 m tall or long, suberect to decumbent from creeping and rooting base, with stems solitary? or clustered, often branched below inflorescence or lower, with branches slender, spread- ing. Stems terete or 2-4(6)-lined, eglandular or occasionally with few reddish glands on lines; internodes 8-50 mm, longer than leaves. Leaves sessile or to 2 mm petiolate; lamina 4-24 x 2- 1 7 mm, ovoid to oblong or elliptic to obovate or oblanceolate, paler or usually glaucous beneath, plane, subchartaceous; apex rounded, margin entire, base cordate to rounded or cuneate, the upper often with black-glandular- ciliate auricles and sometimes also glandular-ciliate proximal margin; venation: 2-4 pairs of main laterals from lower quarter to almost half of midrib, with tertiary reticulation dense; laminar glands pale, punctiform, small to medium, dense; intramarginal glands black, dense. Inflorescence 1-12-flowered, from 1(2) nodes, often with flowering branches from up to 4 nodes below, the whole subcorymbiform; pedicels 1 .5^ mm; bracts and bracteoles narrowly elliptic or linear-lanceolate to linear, black-glandular-ciliate and - auriculate or more rarely entire and then either bracteose or reduced-foliar, persistent. Flowers 10-20 mm in diam., stellate; buds ovoid to broadly ellipsoid, obtuse to acute. Sepals 5, 3.5-7 x 1-2.5 mm, equal to subequal, erect in bud and fruit, ovate-lanceolate or elliptic to linear-lanceolate, subacute to acute or rarely rounded, glandular-ciliate to entire; laminar glands pale or often partly black, linear (with black parts dots to usually streaks) to dots; marginal glands black, on cilia or sessile or absent. Petals 5, bright yellow, sometimes tinged red in bud, 6-10 x 2-4 mm, 1.2-2 x sepals, oblong-elliptic to oblanceolate, rounded, margin entire or rarely with one apical glandu- lar cilium, laminar glands black, striiform, or absent, marginal glands absent or 1-4, black, apical and subapical, sessile or apical on cilium. Stamens 12-30, '3' -fascicled, longest 4-7 mm, 0.65-0.85 x petals; anther gland black. Ovary 3-locular, 2-3.5 x 1.5-2 mm, ovoid to ellipsoid; styles 3(4), 2-3 mm, 0.65-1 (-1.3?) x ovary; stigmas narrowly capitate to narrow. Capsule 3-9 x 2.5-6 mm, broadly ellipsoid, exceeding or equalling sepals. Seeds yellowish-brown, 0.5- 0.6 mm, cylindric; testa densely scalariform-foveolate. 2n = ? 82 N.K.B. ROBSON Map 13 Sect. 9e: 4. H. himalaicum specimens, other record O. Forest and woodland clearings, alpine meadows and rocky or grassy slopes, often in damp places; 2100-3900 m. Himalaya from Pakistan to east Bhutan and China (south-east Xizang, west Yunnan and adjacent? Sichuan), with an isolated population in the Arunachal Pradesh-Myanmar frontier area (see note under 3. H. trigonum). CHINA. Sichuan: no precise locality, 3150 m, 9 August 1964 (fl & fr), Sichuan Exped. 04053 (CDBI). Yunnan: E. flank of Lichiang Range, 27 300, 3300 m, July 1910 (fl), Forrest 6016 (BM, K); Zhongdian Xian, 3200 m, 28 August 1 962 (fl), Zhongdian Exped. 1 598 (KUN, PE); Yangbi Xian, W. side of Diancang Shan, vicinity of Dajiuping, 2500-2600 m, 30 June 1984 (fl), 1984 Sino-Amer. Bot. Exped. 620 ( A, BM, CAS, KUN, US). Xizang: Nage, 3200 m, 3 1 July 1 974 (fl&fr),Qingzang Exped. 74-3797 (PE);Nyingchi Xian, 3040m, 28 July 1965 (fr), Chang & Lang 1024 (PE); Yadong Xian, 3100 m, 15 September 1974 (fl & fr), Qingzang Exped. 74-2561 (KUN, PE). MYANMAR. Kachin: Sources of the Irrawaddy, Adung Valley, 3600 m, 7 August 1931 (fl), Kingdon Ward 9913 (BM). INDIA (East). Arunachal Pradesh: Assam frontier, Mishmi Hills, Delei Valley, Dri La, 3000 m, 13 December 1933 (fr), Kingdon Ward 1 1049 (BM). Benghal: Kalimpong, Labah, 2100 m, September 1904 (fr), Haines B.B. 983 (K); Darjeeling [Darjiling], Singaleleh, 3000 m, 7 October 1870 (fr), Clarke 13513 (K). Sikkim: Tonglo, 3000 m, 5 August 1874 (fl), Treutler 505 (K); Jongri, 3600 m, 15 October 1875 (fr), Clarke 25852 (K). BHUTAN. Central: Thimpu [Thimbu) distr, E. of Thimphu, c. 3000 m, 20 July 1979 (fl), Grierson & Long 2798 (BM, E*). NEPAL. East: Kasuwa Khola, 3450 m, 18 August 1975 (fl & fr), Beer 25327 (BM); Lumding Khola, 3000 m, 13 July 1954 (fl & fr), McCosh 404 (BM). Central: Kali Gandaki, Lete (S. of Tukucha), 3000 m, 9 July 1 954 (fl & fr), Stainton, Sykes & Williams 1666 (BM); above Maikot, 3300 m, 26 June 1 954 (fl), Stainton, Sykes & Williams 3262 (BM). West: near Tarakot, c. 3 1 50 m, lOJuly 1 952 (f\),~Stainton, Sykes & Williams 2415 (BM);TanglaBanyang, 3400 m, 1 7 August 1 973 (fl & fr), Einarsson, Skarby & Watterhall 3396 (BM). INDIA (West). Uttar Pradesh: Kumaon, Kalimundi, 2610 m, n.d. (fl & fr), Strachey & Winterbottom 3 (BM, GH, K); Tihri-Garhwal, Ganges Valley near Jangla, 2400-2700 m, 10 July 1883 (fl), Duthie 976 (BM). Himachal Pradesh: Lahul, Manali 3000 m, 2 August 1941 (fl & fr), Bor 15587 (K); Simla, Bushahr, Pangi, 8 August 1934 (fl & fr), Negi Parmanand 863 (K). Kashmir: Bhadawar Distr., Chithar, Kal Nai, 3000 m, 24 July 1943 (fl), Ludlow & Sherriff9228 (BM); Pahlgam, 2250 m, 15 August 1945 (fr), Stewart 21696 (K). PAKISTAN. Hazara: Thandiani, 2400 m, 14 August 1956 (fl & e.fr), Stewart 27773 (BM, RAW); Murree Hills, DungaGali, 25 August 1962 (fl & fr), Stewart 1561 (RAW). The specimens of H. himalaicum proper that most closely resemble the intermediate population from north Myanmar/ Arunachal Pradesh, and hence are nearest to 3. H. trigonum, are found in central Nepal. From there a westward reduction trend involves merely changes in size, petiole length and complexity. A corresponding eastward trend, however, has resulted in simplification as well as reduction, so that not only do the plants become much smaller in Sikkim, but they become simpler, e.g. the sepals become narrower, acute and entire. Such plants have been confused, by me (Robson, 1977) as well as by Kimura (1966), with the reduced form of//, wightianum in south India that, in turn, had been confused earlier (e.g. Dyer, 1 874; Keller, 1925; Alston, 1931) with the European H. humifusum. The un- doubted resemblances between these populations, however, are clearly due to convergence. The other 'arm' of this eastward trend penetrates into Xizang through the Yadong (= Chambo) gap and continues to north-west Yunnan and north into Sichuan, where it has been known as H. monanthemum var. brachypetalum Franchet and where the sepals are sometimes entire and rounded. 5. Hypericum ludlowii N. Robson in Notes Roy. Bot. Card. Edin- burgh 41: 133 (1983), in Grierson & Long, Fl. Bhutan 1(2): 378 ( 1 984). Type: China, Xizang, Kongbo, Tsangpo Valley, Doshong, 2850 m, 28 July 1938 (fl), Ludlow, Sherriff& Taylor 5461 (BM!- holotype). Fig. 15C, Map 12. H. monanthemum sensu Li Xiwen in Fl. Xizangica 3: 276 (1986) pro parte quoad t. 1 14 ff. 3-6. H. himalaicum sensu Li Xiwen in Fl. R. P. Sinicae 50(2): 57 (1990) pro parte quoad descr. p.p. et t. 12 ff. 1-3. Icones: Li Xiwen in Fl. Xizangica 3: 276, 1. 1 1 4 ff . 3-6 ( 1 986), in Fl. R. P. Sinicae 50(2): 58, t. 12 ff. 1-3 (1990). STUDIES IN HYPER/CUM Perennial herb up to 0.4 m tall or long, erect or ascending from creeping and rooting base, with stems slender, growing through other vegetation, unbranched or with short branches from upper or occasionally several nodes. Stems 2^-lined, eglandular; internodes 7-30 mm, exceeding leaves. Leaves petiolate, with petiole 0.5-1 mm; lamina 3-20 x 1.5-10 mm, triangular-ovate or elliptic to oblong-oblanceolate, paler beneath, not glaucous, subchartaceous; apex rounded, margin entire, plane, base rounded to rarely cuneate; venation: 3(2) pairs of main laterals from lower third to quarter of midrib, with tertiary reticulation dense; laminar glands pale, puncti- form, very small, dense to sparse; intramarginal glands all black or black and pale or reddish, rather dense. Inflorescence 1-9-flowered from 1-2 nodes, subcorymbose or broadly pyramidal to cylindric or bifurcate, sometimes with flowering branches from up to 3 nodes below; pedicels 1.5-3 mm; bracts and bracteoles narrowly oblong and black- or red-glandular-ciliate and -auriculate or reduced-foliar and entire without auricles, persistent. Flowers (6-)9-ll(-15) mm in diam., stellate; buds cylindric-ellipsoid, subacute. Sepals 5, subequal or unequal, 1.5-5 x 0.5-1.5 mm, erect in bud and fruit, narrowly oblong or narrowly elliptic to lanceolate, obtuse to acute, glandular-ciliate to entire; veins 5, unbranched; laminar glands linear, all pale or partly black; marginal glands black or reddish, on cilia or sessile or submarginal, sometimes few. Petals 5, golden yellow, not tinged red in bud, 4-7 x 1-2.5 mm, (1.3-)1.5 x sepals, oblong-oblanceolate, rounded, margin with black or reddish gland in apiculus and sometimes a few subapical sessile black glands, without or rarely with 1-2 laminar black streaks. Stamens c. 20, '3'- fascicled, longest 4-5 mm, c. 0.6-1 x petals; anther gland black. Ovary 3?-locular, 1.6-2.5 x 1-2 mm, ellipsoid to subglobose; styles 3, 2-2.5 mm, 1-1.4 x ovary; stigmas narrowly to scarcely capitate. Capsule 4.5-6 x 2.5-3.5 mm, cylindric-ellipsoid to ellipsoid- subglobose; valves longitudinally vittate. Seeds straw-coloured, 0.4 mm, cylindric; testa foveolate-scalariform. 2n = ? Grassy swamps, streambanks and bogs; 2850-3400(-3600) m. China (Yunnan, Xizang), Bhutan. CHINA. Yunnan: WeixiXian,PantianGe, lONovember 1961 (fr),Kunming Working Stn. s.n. (KUN). Xizang: Tsarung, N. slope of Mt Kenichunpo, May-June, 1932 (fl & fr), Rock 22157 (BM, G, K, NY); Nyingchi Xian, Lunang, 3400 m, 4 August 1975 (fl), Qingzang Exped. 751280 (KUN, PE); Kongbo, Nayu Chu, 3 1 50 m, 7 July 1938 (fl), Ludlow, Sherriff& Taylor 5761 (BM); Mainling Xian, Hongwei Forest Farm, 3200 m, 29 July 1975 (fl), Qingzang Exped. 750980 (KUN, PE); Kongbo, Cha, Gyamda Chu, left bank, 2850 m, 31 July 1947 (fl & fr), Ludlow & Sherriff \42\4 (BM). BHUTAN. Central: Bhumthang distr., Dhur, 3000 m, 23 July 1949 (fl & fr), Ludlow, Sherriff & Hicks 19052 (BM). H. ludlowii shows a reduction trend westward from Yunnan to Bhutan. The easternmost populations are rather similar to the 'inter- mediate' population of H. himalaicum in northern Myanmar and adjacent India, but differ in the smaller overall size and delicate habit and in the size of the leaves and floral parts, as well as in the absence, in the more reduced forms, of black glands (except in the anthers). The upper leaves in this easternmost population, however, are 'trigonous' as in typical H. trigonum. From H. himalaicum it differs in the relatively longer styles and narrower acute sepals that are not black-streaked, and it is usually distinct in the smaller flowers, shorter ascending stems and boggy habitat. The styles are not absolutely longer, but the smaller ovary makes them relatively so. It would therefore appear probable, for morphological as well as geographical reasons, that H. ludlowii is related directly to H. trigonum, not via the H. himalaicum 'intermediate' population. 6. Hypericum daliense N. Robson, sp. nov. Map 12. 83 H. wightiano Wall, ex Wight & Arn. affinis, sed foliis maioribus, omnibus sessilibus, elliptico-oblongis integris, floribus maioribus, capsulis cylindrico-ellipsoideis, inter alia differt. Type: China, Yunnan, Tali Range, east flank, 2400-2700 m, July 1906 (1. fl), Forrest 4291 (BM!-holotype; K!-isotype). Perennial herb 0.15-0.4 m tall, erect from creeping and rooting base, with stems single or few, unbranched (always?). Stems 4- angled when young, soon terete or faintly 2-lined, eglandular; internodes 2-8 mm, exceeding leaves. Leaves sessile; lamina 22^45 x 7-21 mm, elliptic-oblong to (upper) lanceolate, paler beneath, not glaucous, plane, thinly chartaceous; apex rounded, margin entire, base rounded-amplexicaul; venation: 4 pairs of main laterals (lowermost weak) from lower 2/5 of midrib, with tertiary reticula- tion not prominent, rather dense; laminar glands pale, dense, small, punctiform; intramarginal glands dense, black. Inflorescence 5-14-flowered, from 1-2 nodes, densely capitate-subcorymbiform; pedicels c. 3 mm; bracts and bracteoles lanceolate to narrowly elliptic, with margin and auricles reddish- to black-glandular-ciliate or rarely subentire with a few intramarginal black glands, persistent. Flowers c. 10-15 mm in diam., stellate; buds ellipsoid. Sepals 5, 4.5-6.5 x 1-1 .5 mm, free, subequal or equal, erect in bud and fruit, lanceolate to narrowly oblong-elliptic, acute, glandular-denticulate; veins 5, outer pair branched; laminar glands all pale, linear or usually distal part(s) black, punctiform or striiform; marginal glands reddish or black, on denticles. Petals 5, bright yellow, not tinged red in bud, 6-8 x 2.5-3.5 mm, oblong-oblanceolate, margin entire, laminar glands absent, marginal glands few, black, subapical, ses- sile. Stamens 25-35, '3'-fascicled, longest 5-6 mm, c. 0.75-0.8 x petals; anther gland black. Ovary 3-locular?, c. 2 x 1.5 mm, ellip- soid; styles 3, 2-2.5 mm, equalling or slightly longer than ovary, free, divaricate; stigmas narrowly capitate. Capsule c.l x 4.5 mm, cylindric-ellipsoid, slightly exceeding sepals; valves with dense longitudinal vittae. Seeds not seen. 2n = ? 'Open situations amongst scrub in side valleys on the eastern flank of the Tali range' (Forrest 4297); 2400-3100 m. China (north Yunnan). CHINA. Yunnan: Tali [Dali] range, lat. 25 40' N, 2400-2700 m, July 1906 (fl & fr), Forrest 4297 (BM, K); Dali, Mt Dian-chang Shan, E. slope of Zong- he-fong, 2500-3 100 m, 3 August 1 990 (e. fl.), Murata et al. 68 (A); Lichiang [Lijiang] Range, March 1933 (fl & fr), McLaren's Collectors 176B (BM. K): 'Duplicate of 1912-1913', August 1917 (fl), Forrest 15853 (K, W). H. daliense resembles a large form of//, wightianum (to which it is clearly closely related), differing from that species inter alia by its larger flowers, larger, all sessile and mostly elliptic-oblong leaves (which are all entire, i.e. the upper ones are not basally glandular- fimbriate) and larger cylindric-ellipsoid capsule. It appears to provide a morphological link between H. wightianum and the relatively broad-leaved, few-flowered, south-western form of H. przewalskii, and to differ from the other relatives of that form ( 1 . H. monanthemum and 3. H. trigonum respectively) by the combination of numerous small flowers with glandular-denticulate (not glandular-ciliate to entire) sepals and relatively large elliptic-oblong to elliptic leaves. 7. Hypericum wightianum Wall. [Numer. List, no. 4818 (1831)] ex Wight & Arn., Prodr. Fl. Ind. Or.: 99 (1834); Wight, ///. Ind. Bot. 1: t. 34 (1838); Gamble, Fl. Pres. Madras 1: 70 (1915); Fyson, Fl. Nilgiri & Pulney Hill-tops 1: 38 (1915), Fl. S. Indian Hill Stns 1: 47 (1932); N. Robson in J. Jap. Bot. 52: 286, ff. 3^ (1977); Nair & Henry, Fl. Tamil Nadu, ser. 1, 1: 27 (1983); Saldanha, Fl. Karnataka 1: 209 (1984); Robson & Long, Fl. Bhutan 1(2): 377 (1984); Li Xiwen in Fl. R. P. Sinicae 50(2): 55, 84 N.K.B. ROBSON t. 1 1 ff. 1-2 ( 1 990); Biswas in Sharma & Sanjappa, Fl. India 3: 78 (1993), in Fl. West Bengal 1: 265 (1997). Type: India, Madras [Tamil Nadu], ex Herb. Wight., n.d. (fr), Wallich 4818 (K-W!- lectotype, Robson 1977; K!); Madras [Tamil Nadu], Neelgherries, 1828? (fl & fr), Wight 336 (BM!, K!-syntype). Fig. 13C, Map 14. H. rubrum Wight ex Dyer in Hook, f., Fl. Brit. India 1: 255 ( 1 874) pro parte quoad spec. cit. excl. typum. Type as for//, humifusum L. H. humifusum sensu Dyer in Hook, f., FL Brit. India 1: 255 (1874); Gamble, Fl. Pres. Madras 1: 70 (1915); Alston in Trimen, Fl. Ceylon 6: 19 (1931); pro parte omnes excl. typum. H. napaulense sensu Dyer in Hook, f., Fl. Brit. India 1:256 (1874) pro parte quoad syn. H. wightianum et spec. Maderaspatana; H Leveille in Bull. Soc. Bot. France 54: 594 (1908); Gagnepain in Lecomte, Fl. Gen. Indo-chine 1: 286 (1909); Hand.-Mazz., Symb. Sin. 7: 402 (1931); Rehder in J. Arnold Arbor. 18: 225 (1937); Lauener in Notes Roy. Bot. Card. Edinburgh 27: 4 (1966). H. bodinieri H. Leveille & Vaniot in Bull. Soc. Agric. Sarthe 39: 322 (1904); H. Leveille in Bull. Soc. Bot. France 54: 594 (1908), Fl. Kouy-Tcheou: 198 (1914). Type: China, Yunnan, frontiere du Kouy-Tcheou a Kian-ty, bord du fleuve, rive du Yunnan, 9 September 1897 (fl), Bodinier 1517 (P? or E?). H. elodeoides sensu R. Keller in Bot. Jb. 33: 553 (1904). H. delavayi R. Keller in Bot. Jb. 44: 49 (1909); Anon., Iconogr. Cormoph. Sin. 2: 876, f. 3482 (1972). Type: China, Yunnan, Tschen-Fong-Chan, Delavay 5180 (G-BOIS-holotype). H. monanthemum sensu Pax & Hoffm. in Repert. Spec. Nov. Regni Veg.,Beih. 12:438(1922). H. nepaulense sensu Craib, Fl. Siam. Enum. 1: 111 (1925). H. wightianum subsp. wightianum sensu N. Robson in J. Jap. Bot. 52: 286, ff. 3.4 (1977); Biswas in Sharma & Sanjappa, Fl. India 3: 80, f. 28(1993). H. humifusum subsp. humifusum sensu Biswas in Sharma & Sanjappa, Fl. India 3: 67 (1993) pro parte excl. typum et distrib. Europ. Icones: Wight, ///. Ind. Bot. 1: t. 34 ( 1 838); Anon., Iconogr. Cormoph. Sin. 2: 876, f. 3482 (1972); Li Xiwen in Fl. R. P. Sinicae 50(2): 56, t. llff. 1-2(1990). Perennial or annual herb (0.08-)0. 13-0.45 m tall, erect to decum- bent or procumbent from creeping and rooting base, with stems single or few, clustered, usually branched above but rarely through- out, with branches short, spreading-ascending. Stems shallowly 2-lined or terete, eglandular; internodes 9-50(-75) mm, usually longer than leaves. Leaves sessile to shortly petiolate (especially lower ones); lamina (6-)10-30 x (3-)5-15(-19) mm, broadly ellip- tic to obovate or ovate, paler beneath, not glaucous, plane, thinly chartaceous; apex rounded or rarely acute to apiculate-obtuse (Meghalaya), margin entire or (upper) sometimes basally or wholly reddish- to black-glandular-ciliate and with reddish- to black-glan- dular-ciliate auricles, base rounded to cordate; venation: 2-3 pairs of main laterals from lower third of midrib, with tertiary reticulation scarcely visible beneath, dense; laminar glands pale or rarely a few black, punctiform to striiform, varying in size; intramarginal glands dense or irregular, black. Inflorescence \-c. 50- flowered, from 1- 2(3) nodes, the whole laxly corymbiform or broadly pyramidal to capitate-cylindric or bifurcated; pedicels 2-3 mm; bracts and bracteoles ovate or lanceolate to linear, with black-glandular-ciliate margin and auricles, persistent. Flowers 5-8(-ll) mm in diam., Map 14 Sect. 9e: 7. H. wightianum . other records O. STUDIES IN HYPERICUM 85 stellate; buds ellipsoid, subacute. Sepals 5, 2.5-5(-6) x 1.5-3 mm, equal, narrowly to broadly oblong or elliptic, acute to obtuse, glandular-ciliate to -laciniate or very rarely entire; veins 3(5), unbranched; laminar glands all pale or some black, linear to puncti- form, scattered; marginal glands reddish to black, on cilia, sometimes alternating with intramarginal black gland dots, or very rarely all intramarginal. Petals 5, bright yellow, not tinged red in bud, 3-5 x 1.2-1.6 mm, c. 1-1.2 x sepals, elliptic-oblong, rounded to acute, margin entire or apically glandular-ciliate, laminar glands absent, marginal glands black, few, distal and subapical, sessile. Stamens c. 7-11, '3' -fascicled, longest 2.5^4- mm, c. 0.8-0.9 x petals; anther gland black. Ovary 1-locular, 1.5- 3 x c. 1 mm, ovoid to globose; styles 3, 1.5-2.5 mm, 0.9-1 x ovary, divergent; stigmas narrowly capitate. Capsule3-4.5(-6)xc. 34 mm, broadly ovoid to subglobose, equalling or slightly exceeding sepals; valves with c. 7-9 longitudinal vittae. Seeds brown, c. 0.5 mm; testa finely scalariform. 2n = ? Grassy slopes, open woodland, streamsides, marshes, roadsides and rice paddy terraces; 750-3300 m. South China (Guangxi, Sichuan, Guizhou, Yunnan, Xizang), north Laos, north Thailand, Myanmar (Kachin, Chin, Shan), north-east India (Nagaland, Manipur, Mizoram, Meghalaya, Benghal, Sikkim), Bhutan, south India (Tamil Nadu) and Sri Lanka. CHINA. Guangxi: Ling Yun Hsien, Loh Hoh Tsuen, 1 1 50 m, 8 June 1 933 (fl & fr), Steward & Cheo 616 (GH, NY); Lingyun Xian, 7 April 1936 (fl), Guangxi Prov. Mus. 1 1404 (IBSC). Sichuan: Huidong Xian, 1750 m, 3 July 1978 (fl & fr), Zhao Q.S. 5828 (SZ); Leibo Xian, Shazigou, 1600 m, 27 May 1959 (fl & fr), Liangshan WildEcon. PL Exped. 0375 (CDBI). Guizhou: Lo- fou, Pin-fa [Pingfa], April 1906 (fl), Cavalerie 2768 (K); Anlong Xian, 1280 m, 26 May 1960 (fl & fr), Zhang & Chang 3095 (IBSC). Yunnan: East flank of Tali Range, 25 40' N, 3000-3300 m, August 1906 (fl), Forrest 4298 (BM, K); Anning Xian, Wenquan Gongshe, 1890 m, 4 July 1965 (fr), Wu CK 122 (KUN); Pingbian Xian, 1350 m, 20 May 1934 (fl & fr), Tsai H.T. 55271 (A). Xizang: Gyigang [Zayii] Xian, 2350 m, 30 August 1976 (fl & fr), Wu et al. 5266 (KUN). LAOS. Louang Prabang: Luang-prabang [Louangphrabang], Thorel (fide Gagnepain, 1909). THAILAND. Northern: Summit of Doi Angka, 2500 m, 19 August 1927 (fr), Garrett42\ (ABD, BM, C, K); Doi Pa Kao, c. 1600 m, 8 May 1921 (fl), Kerr 5392 (ABD, BK*, BM, K); N. Chiangmai, Doi Inthanon, 2500 m, 30 October 1962 (fr), Smitanand, Poore & Robbins 7643 (BK, F). MYANMAR. Kachin: Irrawadi basin near Tsu-Yo-Ho, 26 N, 1800 m, October 1905 (fr), Forrest 829 (BM, K); Kachin Hills, Palaungketaung, 1050 m, 13 April 1912 (fl), Lace 5789 (K). Shan: Kalaw, 1 May 1931 (fr), Dickason 5707(GH); Namkham, May 1933 (fl & fr), Dickason 5970 (GH). Chin: Mindat, 1350 m, 5 July 1956 (fl & fr), Kingdon Ward 2244B (BM). INDIA. Nagaland: Naga Hills, Kohima, n.d. (fr), Prain s.n. (BM). Manipur: Ukhrul, 1650 m, 13 March 1948 (fl), Kingdon Ward 17083 (A, BM, NY*); Myang Khong, 900-2100 m, February 1882 (fl & fr), Watt 6051 (K). Mizoram: Lushai Hills, Champnai, 1 500 m, July 1 926 (fl & fr), Parry 63 (K). Meghalaya: Khasi Hills, Shillong Peak, 1500-1800 m, 28 July 1949 (fl & fr), Kingdon Ward 18677 (A, BM, NY*); Khasi Hills, Barapani, c. 900 m, 18-27 April 1954 (fl), Chand 754 (L). Benghal: Kalimpong, 1200 m, July 1882 (fr), Gamble 10471 (K); Darjeeling, Laebong, 1500 m, 22 June 1884 (fr), Clarke 35424 (K). Sikkim: Rungbee, 1350 m, 5 June 1870 (fl & fr), Clarke 11793 (BM). Tamil Nadu: Nilgiris Distr., Ootacamund [Udagamandalam], 2100 m, June 1886 (fl & fr), Gamble 14348 (K); Palni Hills, Kodaikanal Distr., Madurai, Berijam, 2100 m, 19 November 1985 (fl & fr), Matthew RHT 42834 (BM, RHT*); Anamally Hills [Anaimalai], pre- 1885 (fl), Beddome 388 (BM). SRI LANKA. Central: Nuwara Eliya Distr., Pattipola 1980 m, 21 June 1972 (fl & fr), Hepper444l (K). BHUTAN. South: Chukka Distr., Chukka and Marichong. Central: Thimpu Distr., Thimpu Chu Valley; Mongar Distr., Sengor. All fide Fl. Bhutan: 378. H. wightianum is clearly closely related to 7. H. daliense and has apparently spread from south-east China westward to Sikkim and south-westward to western Myanmar and Mizoram in India. It then reappears in the south Indian hills and in Sri Lanka. Morphological variation follows these trends: ( 1 ) The erect to basally decumbent form extends along the Himalaya and also occurs in Meghalaya, Myanmar and the Nilgiri Hills. It has sessile leaves, the upper ones having near-basal glandular cilia and glandular auricles. (2) In Thailand form (1) gives rise to a decumbent form with slender stems and shortly petiolate leaves, the inflorescence being a small version of that of form (1). (3) A form similar to (2) occurs in south India (Kodaikanal), except that the leaves vary from sessile to petiolate. This gives rise to a form in the Anaimallai and Nilgiri Hills with petiolate entire leaves and entire sepals and bracts (e.g. Beddome 388) ('//. rubrum'or '//. humifusum' of Indian Floras). One of these Anaimallai specimens (Beddome 386) has sessile leaves. All these variations are continuous. The form that is most similar to 6. H daliense occurs in Khasiya (Meghalaya). ACKNOWLEDGEMENTS. My studies for Part 4(1) have been especially helped by Dr Peter Raven and Dr Ihsan Al-Shehbaz, who organised loans from ten Chinese and four US herbaria and enabled me to have an enjoyable and productive stay in MO to work on them; and by Yang Guang (NAS), who translated most of the labels from Chinese, verifying the data and compiling a database from all those specimens that I had named. The work on the database was co-ordinated by Dr and Mrs William Thai, to whom I am also grateful. The treatments of Hypericoideae for various Chinese Floras by Prof. Li Xiwen (KUN), and his advice, have been very useful. I should like to acknowledge the assistance of various colleagues in the Herbarium and Library at BM, in particular Mike Gilbert, for his expertise on the Flora of China, distributional data and computer matters, Prof. Chris. Humphries, for producing the maps on computer, and Malcolm Beasley, for much library assistance and, especially, for obtaining translations of Chinese labels from the Gunghwa Co. Ltd, Chinese booksellers. My thanks are also due to the directors of the following herbaria for loans and (to some) for study facilities: A. BO, CAS, CDBI, E, Fl, G, GH, HNWP, IBK, IBSC, IFP, JE, K, KATH, KUN, L. LE, MO, NAS, NY, RAW, SING, SZ, TAI, W, WH and WUK. I am deeply indebted to Mrs Margaret Tebbs for most of the drawings and to Phil Rye for the remaining two. My wife Eve has, as always, been a help and support in many ways, especially during our stay at MO. Finally, I must thank two Keepers of Botany, Prof. Stephen Blackmore and then Prof. Richard Bateman, for study facilities in the Botany Department, and the Stanley Smith Trust for a grant towards illustration costs. REFERENCES Alston, A.H.G. 1931. Hypericum. In H. Trimen, Flora of Ceylon 6: 19. Axelrod, D.J., Al-Shebaz, I. & Raven, P.M. 1998. History of the modern flora of China. In A.-L. Zhang & S.-G. Wu, (Eds), Floristic characteristics and diversity of East Asian plants: 43-55. Beijing, Berlin, etc. Brown, M.L. & Brown, R.G. 1984. Herbaceous plants of Maryland. Baltimore. Coulter, J.M. 1 886. Revision of North American Hypericaceae. - 1. Botanical Gazette 2:78-88. Crovello, T.J, Keller, C.A. & Kartesz, J.T. 1983. The vascular plants of Indiana; a computer based checklist. Notre Dame. Dyer, W.T.T. 1874. Hypericinae. In J.D.Hooker, Flora of British India 1: 252-258. Gibson, A.C. 1980. Wood anatomy of Thomea, including some comparisons with other Hypericaceae. International Association of Wood Anatomists Bulletin, N.S. 1: 87-92. (illicit, J.M. & Robson, N.K.B. 1981. The St-John's-worts of Canada (Guttiferae). Publications in Botany, National Museum of Natural Sciences of Canada, No. 11. Ottawa. Gorschkova, S.G. 1949. Guttiferae. In B.K. Shishkin & E.G. Bobrov (Eds), Flora S.S.S.R. 15: 2011-258. 86 N.K.B. ROBSON Grubanov, LA. 1996. Konspekt Flora Vneshnei Mongoliy (Sosudstie Rasteniya). Moscow. Guo, S.-Z. 1 98 1 . On the elevation and climatic changes of the Qinghai-Xizang (Tibet) Plateau based on fossil angiosperms. Studies of Qinghai - Xizang Plateau 1: 201- 205. Beijing. (Not seen; cited from Axelrod, Al-Shebaz & Raven, 1998). Hance, H.F. 1865. Descriptions of four new plants from southern China. Journal of Botany, London 3: 378-381. Handel-Mazzetti, H. 193 1 . Symbolae Sinicae, 7(2) Anthophyta: 401-404 (Guttiferae). Hoar, C.S. & Haertl, E.J. 1932. Meiosis in the genus Hypericum. Botanical Gazette 93: 197-204. Hsii, J. 1983. Late Cretaceous and Cenozoic vegetation in China, emphasizing their connections with North America. Annals of the Missouri Botanical Garden 70: 490- 508. Hulten, E. 1929. Flora of Kamtchatka and the adjacent islands. Guttiferae. Kungliga Svenska Vetenskapsakademiens Handingar, ser. 3, 8(1): 125-127. Jones, S.W. 1980. Morphology and major taxonomy o/Garcinia (Guttiferae). Unpub- lished thesis. University of Leicester. Keller, R. 1 925. Hvpericum. In A. Engler & K Prantl, Die naturlichen Pflanzenfamilien, 2nded. 21: 175-183. Kimura, Y. 1 95 1 . Hypericaceae. In Nakai, T. & Honda, M. (Eds), Nova Flora Japonica 10. Tokyo. 1966. Hypericaceae. In Hara, H., The Flora of Eastern Himalaya: 209-211. Tokyo. Kogi, M. 1984. A karyomorphological study of the genus Hypericum (Hypericaceae) in Japan. Botanical Magazine (Tokyo) 97: 333-343. Komarov, V.L. 1950. Flora Man'chzhurii. Guttiferae. 3: 45-50. Moscow. Krasnoborov, I.M., Rostovtseva, T.S. & Ligus, S.A. 1 980. Chromosome numbers of some plant species of South Siberia and the Far East. Botanicheskiy Zhurnal 65: 659-668. (In Russian) Krogulevich, R.E. 1978. Karyological analysis of the species of the flora of eastern Sayana. In L.I. Malyshev & G.A. Peshcova (Eds), Flora Cisbaikalia: 19^8. Novosibirsk. (In Russian) Krukenberg, A.R. 1983. Temperate floras: the north Pacific connection. Annals of the Missouri Botanical Garden 70: 591-596. Li, H.-L. 1944. New or noteworthy plants from southwestern China. Journal of the Arnold Arboretum 25: 299-318. 1952. Floristic relationships between eastern Asia and eastern North America. Transactions of the American Phiosophical Society N.S. 42: 371^4-29. Reprinted with fore ward (1971) in Morris Arboretum Monographs. Philadelphia. Li Xiwen 1990. Guttiferae. In Li Xiwen (Ed.), Flora Reipublicae Popularis Sinicae 50(2): 1-112. Beijing. McKenna, M.C. 1972. Was Europe connected directly to North America prior to the Middle Eocene? Evolutionary Biology 6: 179-189. Magee, D.W. & Ahles, H.E. 1999. Flora, of the Northeast. Amherst, Mass. Malakhova, L.A. 1990. Kariologocheskiy analiz prirodnykh populaciy redkikh i ischezajushchikh rasteniy na juge Tomskoy Oblasti. Byulleten' Glavnogo Botanicheskogo Sada 155: 60-66. & Markova, G.A. 1994. Chromosome numbers on flowering plants of Tomsk Region. Dicotyledones. Botanicheskiy Zhurnal 79(12): 103-106. (In Russian) Maximowicz, C.J. 1882. Diagnoses plantarum novarum asiaticum, 4. Melanges biologiques 11: 155-350. Melville, R. 1976. The terminology of leaf architecture. Taxon 25: 549-561. 1 983. Remoration: an overlooked process in Angiosperm evolution. Kew Bulletin 37: 613-639. Momiyama, T. 1982. Guttiferae. In Y. Satake et al. (Eds), Wild Flowers of Japan, herbaceous plants (including dwarf subshrubs) 2: 13-119. Tokyo. Nielsen, N. 1924. Chromosome numbers in the genus Hypericum (A preliminary note). Hereditas 5: 378-382. Nishikawa, T. 1990. Chromosome counts of flowering plants of Hokkaido (13). Journal of Hokkaido University, Education, 2B, 40: 19-30. Probatova, N.S. & Sokolovskaya, A.P. 1983. New chromosome numbers for vascular plants from the islands of Peter the Great Bay (Primorye territory). Botanicheskiy Zhurnal 68: 1655-1662. (In Russian). 1 986. Chromosome numbers of the vascular plants from the far east of the USSR. Botanicheskiy Zhurnal 71: 1572-1575. (In Russian) Raven, P.H. & Axelrod, D.I. 1974. Angiosperm biogeography and past continental movements. Annals of the Missouri Botanical Garden 61: 539-673. , Kyhos, D.W. & Hill, AJ. 1965. Chromosome numbers of Spermatophytes, mostly Californian. Aliso 6: 15-113. Robson, N.K.B. 1961. Guttiferae. In A.W Exell. & H. Wild (Eds), Flora Zambesiaca 1: 378^04. 1977. Studies in the genus Hypericum L.(Guttiferae). 1. Infrageneric classifica- tion. Bulletin of the British Museum (Natural History), Botany 5: 291-355. 1981. Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the genus. Bulletin of the British Museum (Natural History), Botany 8: 55-226. 1985. Studies in the genus Hypericum L. (Guttiferae). 3. Sections 1 . Campylosporus to 6a. Umbraculoides. Bulletin of the British Museum (Natural History), Botany 12: 163-325. 1996. Studies in the genus Hypericum L. (Guttiferae). 6. Sections 20. Myriandra to 28. Elodes. Bulletin of the Natural History Museum, Botany 26: 75-217. Sandhu, P.S. & Mann, S.K. 1989. SOCGI plant chromosome number reports - VIII. Journal of Cytology and Genetics 24: 179-183. Stepanov, N. V. 1 994. Chromosome numbers of some higher plants taxa of the flora of Krasnoyarsk region. Botanicheskiy Zhurnal 79: 135-139. Stevens, P.P. (in press) Clusiaceae. In K. Kubitzki (Ed.), The families and genera of vascular plants. Berlin. Sugiura, T. 1944. Studies on the chromosome numbers in higher plants. VI. Cytologia 13: 352-359. Woronov, G. 1906. Guttiferae. In N.Y. Kuznetsov, N. Busch & A. Fomin (Eds), Flora caucasica critica III, 9 : 1-65. STUDIES IN HYPER1CUM 87 SYSTEMATIC INDEX Accepted names are in roman and synonyms in italics; new names and principal references are in bold. An asterisk denotes a figure. A = sect. Ascyreia, B = sect. Bupleuroides, C = sect. Concinna, E = sect. Elodeoida, M = sect. Monanthema, R = sect. Roscyna, S = sect. Sampsonia. Axcyrum L. filicaule Dyer (= Mlb) 77, 78 sibiricum Lam. ex Poiret (=Rla) 54 tetragonum Moench (=R 1 ) 53 Clusioideae Engler 40 Cratoxyleae Engler 38 Cratoxylum Blume 38 Eliea Cambess. 38 Garcinia L. 40 Hypericoideae-Cratoxyleae 38 HypericumL. 37,38,40,41 sect. Adenosepalum sensu N. Robson, pro parte 66 sect. Androsaemum (Duhamel) Spach subsect. Pseudandrosaeinum R. Keller, pro parte 61 sect. Ascyreia Choisy 40, 44, 49 sect. Bupleuroides Stef. 37, 49 sect. Concinna N. Robson 43, 61 sect. Drosocarpium Spach 42, 66 sect. Elodeoida N. Robson 42, 43, 45, 46, 66 sect. Elodes (Adans.) W. Koch 47 sect. Graveolentia N. Robson 42, 43, 63 sect. Euhypericum Godron 41 sect. Hypericum 41,42 sensu N. Robson 61, 63, 66 subsect. Erecta N. Robson 42, 43 subsect. Hypericum 42 series Senanensia N. Robson 42, 43 sect. Inodora Stef. 40 sect. Monanthema N. Robson 42, 43, 45^17, 75 sect. Myriandra (Spach) R. Keller 37 sect. Norysca (Spach) R. Keller 40 sect. Oligostema (Boiss.) Stef. 47 sect. Roscyna (Spach) R. Keller 41, 44, 52 sect. Sampsonia N. Robson 42, 43, 45, 46, 63 sect. Trigynobrathys (Y. Kimura) N. Robson 37 adenophorum Wall, ex Dyer (=E4a) 72 aethiopicum Thunb. subsp. sonderi (Bredell) N. Robson 37 amplexicaule Lam. (=Rlc) 58 ascymides Willd. (=Rlc) 58 ascyronL. (Rl) 41, 45, 52, 55*, 63 sensu Coulter (=Rlc) 58 subsp. ascyron (Rla) 44, 45, 54, 55* subsp. gebleri (Ledeb.) N. Robson (Rlc) 44, 45, 53, 55*, 57, 63 subsp. pyramidatum (Aiton) N. Robson (Rlc) 42, 53, 55*, 58, 63 var. adamii R. Keller (=Rla) 56 var. americanum (=Rlc) 58 var. angustifolium (=Rla) 54 var. ascyron 56 var. brevistylum Maxim. (=Rlb) 54, 57 var. genuinum Maxim. (=Rla) 53 var. giraldii (=R\a.) 53,54 var. hupehensis Pamp. (=R 1 a) 54 var. longistylum Maxim. (= Rla) 53, 54 var. macropetalum hort. ex Vilmorin & D. Bois. 54 var. macmsepalum Ledeb. (=Rla) 53, 54 var. micmpetalum R. Keller (=Rla) 54 var. micmpetalum sensu R. Keller (=A16a) 49 var. punctato-striatum R. Keller (=A16a) 49 var. typicum R. Keller (=Rla) 53 var. umbellatum R. Keller (=Rla) 53, 54 var. vilmorinii. Render (=R la) 54 forma angustifolium Y. Kimura (=Rla) 53 forma vilmorinii (Rehder) Rehder (=Rla) 56 assamicum S.N. Biswas (S2) 42, 46, 48, 64*, 66 bachii (=Mla) 78 hartramium Miller (=Rlc) 58 bequaertii De Wild. 40 biondii R. Keller, pro parte (=Rla) 59 hiondii R. Keller, pro parte (=R2) 54 bodinieri H. Leveille & Vaniot (=M7) 84 bracteatum Kellogg (=C 1 ) 62 bupleuroides Griseb. (Bl) 37,50,51* centiflorum H. Leveille (=E2a) 70 chinense L. var. minutum R. Keller (=R2) 59 concinnum Benth. (Cl) 41, 45, 55*, 61, 63 cordifolium Choisy 44 daliense N. Robson (M6) 47, 49, 83, 85 delavayi R. Keller (=M7) 84 elatoides R. Keller (A16a) 37, 40, 49, 51* electrocarpum Maxim. (=S1) 63 forma parvifolium (=S 1 ) 63 elegans Steph. ex Willd. 42 ellipticifolium H.L. Li) (=Lianthus ellipticifolius) 38 ellipticum Hook. 37 elodeoides Choisy (E4) 42, 43, 46, 47, 72, 73* sensu Hand.-Mazz. (= El) 69 sensu R. Keller (=M7) 84 subsp. elodeoides (E4a) 48, 73*, 74 subsp. wardii N. Robson (E4b) 48, 73*, 74 elodes L. 47 erectum Thunb. 43 esquirolii H. Leveille (=S1) 63 filicaule (Dyer) N. Robson (=Mlb) 78 fl. pentagvnLi, caule tetragono (etc.) Gmelin (=R1) 53 gebleri Ledeb. (=Rlb) 53, 57 graveolens Buckley 42 hemsleyanum H. Leveille & Vaniot (=la) 53, 54 hengshanense W.T. Wang (E3) 46, 48, 69, 72, 74 var. xinlinense Z.Y. Li (=E3) 72 himalaicum N. Robson (M4) 47, 48, 80*. 81-83 sensu Li Xiwen, pro parte (=M5) 82 humifusum L. 47 sensu Dyer (=M7) 84 sensu Y. Kimura (=M4) 81 subsp. humifusum sensu Biswas 84 subsp. orbiculatum S.N. Biswas (=E2b) 71 kamtchaticum Ledeb. 41 var. pibairense Miyabe & Y. Kimura 42 kelleri H. Leveille (=Rla) 41, 54 kingdom! N. Robson (E5) 46, 74 laleriflorum H. Lveille (=E1) 69 longifolium H. Leveille (= Rl) 54 ludlowii N. Robson (M5) 47, 48, 80*, 81, 82, 83 macrocarpum Michx. (=Rlc) 53,58 macmsepalum Rehder (=R2) 42, 46-48, 59, 61 maculatum Crantz subsp. immaculatum (Murb.) A. Frohl. 42 mairei H. Leveille (=E2a) 70 maireiH. Leveille (=M la) 78 majus (A. Gray) Britton 37 monanthemum Hook. f. & Thorns, ex Dyer (Ml) 42,43, 46, 75, 76*, 81,83 sensu Li Xiwen, pro parte (=M3) 79 sensu Li Xiwen, pro parte (=M5) 82 sensu Pax & Hoffm. (=M7) 84 subsp. filicaule (Dyer) N. Robson 46-48, 76*, 78 subsp. monanthemum 46, 48, 76*, 77, 79 var. brachypetalum Franchet (=M4) 81 var. nigmpunctatum Franchet (=Mla) 77 Monanthemum group 81 monogynum L. 40, 50 \ar.franchetii Baroni (=A16a) 49 mororanense R. Keller, pro parte 79 napaulense Choisy (=E4a) 72 sensu Dyer, pro parte (=M4) 81 sensu Dyer, pro parte (=M7) 84 nepaulen.te sensu Craib 84 nervosum D. Don (=E4a) 72 nikkoense Makino 42 obtusifolium R. Keller (=R2) 59 ocymoides Loddiges (=Rlc) 58 oshimaen.se R. Keller (=S1) 65 pollens D. Don (=E4) 72 sensu D. Don pro parte (=M4) 81 paradoxum R. Keller 41 pedunculatum R. Keller (=R2) 41, 42, 47. 48, 59, 60*. 61 perfoliatum Ledeb. (=B 1 ) 50 perforatum L. 42 petiolaium sensu R. Keller (=E2) petiolulatum Hook. f. & Thorns, ex Dyer (E2) 69 subsp. petiolulatum (E2b) 46, 48, 68*, 71 subsp. yunnanense (Franchet) N. Robson (E2a) 46, 48, 68*, 70 var. orbiculatum Franchet (=E2b) 7 1 var. subcordatum (R. Keller) H. Leveille 79 podocarpoides N. Robson 37, 41, 44, 45 prattii N. Robson 41 przewalskii Maxim. (R2) 41, 42, 44, 47. 48, 59, 60*. 66, 81.83 macrosepalum form 60*, 61 pedunculatum form 60*. 61 pseudomaculatum Bush 42 pseudopetiolatum R. Keller var. grandiflorum Pampanini (=E1) 70 pyramidatum Aiton (=Rlc) 54, 58 revolutum Vahl subsp. keniense (Schweinf.) N. Robson 40 rubrum Wight ex Dyer (=M7) 84 sachalinense H. Leveille (=Rlb) 41, 57 sagittifolium Koidz. (=Rla) 56 salicaria Reichenb. (=Rla?) 54 sampsonii Hance (SI) 42, 43, 46, 48, 63, 64* sensu Dyer, pro parte (=S2) 66 scallanii R. Keller (=Rla) 54 scouleri Hooker 42, 43 seleriR. Keller (=C1) 62 senanense Maxim. 43 seniawinii, Maxim. (El) 42, 46, 48, 67, 68*, 69. 72,81 setosum Wall, ex Dyer (=M4) 81 subcordatum (R. Keller) N. Robson 47, 48, 76*. 78 thomsoni R. Keller (=E2) 69 var. subcordatum (=M2) 78 sensu R. Keller pro parte (=M2) 78 trigonum Hand.-Mazz. 47, 48, 79, 80*, 81, 83 Trigonum group 47, 8 1 wightianum Wall, ex Wight & Arn. 47, 49, 73*, 83, 85 subsp. axillare N. Robson (=E5) 74 subsp. wightianum sensu N. Robson (=M7) 84 xylosteifolium (Spach) N. Robson 50 yabei H. Le\eille & Vaniot (=Rla) 54 yamamotoi var. riparium Y. Kimura 42 vunnanense Franchet (=E2a) 70 yunnanense Franchet pro parte (=Mla) 77 88 N.K.B. ROBSON Lianthus N. Robson 38, 40 ellipticifolius (H.L. Li) N. Robson 38, 39* Roscyna Spach (=R) americana Spach 58 ascyron (L.) Y. Kimura (=Rlc) 53 gebleri (Ledeb.) Spach (=Rlb) 57 gmelinii Spach (=R1) 53 japonica Blume (=Rla) 54 Santomasia N. Robson 40 Thornea Breedlove & McClintock 38, 40 Thornea group 38, 40 Thymelaeaceae 40 Triadenum Raf. 38, 40 brevifolium (Wall, ex Dyer) Y. Kimura 40 japonicum (Blume) Makino 40 virginicum (L.) Raf. 40 Bw//. r. Mus. Lond. (Bot.) 31(2): 89-106 Issued 29 November 2001 A bibliography of the scientific work of Boris V. Skvortzov (1896-1980) with commentary on the publications concerning diatoms (Bacillariophyta) DAVID M. WILLIAMS AND GERALDINE REID Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD CONTENTS Introduction 89 A short biography 90 A summary of Skvortzov's diatom collecting localities 91 China 91 Russian Federation 94 Kazakhstan 95 South Korea 95 Japan 95 North America 96 Australia (Fossil) 96 New Zealand 96 Philippines 96 Africa 96 Cuba 96 Afghanistan 96 Sri Lanka 96 India 96 B.V. Skvortzov's bibliography 96 References ... 106 SYNOPSIS. This paper presents a full bibliography for the naturalist Boris V. Skvortzov. Special attention has been given to his diatom (Bacillariophyta) studies. Skvortzov was fortunate to have had access to material from many areas which had never been studied for their diatom flora. In this paper we present a detailed list of those localities. INTRODUCTION The scientific career of Boris Skvortzov [Skvortsow, Skvortsov] (1896-1980) spanned nearly 70 years: he published his first paper in 1916 when just 20 years old and his last papers appeared posthu- mously between 1981 and 1985. As far as we have been able to establish, during that period Skvortzov published some 434 papers, notes and popular botanical accounts mostly relating to the area of Manchuria in north-east China. Skvortzov was predominantly inter- ested in microalgae, particularly diatoms, but published a great deal in many other botanical areas. Contemporary reaction to Skvortzov's work, at least from the perspective of phycologists, is somewhat mixed. Skvortzov did indeed have a penchant for naming many new taxa - indeed most of his floristic diatom accounts are predominantly descriptions of new taxa. Yet this in itself is not the largest obstacle to understanding and coming to some informed judgement of Skvortzov's work. He was able to examine material from parts of the world that had hardly, if ever, been studied before, but practically none of his material is available for study today, and very little has found its way into existing herbaria. Why not, then, simply consign his work to history, regard his new taxa as unverifiable and rely on current accounts made from the same areas? We believe that this option seems a little premature, although it is worth noting that some researchers study- ing flagellates have opted for that approach (Patterson, 1994: table 5). We have developed a slightly different view and this contribution is a preliminary start on gaining an understanding of Skvortzov's work and will ultimately contribute to a greater understanding of diatom diversity in north-east China. Our specific interest in Skvortzov developed from recent studies on Lake Baikal, Siberia under the auspices of a 'Darwin Initiative for the Survival of Species' study (Flower & Williams, 1999). The Darwin Initiative (DI) allowed us to make a systematic collection of diatom samples from the entire lake at regular intervals and regular depths. In addition, we had the opportunity to revisit and re-sample the localities Skvortzov examined. Skvortzov was not the first to write about and describe the many diatoms of Lake Baikal, but he was the first to recognize and identify many of its taxa as endemic. His diligence, with the deep water Lake Baikal sample at least, can The Natural History Museum, 2001 90 D.M. WILLIAMS & G. REID be appreciated by his comment that 'I have examined about a hundred microscopic slides from this place...', (Skvortzov, 1937:297). It would be surprising if anyone working today dealt with such large numbers of preparations from a single site. (Skvortzov (1936: 9) notes the same approach for his study of Kizaki Lake in Japan: 'I have examined a hundred microscopic slides... Half a year was spent in the study of this collection'.) Our own studies suggest that far from being rather optimistic concerning the diversity of Lake Baikal diatoms, Skvortzov may have under-estimated Lake Baikal's benthic diatom diversity (see Mann, 1999). If Skvortzov was reasonably accurate in this study, there seems no reason to doubt his work elsewhere - or at least until such time as relevant material becomes available and can be exam- ined. A second aspect that we felt required attention was that as Skvortzov had written some of the first accounts of north-east Chinese diatoms, and we were struggling to understand the biogeographic relations of the area relative to our Lake Biakal samples, a survey of the areas he did study was of significance. Considerably more attention is now being paid to the idea of regional diversity of diatoms and to their significance in biogeographic studies (Kociolek & Spaulding, 1999). Bearing these ideas in mind, in this paper we have tried to achieve three things that may act as a baseline for further examination and evaluation of Skvortzov's diatom studies. It is of some significance that Chin (1951:151) wrote: 'During the past hundred years Skvortzow was the most ardent worker.' ( 1 ) The biggest outstanding problem when dealing with Skvortzov's work is that very little of his original material is available. To address this issue, we have compiled a list of the material Skvortzov examined. A good deal of this material was given to him by other collectors and the possibility remains that some still exists in various herbaria around the world. We have been able to trace some original material in the diatom collections at the BM and are aware that other sub-samples exist in the California Academy of Science, Bremerhaven in Germany, and the Department of Paleobotany, Swedish Museum of Natural History. A list of geographical areas may allow new or recently collected material to act as neotypes should Skvortzov taxon names be confirmed. We have begun this process by designat- ing neotypes from the DI material for some Lake Baikal diatoms we have examined in detail (Reid & Williams, submit- ted). (2) We have compiled a bibliography of all his writings as far as we have been able to establish. A number of the entries have not been seen by us but have been taken from bibliographies Skvortzov published himself. Tracing Skvortzov's published work is a complex business simply because many of the places he chose to publish at that time were small local natural history societies whose existence was sometimes rather short, exacer- bated by the volatile nature of Manchuria during the years he was living and working there. Fortunately for us, many of these publications are held in the various libraries of The Natural History Museum, London (BM). We have numbered all the items in the bibliographic section and refer to these numbers in the remaining text. For instance, Skvortzov's three biblio- graphies can be found under items 300, 396 and 418. References to work not written by Skvortzov appear at the end of the paper and are referenced in the usual way. With respect to Skvortzov's publications, two articles have been published that review his contributions. Unfortunately, we have been unable to trace copies of either (Uedo, 1940; Baranov, 1959). (3) As our area of interest is diatoms, we have marked those publications which mention or list diatom species, new or otherwise. We have resisted the temptation to list all of Skvortzov's taxonomic names at this stage, simply because the large amount of Chinese literature needs to be taken into account and it would serve no immediate purpose other than to highlight various permutations of names. A SHORT BIOGRAPHY Although Skvortzov lived most of his life in Harbin, north-east China (he was born in Warsaw in 1896, moving to Harbin in 1902), he travelled to St. Petersburg during the years 1914 to 1917 to train as a diatomist under R.W. Kolbe (Selling, 1962) and S. Wislouch. With the exception of some time spent collecting material and learning English in Fuzhou, Fujien, China (1918-1919),' Skvortzov remained in Harbin until 1 962 when he departed for Brazil where he spent his remaining years. During his time at Harbin, he was actively involved in the Manchurian Research Society and was in charge of their crypto- gamic plants (Review of the Manchurian Research Society, 1 926: 13). On his return to Harbin from Fuzhou, Skvortzov noted that from 1919 to 1935 he taught botany. He was heavily involved with the Manchurian Research Society, acting as their secretary, which he probably undertook on a voluntary basis: 'the shortage of funds did not permit to engage a special personnel for the station, and all work was carried on voluntarily by a few members of the Manchuria Research Society in their leasure [sic] hours' (Pavlov, 1925:9). From 1923 to 1928, the period covered by written reports, Skvortzov gave numerous presentations, especially to the 'Young Archaeologists, Naturalists and Ethnographers Association of Harbin' (he gave two presentations in 1923, six in 1924 and seven in 1925). Skvortzov published most of his diatom studies between 1919 and 1939. After that time, and especially while living in Brazil, he worked and published mainly on flagellates. We have no expertise with those organisms and leave any evaluation of that work to those who do. We were unaware of any existing portraits until recently when we discovered a number in the various publications of the Manchuria Research Society. The reports of the society include a number of group photographs that include Skvortzov. In the Review of the Manchurian Research Society (1926, tables 2 and 3), Skvortzov appears in a group photograph of the 'Members of the M.R.S. committee presided over by Doctor Wang Tsin-Tschung. (1924)' and a group photograph of 'Members of the M.R.S. committee presided over by Doctor Hei-Show-Djen (1925)'. Other photo- graphs include Skvortzov with 'The American explorer prof, [sic] Hansen, seated amid the members of the Natural History section attached to the M.R.S.' (Fig. 2) and a portrait of Skvortzov partially hidden by a giant water lily (Figs 1 & 3). In a later report, there are group photographs of the Manchurian Research Society that includes Skvortzov; 'Vice-president of the Chinese Eastern Railway Board of Directors Mr. Lashevich sur- rounded by members of the Manchuria Research Society Committee after having viewed the Museum' and 'Manchuria Research Society Members of the Committee and Revisional Committee, elected in 1927' (Review of the Manchurian Research Society, 1928: tables 2 and 6). 'The Journal of the North China Branch of the Roval Asiatic Society (50:259) listed Skvortzov as a member, giving his address as 'c/o C.M. customs, Foochow'. BORIS V. SKVORTZOV 91 Fig. 1 Skvortzov holding a giant water lily. From the Review of the Manchurian Research Society (1926). Fig. 2 Boris V. Skvortzov. From the Review of the Manchurian Research Society (1926). Fig. 3 Skvortzov partially hidden by a giant water lily. From the Review of the Manchurian Research Society (1928). A further photograph is included in Naturalist Man'chshurii (Naturalist of Manchuria) ( 1 936:64). The caption reads 'Anniversary day of association A.N.E. - July 26 1936. The members of the association and their friends.' A SUMMARY OF THE DIATOM MATERIAL SKVORTZOV STUDIED The majority of Skvortzov's material came from the area known as Manchuria which has been largely understood as north-east China. Manchuria can be separated from the Russian Federation largely by the Amur, Argun, and Ussuri rivers, from North Korea by the Yalu and Tumen rivers, and from Mongolia by the Da Hinggan (Great Khingan) Mountains. Manchuria also includes the Liaodong penin- sula. Provincial divisions have changed frequently and since 1956 Manchuria has comprised Jilin, Heilongjiang, and Liaoning. Much of the region is mountainous. The Da and Xiao Hinggan (Great and Lesser Khingan) in the north and the Changbai in the east are the greatest ranges. Material has been grouped to largely correspond to modern geographical areas. Each entry below is identified by the relevant published reports and a collecting date if given. Future work will concentrate on the new taxa Skvortzov named and the possible material that can be, or has been, used for typification. China Fujian (Fukien) FuZHOU (FOOCHOW, MINHOW). Skvortzov studied several collec- tions from Fuzhou most of which he collected himself in 1918 and 1919 (item 418:389); Chung (1929:125) gives a brief account of Skvortzov's visit. 1917 (item 61): The first article was based on material collected by Mrs W.R. Myers 'in the environs of Foochow. . .the material was examined in the Petrograd Academy of Science in 1917' (item 61:205). Skvortzov examined the material in the Petrograd Acad- emy suggesting that this study may have been under the supervision of Kolbe and Wislouch, leaving open the possibility that this material may still be in St. Petersburg (Petrograd). 1918 (item 139): This was the first account of material collected by Skvortzov during his trip to China: 'Being in Foochow in 1918 some observations were made by me during the winter time on the life of the plankton of the fishponds' (item 139:190). 1918 (item 259): Material collected by Skvortzov 'on the sea- shore of Fukien Province, near Foochow, China, during the winter of 1918' (item 259: 151). 92 D.M. WILLIAMS & G. REID 1918 (item 427): This material is presumably part of the collec- tion made by Skvortzov in 1918. One collection was noted as the type material for Porosularia meisteri Skvortzov and Porosularia merrilli Skvortzov, ('Hab. in Stagno prope oppidum Foochow, prov. Fukien, China australis, Ig. B. Skvortzov, in 1918', item 427:413) and another for the type material of Porosularia borgei Skvortzov ('Hab. in orizetis prope oppidum Foochow, prov. Fukien, China australis, Ig. B. Skvortzov, 1918', item 427:415). February 1919 (item 140): Material collected by 'Mr. C.R. Kellogg, ... at the end of February 1919 in Kokchiang, 70 miles from Foochow...' (item 140:195). No date (item 427): Some additional samples were included in this report, the collections probably also dating to 1918: 'River Ming, in the environs of Foochow City, prov. Fukien, South China' (type of Pinnularia kisselewii var. subacuta Skvortzov) (item 427:400) and '...swamps of Foochow, prov. Fukien, South China' (type material for Pinnularia meyerii var. hinganica Skvortzov) (item 427:400). XIAMEN (AMOY) 1924 (items 222, 386, see also item 194 & 427): Material collected by Dr H.H. Chung in the winter of 1924. More details of the locality were given in a later paper by Skvortzov: '...Hab. In stagnis prope oppidum Amoy, prov. Fukien, China australis, Leg. H.H. Chung, a. 1924' (item 386:378, item 427: 412 & 414). This material served as the type for Porosularia amoyensis Skvortzov and Porosularia kolbei Skvortzov (item 427:412 & 414). TIANJIN (TIENTSIN). Collections made by Skvortov in early 1919 were used for two studies (detailed below). His general botanical work from this period is covered in a series of papers entitled 'Notes on the agriculture botany and zoology of China' (items 59-90, 105- 110, 125-132, 138-140). 17 March 1919 (item 138, 190): 'Being in Tientsin in March 17, 1919 a small collection of Algae was gathered by me in the ponds of the Russian Garden and near brick-kilns' (item 138:189; item 190:102). KULIANG RIVER 1918 (item 228): Material collected by Skvortzov 'at Kuliang, Fukien Province, in South China. This place is 9 kilometers from Foochow, and is 2,400 feet above sea level' (item 228:39). Jiangsu (Kiangsu) SUZHOU (SOOCHOW). Several samples from Suzhou were noted by Skvortzov in item 330, collected by Dr H.L. Li in 1934. Skvortzov later (item 402) gave descriptions of the same taxa, noting the troubled publication history of this study: '...this manuscript was published by the author in Proceedings of the Harbin Nat. Hist Sc. in 1946...' (item 402:59). The 1946 paper (item 3 30) therefore still has priority in spite of it being hard to obtain. 1934 (item 330 & 402): 'China media prope Soochow prov. Kianghsi Dr. H.L. Li, 9/8 34'. Skvortzov described several new taxa from this locality. One locality is given with the date '9/9, 34'. We suspect that the month of collecting in erroneous in one or other of the reports as Skvortzov later states that the new taxa were based on one sample provided by Dr L. Li 'in the lake in the City of Soochow of prov. Kiansu...in 1934' (item 402:59). POYANG October 1929 (item 287): Material from Rev. Umberto Verdini, 'from the Eastern Lake of the city of Poyang, Hunan, China, on lotus leaves' (item 287:465). While the title refers to Hunan, Chin (1951:156) points out that Poyang Lake is in the northern part of Kiangsu and not in Hunan. The material is referred to again in item 405. Zhejiang (Chekiang) HANGZHOU (HANGCHOW) 12 August 1933 (item 297): Material collected from '. . .a sample of mosses...' by Mr I. Kovalchook-Koval 'in a cave on a rock in the environs of Hangchow . . . ' (item 297:2 1 9). Kovalchook-Koval made a number of collections and contributed to the many Manchurian Research Society publications. Liaoning DALIAN (DAIREN) 1926 (item 226): Material from 'my young friend A. Prosowetsky a small collection of sea mud, from oysters collected at Dairen' (item 226:419). Shandong (Shantung) SHANWANG - MIOCENE FOSSIL 1935 (item 308, 330): Skvortzov reported that this material was 'sent to me some time ago by Drs. C.C. Young and P. Teilhard de Chardin of the Geological Survey of China' (item 308:193). He described the material thus: 'diatomaceous earth forms a thick formation at Shanwang near the city of Linchii, eastern Shantung, about half way between Tsinan and Tsingtao, not far south from the railway' (item 308:193). According to Skvortzov, this was the first Chinese freshwater diatom fossil deposit. In an earlier report, Young (1936) provides more details of the deposit and when it was col- lected ('A closer examination of the region was evidently necessary, and I carried it out in May 1935', Young, 1936:172). Young adds in a footnote: 'The papyrus shales are a diatomaceous earth, the botanical study of which has been undertaken by Dr. B. Skvortzov. Most abundant are the frustules of Melosira granulata (Her.) Ralfs. A paper by Dr. Skvortzov will be published in the Bulletin' (Young, 1936:175, footnote 1). As for the age of the material, 'After careful examination, Hu believes that the age of the flora is probably Miocene...' (Young, 1936:177) and '...we have to consider the Shanwang series as Upper Miocene in age' (Young, 1937:280). Material we believe to be from the same source was also studied by Voigt (1937). He says of this material, '...a certain quantity was kindly placed at the disposal of the writer by Dr. W.H. Wong of the National Geological Survey of China...' (Voigt, 1937:311). Voigt also states that 'So far only one authentic deposit of diatomeaceous earth has been reported in China, namely, that from Shan-wang in the Lin-chu district of Shantung, which was discovered by Dr. C.C. Young of Peiping' (Voigt, 1937:311). Some of Voigt's material is housed in the BM and some preliminary electron micrographs have been published (see Williams et al., 1998:59, figure 2-5, electron micrographs of Tetracyclus emarginatus var. cf. parvula Forti, see also p. 55, footnote 8; BM s.n.). Further examples of this material are housed in the Palaeontological Museum of Stockholm. Item 330 contains repeat descriptions of two taxa first described as new in item 308. Sichuan (Szechwan) CHENGDU (CH'ENG-TU) March 1926 (item 317): '...two tubes of diatom material from the environs of that city [Ch'eng-tu], collected in March, 1926' (item 317:479). The material was donated by Dr H.D. Brown of West China Union University, Chengdu, Sichuan. Some further commen- tary and notes are in item 405. BORIS V. SKVORTZOV Heilongjiang (Heilungkiang, Heilungtsiang) VARIOUS LOCALITIES 1950 (item 426): 'Amur prope Oopu', part of A.I. Baranov collect- ing trips (type material for Pinnularia zabelini var. amurensis Skvortzov). 1951 (item 427): Material collected by Skvortzov, 'Hab. In lacu alkalini prope Stationem Sun, prov. Heilungtsiang, Manchuria borealis, China, Skvortzov, 1 5.9. 1 95 1' and used as the type material for Porosularia poroidea Skvortzov and Porosularia subsalsa Skvortzov. No date (item 405): 'In regionis montanis, Prov. Heilungtsiang, Manchuria borealis, China'. HARBIN 1915-1917 (item 160, 171): Skvortzov lists six samples in item 160 and eight samples in item 171. The samples were collected by various people including Skvortzov: B.R. Arnold, S.W. Schernich, M.A. Hintze, N.A. Schemilewitch and M.G. Dorian. Of these collections, the information given is, in most some cases, of little significance in establishing exact locations and dates of collecting. For instance Skvortzov's own collecting is given as '...from the environs of Harbin collected by me...' (our translation) between 1915 and 1917. The most precise dates and details are given for B.P. Arnold (during August 1916), N.A. Schemilewitch (July 1916) and M.G. Dorian (August 1916). 1927 (item 312): Material collected by Skvortzov. Three samples were examined: ( 1 ) from Harbin, 20 September 1 927, 'from the bark of Ulmus manshurica Nakai'; (2) from Eastern Harbin, near Maoershan Railway Station, 20 July 1927, 'on mosses on rocks along a mountain river'; (3) from Eastern Maoershan, near Mifun Station, 5 September 1927, 'in mosses on mountain rocks' (item 312:263). No date (item 66): Material described simply as The algae just described were studied from numerous collections gathered near the railway line in North and Middle Manchuria and mostly in the environs of Harbin...' (item 66:63). SONGHUA (SUNGARI) RlVER, HARBIN 17 March 1923 (item 211): Material collected from the Sungari River for a study on winter phytoplankton. The report lists ten species of diatom. 28 August 1935 (item 305): 'The material was collected 28 August 1935 by one of my enthusiastic collectors Mr. I. Kovalchook- Koval on a shore of a swampy lake on the Sungari river plain, three miles from the Sungari railway bridge' (item 305:783, including a photograph of the site, fig. a). A short account of the expedition is given in Anon. (1936). August 1936 (items 302-304): Plankton samples collected on the 9 August 1936 by Mr N.I. Nikitin, from a 'marshy branches], along sand-dunes' of the Sungari River (item 302:628). The diatoms are listed in part II of a three part study of this area (item 303). Skvortzov used the material for further taxonomic revisions in a later study (item 330): 'Hab. Manshuria bor., prope Cheng st. in stagna fl. Ashiho, M. Nikitin, 10/8 1936'. Beijing Municipality BEIJING (PEKING) March 1926 (item 219): Collections made by Prof. N.G. Gee and Dr H.J. Chu 'in the environs of Peking' (item 219:43). In all, three lakes were sampled. Previously, Gee published a list of diatoms from Suzhou and Ningbo (Gee, 1926). A name change is given in item 330:26 (Nitzschia regula f. pekinensis Skvortzov is replaced with N. regula var. robusta Skvortzov). 93 Shanghai June 1918 (item 70): Material collected by Skvortzov from a pond near the Public Gardens in Shanghai (item 70:66). 19 May 1933 (item 310): Material collected by Mr I. Kovalchuk- Koval 'a sample of mosses collected by him on the bark of a tree in Shanghai...' (item 310:443). Hongkong August 1957 (item 420): Collected by Mrs V.T. Mamchyi 'from mosses grown on the trunks of a tree in Hong-kong, eastern Asia' (item 420:407). Inner Mongolia HlNGGAN LING (KHINGAN MOUNTAINS) 1922 (item 213 and 330): Collections made by Skvortzov from the 'Khingan Mountains in the environs of the Fuleierdi Railway Station' (item 213:39). Further details given in item 330: 'Hab. Manshuria bor., in rivulos fl. Jal prope Barim, Khingan montibus'. 1951 (item 426 & 427): Material from the 'Northern part of Great Khingan Mountains from the western corner of Inner Mongolia, China. Collected during a botanical expedition made in August of 1951 by the Botanical Section of the N.E. China Agricultural Institute of Harbin' (item 426: 111). The material is composed of two samples of moss extracts (details are given in item 426:1 19). The samples were collected by Skvortzov who gives a full description of the collecting trip (item 426: 1 1 2). The paper makes reference to an unpublished manuscript (Bacillariophyta in Illustrated flora of North- Eastern China, 1957, 'deposited in Herbarium of Forest Academy in Harbin, China') which contained the first descriptions of the new taxa subsequently validated in this report. ARGUN RIVER, DALAI-NOR August 1926 (item 227): Twelve tubes of material collected by Mr. P. A. Pavlov '...in the northern part of Dalai-nor Lake in August, 1926'. (item 227:31). 1927 (item 314): Material collected by Mr PA. Pavlov during the summer of 1927 'in the environs of the Chalainor station of the Chinese Eastern Railway, at the source of Argun River, which arises from Dalai-nor or Talaihu Lake' (item 314:43). 1950 (item 427): 'River Argun, A.I. Baranov, 1950' as type material for Cymbella lanceolata var. grossepunctata Skvortzov. VARIOUS COLLECTIONS 1916, 1918, 1928, 1931 (item 289): This paper is divided into four parts. The first part deals with collections made from Bagah Oulan on the 18 September 1918 on Licent's travels to Kansou and Kou kou noor. Some further details of the expedition are given in Licent (1936:5 & 11). The second part deals with collections made from Yen tche in the province of Chansi in 1916, 'Chensi Central: vallee du Wei ho, les Alpes chinoises' (Licent, 1 936:5). The third part deals with collections made in the environs of Kalgan, near a lake. No specific details are given in item 289 but the material may have been collected during the expedition described by Licent (1936:7), of a visit in 193 1 to the Mongolian Interior, Kalgan. The fourth part deals with collections made in the North of Manchuria, in the province of Kirine, to the west of Harbin, on July 1928 (item 289:36; Licent, 1936:7). RIVER IMINGOL 1925 (item 209): Material collected by J.W. Tokmakoff 'in freundlichster und dankenswertester Weise Bacillariaceen-Material aus dem Fluss Imen-gol, 80 Kilometer von Hailar' (item 209:311). 94 D.M. WILLIAMS & G. REID Some further are details given in item 330: 'Mongolia occidentalis, fl. Imingol prope Hailar, J.V. Tokmakov, 1925'. Tibet 1901 (item 69): Material collected by Mr. Ladigin 'during the expedition in Tibet in 1 90 1 . . . ' The collections mentioned were from three lakes: (1) 'plankton... freshwater lake Kurlyk-nor [1 June 1901] in the Tsaidam district...' (2) '...salt lake Toso-nor [3 July 1901]... not far from lake Kukunor...' The samples were '...exam- ined in the Petrograd Academy of Science' (item 69:66). The list of species (containing only two diatoms) was from (3) Khara-nor. Russian Federation Jakoutsk 1912 (item 3): Material collected by Mr G. Dolenko ' . . .pendant une expedition, organisee par le Comite de migration' (item 3:19). No further details have been discovered. Primorsk LAKE TSHLA 5 August 1916 (item 5): Material collected by Lake Tshla 'se trouve pres de 1'embrouchure de 1'Amour et de la ville de Nikolaievsk' (item 5:20) ['close to the mouth of River Amur and the town of Nikolaievsk']. KHANKA (HANKA) LAKE, NEAR VLADIVOSTOK 1924 (5, 16, & 24 July; 4, 11, 15, 20 & 23 September; 15 August) (item 234): Material collected by E.N. Klobukova-Alisova in 1925 and by E.N. Klobukova-Alisova and A.G. Hahina in 1 924 during the South Khanka Botanical Expedition made by the Southern branch of the Geographical Society of Nikolsk-Ussurisk. Seventeen samples are listed with collection numbers. Some further notes are made in item 330. VLADIVOSTOK 1928 (item 258): Material collected by Skvortzov during the sum- mer from 'Golden Horn Bay and from Cape Basargino near the Pacific Fishery Research Station' (item 259:129) on a visit to Vladivostok. No date (item 220): Material collected from Vladivostok ' . . .from sea-weed Laminaria sp.' (item 220:57). No date (item 315): Material received from Mr I.P.Popov, 'a sample of diatoms collected by him during a botanical survey of Primorsk Province in a Carex-Sphagnum peaty bog in Lianchiho River Valley, about 40 kilometres east of Vladivostok, not far from the seashore' (item 315:161). Amursk KHABAROVSOK (HABAROVSK) 1909-1916 (item 40, 43, 66 & 254): Samples collected from the River Amur 'pendant les expeditions de 1909 a 1916, en Extreme Orient, entreprises par le Department d' Agriculture' (item 40:21). In two later reports, Skvortzov adds that the samples were col- lected by W.K. Soldatow 'during the expedition of the Department of Agriculture in 1910 to 1914. ..largely plankton obtained from the Amur river, near Habarosvk' (item 254:69) and 'also from collections of the Amur river made by W.K. Soldatow...' (item 66:63). RIVER AMUR 1928 (item 311): Material collected by Skvortzov during the summer 'in the environs of Okeanskaia station, near Amur Bay, in a forest Hypnum bog' (item 3 1 1 :25 1 ). 1951 (item 386): Skvortzov described several new taxa from Amur River material: 'Hab. In fl. Amour, Siberia, Leg. A.I. Baranov a. 1951' [item 386:375]. RIVER ZEYA (ZEJA) 1909-1910 (item 6): Material collected by Mr J. Abramov 'du fleuve Zeja (province Amourienne). . .dans les environs de la station meterologique de Bomnak...' (item 6:128). 1950 (item 427): Material described by Baranov, 'fl. Zea. Districtus Amurensis, Siberia orientalis, Ig. A. Baranov, 1950' and used by Skvortzov to described new taxa from Amur River' (type of Pinnularia zabelini Skvortzov and var. zeaana Skvortzov, item 427:414 type of Porosularia pseudoviridis Skvortzov). Siberia LAKE BAIKAL 1916, 1925-1926 (item 210): The material came from several expeditions made by C.I. Meyer. Skvortzov listed 36 samples, nine from 1916,11 from 1 925 and 1 6 from 1 926. Meyer ( 1 930) presented his own detailed account of the algae from the samples he collected during these expeditions, as well as presenting his accounts of the samples Skvortzov studied. Meyer's descriptions of the samples are somewhat more detailed than Skvortzov's account. There has been some debate as to whom was actually responsi- ble for the new taxon names in the 1928 publication (item 210). Much later Skabichevskij (1974) established that Skvortzov alone was responsible, as Meyer (1930) presented his own account a few years later in which not only did he differ from Skvortzov in a number of places but also he noted that he was solely responsi- ble for collecting the material (Meyer, 1930:327). A similar statement was made in Skvortzov and Meyer: 'Since 1926 B.W. Skvortzov has joined prof, [sic] C.I. Meyer to work together on the identification and classification of the Diatoms collected by the latter in the Baikal lake and this work is but a preliminary report of the work' (item 210:2, italics added). Hence it seems justified to consider the taxonomic work as Skvortzov's alone. The correct attribution for the new taxa in item 210 should there- fore be 'Skvortzov in Skvortzov & Meyer' (see also Kociolek & Stoermer 1988:96-97). 29 July 1916 (item 307): Nearly ten years after the first paper on Lake Baikal, Skvortzov re-examined in detail one sample at '33 meters near the Ohlon Gate of Baikal Lake...' (item 307:297). This sample is also listed in Skvortzov & Meyer (item 210:2; '7. 29/VII 'Olhinskie vorota'. Haringari bay, at a depth of 33 metres'). A number of taxonomic changes are made in item 330. Some of Meyer's Lake Baikal material is available in the Califor- nia Academy of Science and Bremerhaven (Kociolek & Stoermer 1988:95-96). New collections from Lake Baikal (housed at BM), made as part of the DI (Flower & Williams, 1999), will be ideal for designating neotypes as many of Meyer's collecting sites were revisited and are subsequently being re-studied (Reid & Williams, submitted). No date (item 426 & 427): Some additional material from Lake Baikal was noted in this later report: 'River Kitschera near Lake Baikal, Siberia' (type of Pinnularia kisselewi var. attenuata Skvortzov), 'Lake Baikal' (type of Pinnularia dorogostaiskii var. latior Skvortzov) (both in item 427) and 'River Selenga prope Baikal, Siberia' (type of Pinnularia lata var. intermedia Skvortzov) (item 426). BORIS V. SKVORTZOV 95 KENON LAKE, TRANSBAIKALIA No date (item 313): Material collected by Miss K.V. Okunozova, '. . .near the shore of the lake, from twigs and leaves of Potamogeton sp.' (item 3 13:399). RIVER KHOLOY (CHOLOY) No date (item 43): Material obtained from ' 1' Amour par 1' expedition de Mr. Korotky ...dans le fleuve Choloy en Transbaikalie par 1'expedition de Mr. Korotky...'. (item 43:22). Tyva RIVER YENISSEI 1925-1926 (item 386, 403): Material collected by P.J. Usachow and described as '...Hab. In fl. lenissei prope oppidum Krasnojarsk, Siberia, Leg. P.J. Usachow, 1925-1926' (item 386:376). Many years after receiving the material Skvortzov noted: 'The author received the diatom samples from Yenisei River in 1927 while he was working in Harbin, China from the Hydrobiological station of Krasnojarsk City, Siberia. A series of slides was prepared by the author from these collections and preliminary studies made in 1930- 1931' (item 403:57). The material was used for some taxon descriptions in item 399, which provides more precise dates. The collections include material for Pinnularia viridis var. tubensis Skvortzov, P. viridis var. minuta Skvortzov, ('lenissei prope oppi- dum Krasnojarsk, Col. 17, 11. '26') and P. viridis var. tumida Skvortzov ('In fl. Jenisei, Siberia media, Ig. 26.6. '26'.) The Altai Mountains The Altai Mountains in southern Siberia form the major mountain range in the western Siberia region and provides the sources of the rivers Ob and Irtysh. The main areas of the region are Altaysky, Zapovednik and an area surrounding Teletskoye Lake. The entire area is enormous and Skvortzov's samples are limited especially as most of his reports lack useful details. 1897 (item 41): Material collected by Mr Silantiev 'dans les sources "Rachmanovsky" de 1' Altai.' (item 41:21). Kaolingtze 15 July 1926 (item 316): Material collected by Skvortzov, on 15 July 1926 'in the mountain ranges near Kaolingtze station of the Chinese Eastern Railway, about 300 miles east from Harbin, I collected a sample of diatoms from a little mountain bog of forest mosses' (item 316:343). The material was probably used in item 427 as the type material for Pinnularia fritschiana Skvortzov. Kazakhstan Akmolinsk KURGAL'DZIN LAKE. These early collections were probably stud- ied in St. Petersburg and may have been retained in that institute. June 1899 (item 42): Samples of Chaetoceras wighamii Brightwell from 'le bassin du lac Kourgaldjin de la province d' Amolinsk.' As noted in item 252, 'The presence in the Kokai Lake and Nura River of Chaetoceras Wighamii Brightwell... is interest- ing' (item 42:33). The collecting date for Nura River is given as 8 June 1899 and for Kokai Lake is given as 23 June 1899. June 1912 (item 7): Collected by Mr Ganeshinn (Ganesin) 'provenant du lac Courgaldgin' (item 7:128). VARIOUS LAKES INCLUDING TELETSKOI (TELETSKOE) AND KURGAL'DZIN 1899, 1901 (item 252, 246): Material provided by Prof. G.I. Vereshagin, collected by PT. Ignatow 'in lakes of the Akmolinsk district of Siberia in 1899 and 1901' (item 252:33). There were 20 samples, all listed. Item 246 is a short report on a sample from Lake Teletskoi taken in 1901 by Ignatow. BLACKHACH (BALKACHE) LAKE No date (item 42): Material collected from 'le lac salin d Ala-Koul, situe dans le partie ouest du lac Balkache' (item 42:22). ZAYSAN (ZAISAN) LAKE 1905 (items 193, 203): Material collected by A.N. Sedelnikoff (Sedelnikov) 'in lakes in the Altai Mountains of South Siberia. All the samples were gathered in the environs of Zaisan Lake...' (item 193:249). Further comments on the material can be found in item 203. There is some overlap of material studied in these accounts but they are largely complementary suggesting they are all from the same source. Skvortzov refers to a detailed account of Zaysan Lake written by Sedelnikoff ( 1 905), a book we have been unable to trace. MARKAKOL' (MARKA-KUL) LAKE 8 July 1912 (item 8): Material collected by Mr A.N. Sedelnikov 'dans le grand lac Marka-Kul...' (item 8:128). 1914, 1916 (item 253): Material collected by Sedelnikov from 'several mountain lakes of the Altai district in the summers of 1914 and 1916' (item 253:36). South Korea SEIKO LAKE 1917 (item 223): Material collected from 'leaves and stems ofTrapa natans received from Chosen (Korea)...' from 'Seiko lake near Suigen,II, 1917' and received by Skvortzov from the Director of the Agricultural Experimental Station, in Suigen, Keiki-Do (item 223:9). Some further taxonomic revisions are undertaken in item 406. SEIRIORI LAKE March-July 1926 (item 225): Collections made by Prof. Tamezo Mori 'in the lake at Seiriori near Seoul between March and July, 1926' (item 225:283). Some further taxonomic revisions are under- taken in item 330 and 406. KOREAN STRAIT 1925-26 (item 256): Five samples obtained from Dr Yojiro Wakiya. VARIOUS FOSSIL MATERIAL 1927 (item 292): Material received from Sigetaro Kawasaki, la- belled 'diatom earth from the younger Tertiary, Bunzan-Men, Ampen-Gun, South Kankyo-Do' (item 292:9). Japan Honshu Island KAMAGAIKE 2 October 1930 (item 318): 'Diatoms collected by Mr. Yoshikazu Okada in a mountain bog from Kamagaike, Mt. Kirigamine, Prov. Sinaro, Central Nippon, October 2. 1930' (item 318:53). KIZAKI LAKE July 1927 (item 293): Material collected by Mr K. Kiuchi (donated by Prof. T. Kawamura) from Kizaki Lake, Shinano Province. 'I have examined a hundred microscopic slides... Half a year was spent in the study of this collection' (item 293:9). Some further details are given in items 330, 404 and 405 along with some taxonomic revisions. BIWA LAKE No date (item 294): One tube of diatom clay from Biwa Lake, north 96 of Osaka. Sent to Skvortzov by Prof. Tamiji Kawamura. Further details are given in items 330, 404 and 405 along with some taxonomic revisions. KANAZAWA OYSTER EXPERIMENTAL STATION December 1927 (item 257): Material from Dr Juzo Hori (Kanazawa Oyster Experimental Station). Kyushu Island IKEDA LAKE January 1923 (item 306): Material from 'Ikeda Lake, Satsuma Province, Kiusiu Island, from the southern part of Nippon, collected by Dr. T. Kawamura in January, 1923' (item 306:191). Further details are given in items 330, 404 and 405 along with some taxonomic revisions. Sea of Japan 1921-1925 (item 273): A series of collections made by the Imperial Fisheries Institute of Tokyo, Japan from the Sea of Japan and donated to Skvortzov by Dr K. Okamura. The collections were made on 10 July 1921, 14 November 1923, 24 November 1925 and 21 December 1925. Wamura, Nagano Prefecture (Honshu) (item 298) and Saga Prefecture (Kyushu) (item 299) VARIOUS FOSSIL MATERIAL. Skvortzov examined several sam- ples of Neogene diatoms received from Prof. Tamij Kawamura. Item 406 is a summary article with some taxonomic revisions. Ueyama & Kobayashi (1983) have studied material from Wamura and re- assessed Skvortzov 's work. North America January 1936 (item 295): Material from 'the Field Museum of Natural History at Chicago. The material was obtained by filtering Lake Michigan water from the city mains of Chicago in January, 1936... Type material of new forms is in the author's personal collection' (item 295:652). Australia (Fossil) Item 301: Material from New South Wales '...situated five miles from Cooma, and one and a half miles from Bunyan Platform...' sent by Mr F.S. Mance of the New South Wales Department of Mines, Sydney (item 301:175). New Zealand Item 319: Material from a series of collections sent over a period of years. The notes suggest 12 different localities. Item 320: Material received from Mr Ian C. Edmundson, The sample consists of greenish masses of immense growth of Scenedesmus brasliensis Bohlin' (item 320:41 1), collected from an aquarium. Philippines 29 January 1936 (item 309): Sample sent by Dr Eduardo Quisumbing (curator of the Phillippine National Herbarium, Ma- nila), 'based on a sample collected January 29, 1936, from filter No. 8 in Balara, Rizal Province' (item 309:287). D.M. WILLIAMS & G. REID Africa 1936 (item 330): Material is noted only in the review article (item 330), both localities collected by F. Pringle: 'Africa australis, in stratum lacustre prope Franzenkop et Pieska, Cape prov.' and 'Hab. Africa australis, prope Port Elizabeth, epiphytice in lichenes F. Pringle, 27/5 1936.' Cuba 1936 (item 330): Material noted only in the review article (item 330). Both localities collected by Rev. B. Robert: 'Hab. America australis, Cuba, Habana, prope Vedado, epipytice apud radices' and 'Hab. America australis, Cuba, Habana, prope Vedado, epipytice in truncos arbores. Rev. B. Robert, 40/6 1936'. Afghanistan 1916 (item 4): Material collected by Mr V. Lagatov 'dans les canaux de Mourgab pres de la frontiere d' Afghanistan et de Perse dans le desert sablonneux de Kara-Koum' (item 4:19). Sri Lanka 1926 (item 235): Material from several collections made by A.H.G. Alston. Much of Alston's material is in the BM. A search, so far, has revealed nothing of direct relevance. 1926 (item 265): Material collected by A.H.G. Alston, 'near the sea shore at Gintota from the twigs of Ceramium clavulatuni 1 on the 16 August 1926. No date (items 330 & 361): Material donated by Dr N.G. Ball 'in a swamp in mountain region'. India February 1926 (item 288): Two samples from Prof. S.R. Bose; one from mosses and algae from tree bark (15 Feburary 1926), the other from mud in a channel. B.V. SKVORTZOV'S BIBLIOGRAPHY As far as we are aware this is a complete listing of Skvortzov's published work. It is probable that some other publications may be in inaccessible natural history journals and magazines printed in Manchuria. The authorship is Skvortzov unless otherwise stated. An asterisk (*) denotes a publication that includes diatom species. We have not seen the last six entries in this list and presume they were published posthumously. 1. Skvort/ov, B.V. 1916. Les algues de la Mandjourie et recherches sur la vegetation aquatique dans la vallee de Soungari. Zhurnal Mikrobiologii 3 (3_4); 443. 2. Skvortzov, B.V. 1917. Uber Flagellata aus Mandschurei. Part 1 . Zhurnal Mikrobiologii 4 (1-2): 57-78. (French summary p. 287). 3. * 1917. Materialy po flory vodoroslej Aziatskoj Rossii. I. Vodorosli iz' Akutskoj oblasti. [Contributions a la flore des algues de la Russie d'Asie. I. Algues de la province de Jakoutsk.] Zhurnal Russkogo Botanlcheskogo Obshchestva pri Akademii Nauk 2: 10-12. (French summary, p. 19). 4. * 1917. Materialy po flory vodoroslej Aziatskoj Rossii. II. Vodorosli iz' Zakaspijskoj oblasti. [Contributions a la flore des algues de la Russie d'Asie. II. Algues de la province Transcaspienne.] Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 2: 13-15. (French summary, p. 19). 5. * 1917. Materialy po flory vodoroslej Aziatskoj Rossii. III. O fitoplanktony oz. Chla Priamurskoj oblasti. [Contributions a la flore des BORIS V. SKVORTZOV 97 algues de la Russie d'Asie. III. Le phytoplancton du lac Tshla de la province Priamourskaya.] Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 2: 15-19. (French summary, p. 20). 6. * 1917. Materialy po flory vodoroslej Aziatskoj Rossii. IV. Vodorosli verhov'ev" raki zei Amurskoj oblasti. [Contributions a la flore des algues de la Russie d'Asie. IV. Algues des sources du fleuve Zeja (Province Amourienne)]. Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 2: 1 17-120. (French summary, p. 128). 7. * 1917. Materialy po flory vodoroslej Aziatskoj Rossii. V. Vodorosli iz" Akmolinskoj oblasti. [Contributions a la flore des algues de la Russie d'Asie. V. Algues de la province d'Akmolinsk.] Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 2: 120-125. (French summary, p. 128). 8. * 1917. Materialy po flory vodoroslej Aziatskoj Rossii. VI. O fitoplanktony oz. Marka-kul' Kirgizskago kraa. [Contributions a la flore des algues de la Russie d'Asie. VI. Le phytoplancton du lac Marka-Kul]. Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 2: 125-127. (French summary, p. 128). 9. 1918. On timber industry in the Far East. Sel'skoe Khozial'stvo v Sievemoi Man'chzhurii 5-6: 19-20. (In Russian). 10. 1918. Are there truffles in Manchuria? Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 5-6: 20. (In Russian). 11. 1918. Lespedeza bicolour. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 5-6: 20-21. (In Russian). 1 2. 1 9 1 8. On little known vegetable tuber plants of Manchuria. Set 'skoe Khozial'stvo v Sievernoi Man'chzhurii 5-6: 26. (In Russian). 13. 1918. The duckweed as food product for poultry. Sel'skoe Khozial'stvo v Sievernoi' Man'chzhurii 56: 2627. (In Russian). 14. 1918. Plagiospermum chinensis as a fruit shrub. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 5-6: 9-10. (In Russian). 15. 1 9 1 8. On some wild Manchurian plants, used by Chinese and natives as vegetables. Sel 'skoe Khozial'stvo v Sievernoi Man 'chzhurii 7-8: 24-26. (In Russian). 1 6. 1 9 1 8. On little known vegetable tuber-plants of Manchuria. Sel 'skoe Khozial'stvo v Sievernoi Man'chzhurii 7-8: 26. (In Russian). 17. 1918. The duckweed as food product for poultry. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 78: 2627. (In Russian). 18. 1918. Amaranth as a field plant. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 7-8: 27-28. (In Russian). 19. 1918. The Amurian velvet-tree in Chinese medicine. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 7-8: 28-29. (In Russian). 20. 1918. Chinese hemp as a technical plant. Sel'skoe Khozial'stvo v Sievemoi Man'chzhurii 7-8: 30-31. (In Russian). 2 1 . 1918. The dye plant in Manchuria. Sel 'skoe Khozial'stvo v Sievernoi Man'chzhurii 7-8: 31-34. (In Russian). 22. 1918. The use of different timber trees of Manchuria forest. Sel'skoe Khozial'stvo v Sievernoi Man 'chzhurii 7-8: 34-35. (In Russian). 23. 1918. Cultivation of Daurian ivy. Sel 'skoe Khozial'stvo v Sievemoi Man'chzhurii 7-8: 35-36. (In Russian). 24. 1918. On wild vegetables of the Far East. Sel'skoe Khozial'stvo v Sievemoi Man'chzhurii 9-10: 25-28. (In Russian). 25. 19 18. On garden curcubitaceous plants of Manchuria and Primorsk Province of Russia. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 9- 10: 28-32. (In Russian). 26. 1918. Cultivation of the hawthorn in Northern China. Sel'skoe Khozial'stvo v Sievemoi Man'chzhurii 9-10: 32. (In Russian). 27. 1918. Rice cultivation in the Middle and in the South Manchuria. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 9-10: 33. (In Russian). 28. 1918. Manchurian Actinidias as fruit plants. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 9-10: 33-34. (In Russian). 29. 1918. Chemical analysis of Manchurian soy-beans. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 9-10: 35-36. (In Russian). 30. 1918. The trade important fishes of Manchuria and Amur District. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 11-12: 18-23. (In Russian). 3 1 . 1918. Melliferous plants of Manchurian forest. Sel 'skoe Khozial'stvo v Sievernoi Man'chzhurii 11-12: 26-27. (In Russian). 32. 1918. The tanning plants of Manchurian forest. Sel 'skoe Khozial'stvo v Sievemoi Man'chzhurii 11-12: 34-36. (In Russian). 33. 1918. Chinese nutgalls. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 11-12: 36. (In Russian). 34. 1918. The nuts of Manchurian walnut and their utilization. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 11-12: 37-38. (In Russian). 35. 1918. The further note on cultivated cucumbriaceous plants of Manchuria. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 11-12: 38. (In Russian). 36. 1918. Manchurian soybeans. Vestnik Man'chzhurii (Manchurian Monitor 2 ) 153. (In Russian). 37. 1918. The fruit-gardening in Far East. Vestnik Man 'chzhurii (Man- churian Monitor) 209. (In Russian). 38. 1918. The Chinese medicine trade in the Far East. Vestnik Man 'chzhurii (Manchurian Monitor) 212. (In Russian). 39. 1918. The berry-gardening in Manchuria. Vestnik Man'chzhurii (Manchurian Monitor) 217. (In Russian). 40. * 1918. Materialy po flory vodoroslej Aziatskoj Rossii. VII. Perbya svydynia o fitoplanktony r. Amura. [Contributions a la flore des algues de la Russie d'Asie. VII. Notions preliminaires sur le phytoplancton de d' Amour. ] Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 3: 1-9. (French summary, p. 21). 41. * 1918. Materialy po flory vodoroslej Aziatskoj Rossii. VIII. Vodorsli iz" Altaa. [Contributions a la flore des algues de la Russie d'Asie. VIII. Les algues de 1' Altai]. Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 3: 10-12. (French summary, p. 21). 42. * 1918. Materialy po flory vodoroslej Aziatskoj Rossii. IX. O Chaetoceras iz" Zapadnoj Sibiri. [Contributions a la flore des algues de la Russie d'Asie. IX. Sur les Chaetoceras maritimes de la Siberie d'ouest]. Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 3: 12-18. (French summary, p. 22). 43. * 1918. Materialy po flory vodoroslej Aziatskoj Rossii. X. Ky poznaniu vodoroslej Amurskoj i Zabajkal'skoj oblastej. [Contributions a la flore des algues de la Russie d'Asie. X. Sur quelques algues des province de 1'Amour et de la Transbaikalie]. Zhurnal Russkogo Botanicheskogo Obshchestva pri Akademii Nauk 3: 18-20. (French summary, pp. 22-23). 44. 1919.ThewaterplantMrya/e/eroxSalisb.asafoodplantofChinese. Sel'skoe Khozial'stvo v Sievernoi' Man 'chzhurii 12: 22. (In Russian). 45. 19 19. The edible honey-suckle of Manchuria. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 1-2: 23-25. (In Russian). 46. 1919. On some Manchurian plants used by Chinese for various handworks. Sel'skoe Khozial'stvo v Sievernoi' Man'chzhurii 1-2: 23-25. (In Russian). 47. 1919.ThegardenbeansofManchuriaandNorthernChina.Se/ifc0e Khozial'stvo v Sievernoi Man'chzhurii 1-2: 26-28. (In Russian). 48. 1919.ThegardenbeansofManchuriaandNorthernChina.5e/ifcoe Khozial'stvo v Sievernoi Man'chzhurii 3-4: 10-13. (In Russian). 49. 1919. The varieties of kaoliang and barley of Manchuria. Sel 'skoe Khozial'stvo v Sievernoi Man'chzhurii 3-4: 13. (In Russian). 50. 1919. The utilization of horse-tail (Equisetum) in China. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 3-4: 13-14. (In Russian). 5 1 . 1919. Freshwater algae used as food in China. Sel 'skoe Khozial'stvo v Sievernoi Man 'chzhurii 3-4: 14-15. (In Russian). 52. 1919. The awakening of the vegetation at Harbin in 1918. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 5-6: 27. (In Russian). 53. 1919. On some plants cultivated in South Manchuria. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 5-6: 28. (In Russian). 54. 1919. Apiculture and wax trade in China. Sel'skoe Khozial'stvo v Sievernoi Man'chzhurii 5-6: 29-31. (In Russian). 55. 1919. The results of cultivation of some Chinese plants at Harbin in 1919. Sel'skoe Khozial'stvo v Sievernoi Man 'chzhurii 5-6: 34-36. (In Russian). -We have been unable to examine any issues of the Manchurian Monitor. A note in one of the Research Society's reports says: 'An arrangement was concluded with the "Manchurian Monitor", published by the Chinese Eastern Railway, whereby the Department devoted to Research provided space on its pages for any literary labour performed by members belonging to the Manchuria Research Society' (Review of the Manchurian Research Society, March, 1928:9). 98 D.M. WILLIAMS & G. REID 56. & Shkurkin P.W. 1919. A brief Russian-Chinese dictionary of agricultural plants and vegetable products of Manchuria. Sel'skoe Khoziaistvo v Sieve rnoi Man 'chzhurii 5-6: 48-53. (In Russian). 57. 1919. Observations on melliferous plants in Northern Manchuria in 1919. Sel 'skoe Khoziaistvo v SievernoY Man 'chzhurii 10-12: 1 5- 1 6. (In Russian). 58. 1919. On some varieties of horn-cherries (Plagiospermum) in Northern Manchuria. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 10-12: 28-29. (In Russian). 59. 1919. Notes on the agriculture botany and zoology of China. I. Dye-plants and dye-stuffs of Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 49-52. (See item 21). 60. 1919. Notes on the agriculture botany and zoology of China. II. On the fossil animals of North Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 52-53. 61. * 1919. Notes on the agriculture botany and zoology of China. III. The fresh-water algae from the ponds of South China. Journal of the North China Branch of the Royal Asiatic Society 50: 53-56. 62. 1919. Notes on the agriculture botany and zoology of China. IV. On the study of the Manchurian wheat. Journal of the North China Branch of the Royal Asiatic Society 50: 56-57. 63. 1919. Notes on the agriculture botany and zoology of China. V. On the beetles and butterflies of the Far East. Journal of the North China Branch of the Royal Asiatic Society 50: 57-60. 64. 1919. Notes on the agriculture botany and zoology of China. VI. Observations on the river soft tortoise in North Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 60-61. 65. 1919. Notes on the agriculture botany and zoology of China. VII. On the fresh water shrimps in Manchuria and China. Journal of the North China Branch of the Royal Asiatic Society 50: 6 1 . 66. * 1919. Notes on the agriculture botany and zoology of China. VIII. On the exploration of the fresh water algae in Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 62-63. 67. 1919. Notes on the agriculture botany and zoology of China. IX. On some Chinese medical plants of the Far East. Journal of the North China Branch of the Royal Asiatic Society 50: 63-64. 68. 1919. Notes on the agriculture botany and zoology of China. X. Some observations on the growth of weeds and algae in rice fields at Foochow. Journal of the North China Branch of the Royal Asiatic Society 50: 64-66. 69. * 1919. Notes on the agriculture botany and zoology of China. XI. The phytoplankton of some Tibetan lakes. Journal of the North China Branch of the Royal Asiatic Society 50: 66. 70. * 1919. Notes on the agriculture botany and zoology of China. XII. On some freshwater algae collected in Shanghai. Journal of the North China Branch of the Royal Asiatic Society 50: 66-67. 7 1 . 1919. Notes on the agriculture botany and zoology of China. XIII. The use of Nostoc as food in N. China. Journal of the North China Branch of the Royal Asiatic Society 50: 67. 72. 1919. Notes on the agriculture botany and zoology of China. XIV. The bibliography of the algae of China and neighbouring countries. Journal of the North China Branch of the Royal Asiatic Society 50: 67- 69. 73. 1919. Notes on the agriculture botany and zoology of China. XV. On the study of the wild vegetables of Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 69-73. 74. 1919. Notes on the agriculture botany and zoology of China. XVI. Dimensions of trees in the Manchurian forests. Journal of the North China Branch of the Royal Asiatic Society 50: 73-74. 75. 1919. Notes on the agriculture botany and zoology of China. XVII. The origin of the Manchurian fishes. Journal of the North China Branch of the Royal Asiatic Society 50: 74-78. 76. 1919. Notes on the agriculture botany and zoology of China. XVIII. The little known and new oil plants in Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 78-79. 77. 1919. Notes on the agriculture botany and zoology of China. XIX. The principal tanning plants of Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 79-80. 78. 1919. Notes on the agriculture botany and zoology of China. XX. On the cultivation of water plants in China. Journal of the North China Branch of the Royal Asiatic Society 50: 80-81. 79. 1919. Notes on the agriculture botany and zoology of China. XXI. On the growth of panic grass in China. Journal of the North China Branch of the Royal Asiatic Society 50: 8 1 . 80. 1919. Notes on the agriculture botany and zoology of China. XXII. The water plant - Euryale ferox in North Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 81-82. 81. 1919. Notes on the agriculture botany and zoology of China. XXIII. On the study of the flowers of the Manchurian wild apricot. Journal of the North China Branch of the Royal Asiatic Society 50: 82- 83. 82. 1919. Notes on the agriculture botany and zoology of China. XXIV. Observations on banana trees at Foochow. Journal of the North China Branch of the Royal Asiatic Society 50: 83-84. 83. 1919. Notes on the agriculture botany and zoology of China. XXV. A study of the rice cultivated at Foochow. Journal of the North China Branch of the Royal Asiatic Society 50: 84-90. 84. 1919. Notes on the agriculture botany and zoology of China. XXVI. On the kaoliang and barley, cultivated in Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 90-9 1 . 85. 1919. Notes on the agriculture botany and zoology of China. XXVII. Kaoliang and maize growing in Foochow. Journal of the North China Branch of the Royal Asiatic Society 50: 91-92. 86. 1919. Notes on the agriculture botany and zoology of China. XVIII. List of plants growing in Foochow. Journal of the North China Branch of the Royal Asiatic Society 50: 92-93. 87. 1919. Notes on the agriculture botany and zoology of China. XXIX. The insect trade in South China and some methods of catching insects. Journal of the North China Branch of the Royal Asiatic Society 50: 94-95. 88. 1919. Notes on the agriculture botany and zoology of China. XXX. The use of the horse-tail (Equisetum hyemale L.) in China. Journal of the North China Branch of the Royal Asiatic Society 50: 95-96. 89. 1919. Notes on the agriculture botany and zoology of China. XXXI. On new flagellata from Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 96-104. 90. 1919. Notes on the agriculture botany and zoology of China. XXXII. The bibliography on the botany, zoology and the rural economy of Manchuria. Journal of the North China Branch of the Royal Asiatic Society 50: 104-107. 9 1 . 1919. The tanning plants of Far East. Vestnik Man 'chzhurii (Man- churian Monitor) (In Russian). 92. 1919. The fossil animals from Northern Manchuria. Vestnik Man' chzhurii (Manchurian Monitor) 87. (In Russian). 93. 1919. The sugar industry in China and Manchuria. Vestnik Man' chzhurii (Manchurian Monitor) 109. (In Russian). 94. 1919. Rice cultivation in Northern Manchuria. Vestnik Man 'chzhurii (Manchurian Monitor) 109. (In Russian). 95. 1919. The European drug plants in the Far East. Vestnik Man 'chzhurii (Manchurian Monitor) 132. (In Russian). 96. 1919. Sericulture in Manchuria and in Russian Far East. Vestnik Man' chzhurii (Manchurian Monitor) 161. (In Russian). 97. 1920. A study of Chinese vegetable gardening in northern Man- churia. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 1-4: 4-13. (In Russian). 98. 1920. The awakening and growth of the vegetation at Harbin in 1919. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 1-4: 42-43. (In Russian). 99. 1920. Weeds of north Manchuria and their forage qualities. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 5-8: 43^7. (In Russian). 100. 1920. Experiments on the cultivation of the forage plant Symphytum asperimum at Harbin. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 5-8: 47-48. (In Russian). 101. 1920. Varieties of wheat growing in 1919 on the experimental farm of the Manchuria Agricultural Society at Harbin. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 5-8: 48-49. (In Russian). 1 02. 1 920. The Chinese millet 'paitzu' , their varieties and cultivation in the Far East. Journal 'Primorski Hosiaen ' no. 2. (In Russian). 103. 1 920. A study of tanning plants of Far East. Journal 'Primorski Hosiaen' no. 3. (In Russian). BORIS V. SKVORTZOV 99 104. 1920. Vegetable marrows and their cultivation by Chinese in Northern Manchuria. Journal 'Primorski Hosiaen ' no. 7. (In Russian). 105. 1920. Notes on the agriculture botany and zoology of China. XXXIII. The cultivation of water vegetables at Foochow. Journal of the North China Branch of the Royal Asiatic Society 51: 135-142. 106. 1920. Notes on the agriculture botany and zoology of China. XXXIV. On some varieties of peanuts grown in China. Journal of the North China Branch of the Royal Asiatic Society 51: 142-144. 107. 1920. Notes on the agriculture botany and zoology of China. XXXV. The horn-sherry of northern Manchuria. Journal of the North China Branch of the Royal Asiatic Society 51: 145-146. 108. 1920. Notes on the agriculture botany and zoology of China. XXXVI. On some wild vegetables of northern Manchuria. Journal of the North China Branch of the Royal Asiatic Society 51: 146-148. 109. 1920. Notes on the agriculture botany and zoology of China. XXXVII. The wild pears of northern Manchuria. Journal of the North China Branch of the Royal Asiatic Society 51: 148-149. 110. 1920. Notes on the agriculture botany and zoology of China. XXXVIII. The gourds melons and their cultivation by Chinese in Northern Manchuria. Journal of the North China Branch of the Royal Asiatic Society 51: 149-158. 111. 1 920. Forage plants of Manchuria and the Russian Far East with the Russian-Chinese dictionary of forage plants and products. Mongo- lian Expedition. Reports of 1915-1918 Volume II Application 4: 1-80. (In Russian). 112. 1920. Poppy cultivation in Northern Manchuria. Russki Golos 41. (In Russian). 113. 1920. Apricots and apricot-gardens at Harbin. Vestnik Man'chzhurii (Manchurian Monitor) no. 8. (In Russian). 1 14. 1920. The Chinese fruit-orchards at Ashiho. Vestnik Man 'chzhurii (Manchurian Monitor) no. 11. (In Russian). 115. 1921. Cultivation of Chinese onion in Manchuria. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 1-2: 1-18. (In Russian). 116. 1921. The awakening of the vegetation at Harbin in 1920. Sel'skoe Khoziaistvo v Sievernoi Man'chzhurii 1-2: 59-62. (In Rus- sian). 117. 1 92 1 . Manchurian trees and shrubs introduced in Europe, America and Japan. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 3: 71-77. (In Russian). 118. 1921. The wheat varieties growing near Harbin. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 4: 32-36. (In Russian). 119. 1921. The nursery and the experimental farm of Mr. N.N. Prikashikov at laomen station of Chinese Eastern Railway. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 4: 37^5. (In Russian). 120. 1921. Report on plant cultivated in 1920 on the Experimental Farm of the Manchurian Agricultural Society of Harbin. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 6-7: 12-18. (In Russian). 121. 1 92 1 . Water rice cultivation in north Manchuria and in neighbour districts. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 8-9: 46-80. (In Russian). 122. 1921. Memory of P.M. Karvovskii. Sel'skoe Khoziaistvo v Sievernoi Man' chzhurii 8-9: 93-94. (In Russian). 123. 1921. The awakening of the vegetation at Harbin in 1921. Sel'skoe Khoziaistvo v Sievernoi Man'chzhurii 10-12: 35-37. (In Russian). 1 24. 1 92 1 . The tree nursery in Harbin. Sel 'skoe Khoziaistvo v Sievernoi Man'chzhurii 10-12: 37-39. (In Russian). 125. 1921. Notes on the agriculture botany and zoology of China. XXXIX. The poppy culture in North Manchuria Journal of the North China Branch of the Royal Asiatic Society 52: 79-82. 1 26. 1 92 1 . Notes on the agriculture botany and zoology of China. XL. The mountain rice in North Manchuria. Journal of the North China Branch of the Royal Asiatic Society 52: 82-87. 1 27. 1 92 1 . Notes on the agriculture botany and zoology of China. XLI. The apricots grown in Harbin. Journal of the North China Branch of the Royal Asiatic Society 52: 87-92. 128. 1921. Notes on the agriculture botany and zoology of China. XLII. Fruit culture at Foochow. Journal of the North China Branch of the Royal Asiatic Society 52: 93-96. 129. 1921. Notes on the agriculture botany and zoology of China. XLIII. Plums of north Manchuria. Journal of the North China Branch of the Royal Asiatic Society 52: 96-101 . 130. 1921. Notes on the agriculture botany and zoology of China. XLIV. Micromeles alnifolia Kochne in North Manchuria. Journal of the North China Branch of the Royal Asiatic Society 52: 102-103. 131. 1 92 1 . Notes on the agriculture botany and zoology of China. XLV. Experiments in sericulture in North Manchuria. Journal of the North China Branch of the Royal Asiatic Society 52: 103-104. 132. 1921. Notes on the agriculture botany and zoology of China. XLVI. Bibliography on the botany zoology and rural economy of Manchuria. Second part. Journal of the North China Branch of the Royal Asiatic Society 52: 1 04-1 1 1 . 133. 1922. Licorice in Manchuria Mongolia and in China. Sel'skoe Khoziaistvo v Sievernoi Man'chzhurii 1-2: 7-18. (In Russian). 134. - - 1922. On some varieties of forage soybeans, cultivated by Chinese in Manchuria and in North China. Sel'skoe Khoziaistvo v Sievernoi Man'chzhurii 1-2: 76-80. (In Russian). 135. 1922. The wild amurian grap[e]. Sel'skoe Khoziaistvo v Sievernoi Man'chzhurii 1-2: 83-98. (In Russian). 136. 1 922. Japanese vegetables cultivated in North Manchuria. Sel 'skoe Khoziaistvo v Sievernoi Man'chzhurii 1-2: 99-104. (In Russian). 137. 1922. Mountain rice cultivation in North Manchuria. Viestnik Azii [The Herald of Asia] (Bulletin of the Russian Orientalist 's Society of Harbin) 48: 106-1 19. (In Russian). 138. * 1922. Notes on the agriculture, botany and zoology of China. XLVII. On the phytoplankton from the ponds of Tientsin. Journal of the North China Branch of the Royal Asiatic Society 53: 189-190. 139. * 1922. Notes on the agriculture, botany and zoology of China. XLVIII. On the winter phytoplankton of the fish-ponds of Foochow. Journal of the North China Branch of the Royal Asiatic Society 53: 190-195. 140. * 1922. Notes on the agriculture, botany and zoology of China. XLVIX. On some freshwater algae, collected by Mr. C.R.Kellogg in Hokchiang, Fukien. Journal of the North China Branch of the Royal Asiatic Society 53: 195. 141. 1923. The agriculture and experimental stations in the Jubilee Exhibition of Chinese Eastern Railway Company at Harbin in 1923. Sel'skoe Khoziaistvo v Sievernoi Man'chzhurii 7-10: 27-40. (In Rus- sian). 142. &GlouchovN.V. \923.Grow the wM grape. Sel'skoeKhoziai'stvo v Sievernoi Man'chzhurii 7-10: 47-50. (In Russian). 143. 1923. Fauna and Flora of Manchuria and the Russian Far East. Viestnik Azii [The Herald of Asia] (Bulletin of the Russian Orientalist 's Society of Harbin) 50: 161-278. (In Russian). 144. 1923. The agricultural experimental station of South Manchuria Railway Co. Viestnik Azii [The Herald of Asia] (Bulletin of the Russian Orientalist's Society of Harbin) 51: 291-295. (In Russian). 145. 1923. The wheats of Mudagian river valley north Manchuria. Viestnik Azii [The Herald of Asia] (Bulletin of the Russian Orientalist 's Society of Harbin) 51: 297-301. (In Russian). 146. 1923. The forgotten explorer of northern Manchuria. N.A. Baikov. Russki Golos (In Russian). 147. 1924. Farblose Euglenaceen aus nord-Mandchurei (China). Archivfur Protistenkunde 48: 1 80- 1 86. 148. 1924. Does Manchurian wheat degenerate? Ekonomicheskii vestnikMan 'chzhurii (Economical Monitor of Manchuria of China) 4: 3-5 (In Russian). 149. 1924. Vegetable oils in Manchuria and in China. Ekonomicheskii vestnik Man 'chzhurii (Economical Monitor of Manchuria of China) 6: 9-11 (In Russian). 150. 1924. Perilla or 'sutzu' in northern Manchuria and its future. Ekonomicheskii vestnik Man'chzhurii (Economical Monitor of Man- churia of China) 8: 9-11. (In Russian). 151. 1925. Uber einige Susswasseralgen der Umgegend von Peking (China). Archivfur Hydrobiologie (und Planktonkunde) 16 (2): 337- 340. 152. 1925. Zur kenntnis der Phycomycetes aus Nordmandschurei, China. Archivfur Protistenkunde 51: 428-433. 153. 1925. Zur Kenntnis der Mandschurischen Flagellaten. Beihefte zum Botanischen Zentralblatt 41 (3): 311-315 [July]. 100 D.M. WILLIAMS & G. REID 154. 1925. Uber neue und wenig bekannte formen der Euglenaceengattung Trachelomonas Ehrenberg. Bericht derDeutschen Botanischen Gesellschaft 43 (7): 306-3 15. 1 74. 155. 1925. The grain products of northern Manchuria. Book of Mer- chandise (Chinese Eastern Ray Co. Harbin). (In Russian). 175. 156. 1925. Description of new species of Trachelomonas, Ehrenberg from north Manchuria, China. China Journal of Science & Arts 3 (6): 176. 336-341 [June]. (The plate is reproduced in Sowerby (1930): 217, pi. XLII). 157. 1925. K Flore Man'chzhurii. I. De Malo manshurica (Max.) Kom. [Fragmenta Florae Manshuriae. I. De Malo manshurica (Max.) Kom.] Izvestiya Glavnogo Botanicheskogo Sada SSSR 24: 145-146 177. (German summary, p. 150). 158. 1925. K Flore Man'chzhurii. I. De Pyro ussuriensi Max. [I. Fragmenta Florae Manchuriae. II. De Pyro ussuriensi Max.] Izvestiya 178. Glavnogo Botanicheskogo Sada SSSR 24: 146-149. (German sum- mary,?. 150). 179. 159. 1925. On Trachelomonas hispida (Petty) Stein and its varieties. New Phytologist 24 (5): 299-305 [Published 3 1 December 1925]. 1 80. 160. * 1925. BeitragezurKenntnissderMandschurischendiatomaceen. Nuova Notarisia 36: 255-269. 181. 161. 1925. Gigantskoi kuvsiika Sungariiskih. [The Giant Water Lily of the Sungari lakes]. Obshchestvo Izucheniya Man 'chzhurskogo Kraya 1 82. Sekcia Estestvoenania (Miscellaneous Series, Manchuria Research Society, Natural History Section) A. 2: 1-9. 183. 162. 1925. Tykveiiye kul'tury severnoj Man'chzhurii. [Gourd plant cultivation by the Chinese in north Manchuria]. Otdel'noe Izdanie. 184. Obshchestvo Izucheniya Man' chzhurskogo Kraya Sekcia Estestvoenania (Miscellaneous Series, Manchuria Research Society, 185. Natural History Section) A. 4: 1-16. 163. 1925. Silva v severnoj Man'chzhurii. [The plum-tree in northern 186. Manchuria]. Obshchestvo Izucheniya Man 'chzhurskogo Kraya Sekcia Estestvoenania (Miscellaneous Series, Manchuria Research Society, 187. Natural History Section) A. 7: 1-16. 164. 1925. Wislouchiella planctonica, novyj rod vid iz gruppy Volvocales. [Wislouchiella planctonica nov. gen. et sp. of Volvocales from north Manchuria]. Obshchestvo Izucheniya Man' chzhurskogo 188. Kraya (Manchurian Research Society, Proceedings of the Sungari River Biological Station), B. 1 (1): 27-29. 165. 1925. Die Euglenaceengattung Trachelomonas Ehrenberg. Eine 189. systematische Ubersicht. Obshchestvo Izucheniya Man 'chzhurskogo Kraya (Manchurian Research Society, Proceedings of the Sungari River Biological Station. Gesellschaft zur Eforschung der Mandschurei. Arbeiten der Biologischen Sungari-Station), B. 1 (2): 1-101. 190. 166. 1925. Novyi presnovodnyi vid Ampidinium Cl. Et Lach. Iz Severnoi Manchzhurii. Enie [sic] neue Siisswasserart der Gattung 191. Amphidinium Clap. U. Lachm. aus der Nord-Mandschurei. Russkii Gidrobiologicheskii Zhurnal Izdavaemyi pri Volzhskoi 4 (7-9): 146- 192. 148. 167. 1925. The spring at Harbin in 1924. The Pedagogical Magasin 193. 'Voprosi Schkolnoi Juzni' 1: 76-78. (In Russian). 168. 1925. The giant water lily of the Sungari lakes. Vestnik Man'chzhurii (Manchurian Monitor) 3-4: 37^5. (In Russian). (See 194. item 161). 169. 1 925. Gourd-plants cultivation by Chinese in North Manchuria. Vestnik Man'chzhurii (Manchurian Monitor) 5-7: 11-26. (In Rus- 195. sian). (See item 162). 170. 1925. The plum tree in Northern Manchuria. Vestnik Man'chzhurii (Manchurian Monitor) 8-10: 45-60. (In Russian). (See item 163). 171. * 1 926. Uber einige Siisswasseralgen aus der nord-Mandschurei im Jahre 1916 gesammelt. Archiv fur Hydrobiologie (und Planktonkunde) 16 (3): 421^36. 196. 172. 1 926. Uber neue wenig bekannte Formen der Euglenaceengattung Trachelomonas Ehrenb. II. Bericht der Deutschen Botanischen Gesellschaft 44 (9): 603-621. 173. 1926. Beitrage zur Susswasseralgaen von Primorsk Gouvernement, Sibirien. I. Uber neue und bekannte Formen der 197. Euglenaceengattung Trachelmonas Ehrenb. der Umgegenden der Stadt Nikolsk-Ussurisk. Izvestiia luzhno-Ussuri'iskogo (Otdelen'iia Priamurskogo) Otdela Gosudarstvennago Russkago Geograficheskago Obshchestva (Bulletin of the Southern Ussuri branch of the Russian Geographical Society) 13: 227-232. 1926. On fresh-water algae collected by Dr. H. J. Chu at Peking, China. China Journal of Science & Arts 5 (4): 197-199 [October]. 1926. A contribution to the desmids of North Manchuria. Journal of Botany, British & Foreign 64 (761): 121-132. 1926. Polev'u Kul'turnye Rastenia severnoj Man'chzhurii. [The field Crops of Northern Manchuria]. Obshchestvo Izucheniya Man' chzhurskogo Kraya Sekcia Estestvoenania (Miscellaneous Se- ries, Manchuria Research Society, Natural History Section) A. 14: 1-18. (See item 178). 1926. Uber neue Euglenaceen der Gattung Trachelmonas Ehrenberg aus Belgien und Frankreich. Russkii Gidrobiologicheskii Zhurnal Izdavaemyi pri Volzhskoi 5 (1-2): 24-26. 1926. Field crops of northern Manchuria. Vestnik Man 'chzhurii (Manchurian Monitor) 10: 1-18. (In Russian). 1926. Cultivated meadow plants of northern Manchuria. Vestnik Man'chzhurii (Manchurian Monitor) 54. (In Russian). 1926. New flagellata from north Manchuria, China. Philippine Journal of Science 31 (2): 229-232 [Published 21 October 1926]. 1 927. Uber neue Chlamydomonden aus der Mandschurei. Archiv fur Hydrobiologie (und Planktonkunde) 18(1): 133-134. 1927. Uber einige Phycometes aus China. Archiv fur Protistenkunde 57 (1): 204-206. 1 927. Trachelomonas species from Nanking, China. China Jour- nal! (2): 87-91 [August]. 1927. Uber einige Peridinaceen aus der Nordmandschrei. Hedwigia 67 (3): 121-124 [July]. 1927. Einige Siisswasseralgan ausTobolske (Siberien). Hedwigia 67 (4-5): 246 [October]. 1927. Uber einige Siisswasseralgen aus Pamir (Asien). Hedwigia 67 (4-5): 247-248. 1927. K Flore Man'chzhurii. O Man'chzhurskom orehe (Juglans manshurica Maxim.). [Fragmenta Florae Manshuriae. (III. De Juglans manshuriae Max.)] . Izvestiya Glavnogo Botanicheskogo Sada SSSR 26 (1): 81-84. 1927. Sur la Vegetation lacustre de la vallee Sungari en Manshourie. Izvestiya Glavnogo Botanicheskogo Sada SSSR 26 (3): 268-284. 1 927. K Flore Man'chzhurii. II. Man'chzhurskie Formy vodanogo opeha. [Fragmenta Florae Manshuriae. II. Trapa generis formae manshuricae.] Izvestiya Glavnogo Botanicheskogo Sada SSSR 26 (6): 628-630. * 1927. Diatoms from Tientsin, north China. Journal of Botany, British & Foreign 65 (772): 102-109 [April]. 1927. A list of Cyanophyceae from North Manchuria, China. Journal of Botany, British & Foreign 65 (775): 195-198. 1927. On some algae from Sorei Lake, Sumatra. Journal of Botany, British & Foreign 65 (775): 198-199. * 1927. Freshwater algae and phytoplankton of the lakes and rivers of the Zaisan district, Altai Mountains, Siberia. Journal of Botany, British & Foreign 65 (777): 249-254. * 1927. On some fresh- water algae, collected by Dr. H.H. Chung in Amoy, China. Lingnaam Agricultural Review 4(1): 54-56 [March, published 14 July 1927]. 1927. Nabludenia nad zhnzn'u nitchatok - Zygnemaceae v okrestnostah g. Kharbina, v Severnoj Man'chzhurii. [Studies on the occurence [sic] and reproduction of Zygnemaceae in the environs of Harbin, North Manchuria.] Obshchestvo Izucheniya Man 'chzhurskogo Kraya (Manchurian Research Society, Proceedings of the Sungari River Biological Station) B. 1 (4): 1-24. (In Russian). 1927. The soybeans wild and cultivated in Eastern Asia. Obshchestvo Izucheniya Man' chzhurskogo Kraya Sekcia Estestvoenania (Miscellaneous Series, Manchuria Research Society, Natural History Section) A. Istoriko-Etnograficheskaya 22: 1-18. (See item 200 and 201). 1927. Manc'chzhurskaa Psenica. [Manchuria Wheat (abridged sketch)]. Otdel'noe Izdanie. Obshchestvo Izucheniya Man'chzhurskogo Kraya (Manchuria Research Society, Commercial and Industrial Section) A. Torgovo-Promyshleiiaa Sek. 18: 1-29. (See items 198 and 199). BORIS V. SKVORTZOV 101 1 98. 1 927. North Manchurian wheat. Vestnik Man 'chzhurii (Manchu- rian Monitor) 4: 47-57. (In Russian). 1 99. 1 927. North Manchurian wheat. Vestnik Man 'chzhurii (Manchu- rian Monitor) 5: 44-55. (In Russian). 200. 1927. The soybean wild and cultivated in Eastern Asia. Vestnik Man 'chzhurii (Manchurian Monitor) 9: 35-43. (In Russian). 201. 1927. The soybean wild and cultivated in Eastern Asia. Vestnik Man' chzhurii (Manchurian Monitor) 10: 20-25. (In Russian). 202. 1928. On some freshwater algae from Ceylon, collected by W.P. Lipsky in i 908. Annals of the Royal Botanic Gardens ofPeradeniya 11 (1): 109. [Published 28 May 1928]. 203. * 1928. Uber das Phytoplankton des Zaisansees, Siidsiberien. Archivfur Hydrobiologie (und Planktonkunde) 19(1): 165-171. 204. 1928. Die Euglenaceengattung Phacus Dujardin. Eine systematsche Ubersicht. Bericht der Deutschen Botanischen Gesellschaft 46: 105-125. 205. 1928. Some new and little known species of Trachelomonas Ehrenb. from north Manchuria China. Botanical Gazette, Chicago 85 (1): 90-96 [March]. 206. 1928. The botanical investigations in the Kirin and Zizikar provinces during the period of 1886-1923. Bulletin of the Manchuria Research Society 7: 62-66. (In Russian). 207. 1928. On phytoplankton of Alakkul lake. Bulletin of the Middle Asian Branch of Russian Geographical Society 18: 7-8. (In Russian). 208. 1928. On some desmids from Amoy, south China. China Journal 8 (3): 145-147 [March]. 209. * 1928. Ein Beitrag zur Bacillariaceen-Flora der nordostlichen Mongolei. Hedwigia 68: 311-314. 210. * & Meyer, C.I. 1928. A contribution to the diatoms of Baikal lake. Obshchestvo Izucheniya Man 'chzhurskogo Kraya (Manchurian Research Society, Proceedings of the Sungari River Biological Sta- tion), B. 1 (5): 1-55. 211. * 1928. K izucheniu zimnego fitoplanktona r. Sungari. [On winter phytoplancton [sic] of the Sungaree river in north Manchuria, China. Obshchestvo Izucheniya Man 'chzhurskogo Kraya (Manchurian Re- search Society, Proceedings of the Sungari River Biological Station), B. 1 (6): 17-20. 212. 1928. The Manchurian Walnut. Obshchestvo Izucheniya Man 'chzhurskogo Kraya (Manchuria Research Society) A. Istoriko- Etnograficheskaya 32: 1-11. (See item 214). 213. * 1928. Diatoms from Khingan, North Manchuria, China. Philip- pine Journal of Science 35 (1): 39-51 [Published 2 March 1928]. 214. 1928. The Manchurian Walnut (abridged sketch). Vestnik Man'chzhurii (Manchurian Monitor) 9: 54-60. (In Russian). 215. 1929. K Flore Man'chzhurii. 5. O Formah Ulmus pumila L., rastusih v severnoi Man'chzhurii. [Fragmenta Florae Manshuriae. 5. Ulmi pumilae formae manshuricae.] Izvestiya Glavnogo Botanicheskogo Sada SSSR 28 (5-6): 543-545. (German summary, p. 548). 216. 1929. K Flore Man'chzhurii. 6. Oberezah severnoi Man'chzhurii. [Fragmenta Florae Manshuriae. 6. Betulae generis formae manshuricae.] Izvestiya Glavnogo Botanicheskogo Sada SSSR 28 (5- 6): 545-548 (German summary, p. 548). 217. 1 929. Uber drei neue farblose Flagellaten aus China. Archivfur Hydrobiologie (und Planktonkunde) 20 (2): 322. 218. 1929. Einige neue und wenig bekannte Chlamydomonadaceae aus Manchuria. Archivfur Protistenkunde 66: 160-163. 219. * 1929. Diatoms from ponds of Peking. Bulletin of the Peking Society of Natural History 3: 43-48 [Published March 1929]. 220. * 1929. On some marine diatoms from Siberian shore of Japanese sea. Botanical Magazine, Tokyo 43 [506]: 57-59. 221. 1 929. Notes on trees and shrubs of northern Manchuria. 1 . Genus Populus L. China Journal 10 (6): 334-337 [June]. 222. * 1929. Freshwater diatoms from Amoy, south China. China Journal 11 (1): 40-44 [July]. 223. * 1929. On some diatoms from Seiko lake of Chosen, Japan. Journal of the Chosen Natural History Society 8: 9-14 [30 June 1 929]. 224. 1929. The Manchurian crab-apple tree in the Kirin Province of North Manchuria. Lingnan Science Journal 5 (4): 345-348 [April, 1928, published 28 December 1929] 225. * 1 929. Freshwater diatoms from Korea, Japan. Philippine Jour- nal of Science 38 (3): 283-291 [March, published 8 April 1929]. 226. * 1929. Marine diatoms from Dairen, South Manchuria. Philip- pine Journal of Science 38 (4): 419^30 [April, published 18 May 1929]. 227. * 1 929. Diatoms from Dalai-nor Lake, eastern Mongolia. Philip- pine Journal of Science 41 (1): 31-37 [published 7 December 1929]. 228. * 1929. Alpine diatoms from Fukien Province, south China. Philippine Journal of Science 41 (1): 39-49 [Published 7 December 1929]. 229. 1929. Materialy k poznaniu dikih plodovyh rastenii dal'nego Vostoka. I. Kizucheniu lesin Severnoi Man'chzhurii. [Materials to the knowledge of the wild growing fruit-plants of far East Asia I. On the hazel nut of north Manchuria.] Trudy po Prikladnoi Botanike Genetike i Selektsii (Prikladnoi Botanike i Selektsii) 22 (3): 213-218. (English summary p. 238). 230. 1929. Materialy k poznaniu dikih plodovyh rastenii dal'nego Vostoka. II. O Fragaria orientalis A. los. Rastusei v sev. Man'chzhurii. [Materials to the knowledge of the wild growing fruit-plants of far East Asia II. On Fragaria orientalis grown in North Manchuria.] Trudy po Prikladnoi Botanike Genetike i Selektsii (Prikladnoi Botanike i Selektsii) 22 (3): 218-222. (English summary p. 238). 231. 1929. Materialy k poznaniu dikih plodovyh rastenii dal'nego Vostoka. III. O dikih I kul'turnyh abrikosah Severnoi Man'chzhurri. [Materials to the knowledge of the wild growing fruit-plants of far East Asia III. On wild and cultivated apricot trees of North Manchuria.] Trudy po Prikladnoi Botanike Genetike i Selektsii (Prikladnoi Botanike i Selektsii) 22 (3): 222-229. (English summary p. 238-9). 232. 1929. Materialy k poznaniu dikih plodovyh rastenii dal'nego Vostoka. IV. K izuchenie visen, rastusih v Girinskoi provincii v sev. Man'chzhurii. [Materials to the knowledge of the wild growing fruit- plants of far East Asia IV. On cherries of Kirin Province of north Manchuria.] Trudy po Prikladnoi Botanike Genetike i Selektsii (Prikladnoi Botanike i Selektsii) 22 (3): 229-237. (English summary p. 239). 233. 1929. The elm-tree in northern Manchuria. Vestnik Man 'chzhurii (Manchurian Monitor) 11: 52-58. (In Russian). 234. * 1929. Materialy po Izucheniu Vodoroslej Primorskoj Gubernii. Diatomovye Vodorosli Ozera Khanka. [A contribution to the algae, Primorsk district of Far East, USSR. Diatoms of Hanka Lake. Beitraege zur Suesswasseralgan von Primorsk Government, Siberien. II. Die Bacillarien des Hankasees.] Zapiski Uzhno-Ussurijsk Old. Cos. Russk. Oeograf. Obs. (Memoirs of the Southern Ussuri Branch of the State Russian Geographical Society) 3: 1-66. (German summary). 235. * 1930. Notes on Ceylon diatoms. I. Annals of the Royal Botanic Gardens ofPeradeniya 11 (3): 251-260 [published 30 January 1930]. 236. 1 930. Notes on trees and shrubs of Northern Manchuria. 2. Genus Actinidia Lindl. China Journal 12 (4): 214-216 [April]. 237. 1930. Notes on trees and shrubs of Northern Manchuria. 3. Genus Acer L. China Journal 12 (6): 358-362 [June]. 238. 1930. K Flore Man'chzhurii. 7. Man'chzhurskie boarysniki. [Fragmenta Florae Manshuriae. 7. De Crataego Manshuriae.] Izvestiya Glavnogo Botanicheskogo Sada SSSR 29 (3^*): 382-384. (German summary, p. 390). 239. 1930. K Flore Man'chzhurii. 8. O ipah severnoi Man'chzhurii. [Fragmenta Florae Manshuriae. 8. De Tiliis Manshuriae.] Izvestiya Glavnogo Botanicheskogo Sada SSSR 29 (3^4): 384-387. (German summary, p. 390). 240. 1930. K Flore Man'chzhurii. 9. Rod Actinidia Lindl. V severnoi Man'chzhurii. [Fragmenta Florae Manshuriae. 9. Actinidiae generis formae manshuricae.] Izvestiya Glavnogo Botanicheskogo Sada SSSR 29: 388-389. (German summary, p. 390). 24 1 . 1 930. New plants from North Manchuria, China. Lingnan Science Journal 6: 205-220 [September 1928; published 28 January 1930]. 242. & Nedelskii A.F. 1 930. Soybeans in the United States of America. Vestnik Man 'chzhurii (Manchurian Monitor) 3: 1-1 1 . (In Russian). 243. 1930. Changes in the vegetation in the surroundings of Harbin. Vestnik Man'chzhurii (Manchurian Monitor) 4: 60-67. (In Russian). 244. 1930. A contribution to the vegetation of Barga. Vestnik Man 'chzhurii (Manchurian Monitor) 6: 59-66. (In Russian). 102 D.M. WILLIAMS & G. REID 245. 1 930. A contribution of the vegetation of the eastern slopes of the Great Hingan Mountains in the Kirin province of northern Manchuria. 272. Vestnik Man 'chzhurii (Manchurian Monitor) 10: 40-57. (In Russian). 246. * 1 930. Materialy po flore vodoroslej Aziatskoj chasti SSSR. I. O fitoplanktone oz. Teleckogo. Zhurnal Russkogo Botanicheskogo 273. Obshchestva pri Akademii Nauk 15 (3-4): 91-92. 247. 1 93 1 . Notes on trees and shrubs of Northern Manchuria. 4. Genus Pyrus L. China Journal 14 (6): 329-33 1 [June]. 274. 248. 1931. Notes on trees and shrubs of Northern Manchuria. 5 . Genus Syringa L. China Journal 15 (2): 98-100 [August]. 275. 249. 1931. Notes on trees and shrubs of Northern Manchuria. 6. Genus Vitis L. and Ampelopsis Michx. China Journal 15(4): 200-20 1 [Octo- ber]. 276. 250. 193 1 . Notes on trees and shrubs of Northern Manchuria. 7. Genus Sambucus L. and Viburnum L. China Journal 15 (6): 304-307 [De- 277. cember]. 251. 1931. The cultivation of sugar-sorgo in Northern Manchuria. 278. Viestnik Kitaisko-vostochnoi zheleznoi dorogi (Bulletin of Chinese Eastern Railway Company) 11: 5-6. (In Russian). 279. 252. * 193 1 . Phytoplankton from Siberia. I. From the Akmolinsk lake district, collected by P.T. Ignatow in 1899 and in 1901. Journal of 280. Botany, British & Foreign 69 (818): 33-36. 253. * 1931. Phytoplankton from Siberia. II. From the Altai mountains 281. of Siberia, collected by A.N. Sedelnikov in 19 14 and \9\6.Journalof Botany, British & Foreign 69 (818): 36-38. 282. 254. * 1931. Phytoplankton from Siberia. III. From the Amur river. Journal of Botany, British & Foreign 69 (819): 69-72. 283. 255. 1931. Mycetozoa from Northern Manchuria, China. Philippine Journal of Science 46 (1): 85-93 [September, published 21 August 1931]. 284. 256. * 1931. Pelagic diatoms of Korean strait of the Sea of Japan. Philippine Journal of Science 46 (1): 95-122 [September, published 285. 21 August 1931]. 257. * 1 93 1 . Marine diatoms from the Kanazawa Oyster Experimental 286. Station of Japan. Philippine Journal of Science 47 (1): 119-127 [January, published 9 December 193 1 ]. 287. 258. * 1931. Marine littoral diatoms from environs of Vladivostok. Philippine Journal of Science 47 (1): 129-150 [January, published 8 December 1931]. 288. 259. * 1931. Marine diatoms from Formosa Strait. Philippine Journal of Science 47(1): 151-161 [January, published 9 December 1931]. 289. 260. 1 93 1 . Vegetation of the Manchurian plain. Vestnik Man 'chzhurii (Manchurian Monitor) 1: 79-87. (In Russian). 261. 1931. Vegetation of the lower part of the Ashiho river valley in northern Manchuria. Vestnik Man 'chzhurii (Manchurian Monitor) 4: 290. 77-81. (In Russian). 262. 1 93 1 . Vegetation of small Hingan Mountains in northern Man- churia. Vestnik Man' chzhurii (Manchurian Monitor) 7: 43^6. (In 291. Russian). 263. 1 931. The dissappearing forests of Dachungshan, Kirin Province 292. in Manchuria. Vestnik Man' chzhurii (Manchurian Monitor) 7: 46. (In Russian). 264. 1931. Soils of Manchuria. Vestnik Man' chzhurii (Manchurian 293. Monitor) 8: 53-59. (In Russian). 265. * 1932. Notes on Ceylon Diatoms. 2. Annals of the Royal Botanic Gardens ofPeradeniya 11 (4): 333-338 [published 20 Febuary 1932]. 294. 266. 1932. Flagellaten aus der Nordmandschurei (im Jahre 1931 gesammelt). Archivfur Protistenkunde 77: 522-528 [Abstract inActa Phytotaxonomicat el Geobotanica 2: 66-67, 1 February 1933]. 295. 267. 1932. The Manchurian floristic region and its geographical landscapes. Bibliograficheskii Sbornik Biblioteki Vostochno'i Kitaiskoi 296. Zheleznoi Dorogi 2 (5): 1-24. (In Russian). 268. 1 932. Notes on trees and shrubs of Northern Manchuria. 8. Genus Mahus Mill. China Journal 17 (2): 84-88 [August]. 297. 269. 1932. The experiments in cultivation of Hibiscus cannabinus L. in Northern Manchuria. Economical Bulletin of Chinese Eastern Rail- way Company 23-24: 2. (In Russian). 298. 270. 1932. Beitrage zur kenntnis der Flagellaten aus Mukden, Mandschurei, China. Journal of Oriental Medicine 14: 1-4. 271. 1932. Flagellaten aus Korea, Japan. Journal of the Chosen 299. Natural History Society 13: 8-10 [20 April 1932]. 1932. Mushrooms and its commercial importance in China. Vestnik Man'chzhurii (Manchurian Monitor) 6-7: 1 15-123. (In Rus- sian). * 1932. Diatoms from the bottom of the Sea of Japan. Philippine Journal of Science 47 (2): 265-280 [February, published 12 January 1932]. 1932. Desmidsfrom Korea, Japan. Philippine Journal of Science 49 (2): 147-158 [October, published 22 September 1932]. 1932. The perishing forests of the ridge Tachin-Shan of Kirin province. Vestnik Man 'chzhurii (Manchurian Monitor) 2: 65-69. (In Russian). 1932. Ginseng and its commercial importance in China. Vestnik Man'chzhurii (Manchurian Monitor) 3: 80-89. (In Russian). 1933. Sweet potatoes taro and yam and their cultivation in China. Vestnik Man 'chzhurii (Manchurian Monitor) 1 : 49-60. (In Russian). 1933. Native medicine and drug trade in China. Vestnik Man'chzhurii (Manchurian Monitor) 4: 7285. (In Russian). 1933. Native medicine and drug trade in China. Vestnik Man 'chzhurii (Manchurian Monitor) 5: 35-42. (In Russian). 1933. Native medicine and drug trade in China. Vestnik Man'chzhurii (Manchurian Monitor) 6: 59-69. (In Russian). 1933. Native medicine and drug trade in China. Vestnik Man'chzhurii (Manchurian Monitor) 7: 55-66. (In Russian). 1933. Native medicine and drug trade in China. Vestnik Man'chzhurii (Manchurian Monitor) 89: 72-78. (In Russian). 1933. Agricultural practices in Manchurian village of Aschiho district, Kirin Province, Manchuria. Vestnik Man 'chzhurii (Manchu- rian Monitor) 12: 49-5 1 . (In Russian). 1934. Fauna and flora. Manchuria economical and geographi- cal. Description, Part 1: 57-74. (In Russian). 1 935 . Notes on trees and shrubs of Northern Manchuria. 9. Genus Quercus L. China Journal 23 (1): 54-56 [July]. 1935. Notes on trees and shrubs of Northern Manchuria. 10. Genus Euonymus L. China Journal 23 (3): 176-178 [September]. * 1935. Diatoms from Poyang lake, Kiangsi Province, China. Philippine Journal of Science 57 (4): 465^78 [August, published 5 October 1935]. * 1935. Diatoms from Calcutta, India. Philippine Journal of Science 58 (2): 179-192 [October, published 5 November 1935]. * 1 935. Diatomees recoltees par le Pere E. Licent au cours de ses voyages, dans le Nord de la Chine, au bas Tibet, en Mongolie et en Mandjourie. Publications du Musee Hoango ho-Pai ho Tien-Tsin 36: 1-43. 1936. Notes on trees and shrubs of Northern Manchuria. 12. Genera Chosenia Nakai and Salix Linnaeus. China Journal 25 ( 1 ): 42- 57 [July]. 1936. Notes on trees and shrubs of Northern Manchuria. 13. Conifers. China Journal 25 (3): 170-171 [September]. * 1936. The Neogene diatoms from the Ampen District, S. Kankyo-do, eastern coast of Tyosen. Bulletin of the Geological Survey ofTyosen 12: 137. * 1936. The diatoms from Kizaki Lake, Honshu Island, Nippon. Philippine Journal of Science 61 (1): 9-73 [September, published 6 November 1936]. * 1936. Diatoms from Biwa Lake, Honshu Island, Nippon. Phil- ippine Journal of Science 61 (2): 253-296 [October, published 15 December 1936]. * 1937. Diatoms of Lake Michigan. I. American Midland Natu- ralist 18 (4): 652-558 [July]. 1937. Contribution to our knowledge of the freshwater algae of Rangoon Burma, India. I. Euglenaceae from Rangoon. Archiv fur Protistenkunde 90: 69-87. * 1937. Subaerial diatoms from Hangchow, Chekiang Province, China. Bulletin of the Fan Memorial Institute of Biology (Botany) 7 (6): 2 19-230 [20 March 1937]. * 1937. Neogene diatoms from Wamura, Nagano Prefecture, Central Nippon. Memoirs of the College of Science, Kyoto Imperial University B 12 (2): 137-156 [March]. * 1937. Neogene diatoms from Saga Prefecture, Kiushiu Island, BORIS V. SKVORTZOV 103 Nippon. Memoirs of the College of Science, Kyoto Imperial University B 12(2): 157-1 74 [March]. 300. 1 937. Notes and papers which have been printed and mostly used separately from 1917 to 1937. Naturalist Man 'chshurii (Naturalist of 325. Manchuria) 2: 38-48. 301. * 1937. Notes on fossil diatoms from New South Wales, Aus- tralia. I. Fossil diatoms from diatomaceous earth, Cooma, N.S.W. 326. Proceedings of the Linnean Society of New South Wales 62 (3-4): 1 75- 1 80 [Published 1 5 September] . 302. 1937. Notes on algal flora of Manchoukuo. [I. Midsummer 327. phytoplankton of a marshy river branch in the Sungari Plain near Harbin.] Botanical Magazine, Tokyo 51 (607): 627-635 [July]. 303. * 1937. Notes on algal flora of Manchoukuo. II. Botanical Magazine, Tokyo 51 (608): 677-88 [August]. 328. 304. 1 937. Notes on algal flora of Manchoukuo. III. Botanical Maga- zine, Tokyo 51 (609): 738-42 [September]. 305. * 1937. Notes on algal flora of Manchoukuo. IV. [II List of algae 329. from a single growth of Spirogyra, Collected in Environs of Harbin.] Botanical Magazine, Tokyo 51 (610): 783-90 [October]. 306. * 1 937. The diatoms from Ikeda Lake, Satsuma Province, Kiusiu Island, Nippon. Philippine Journal of Science 62 (2): 191-218 [Febru- 330. ary, published 13 May 1937]. 307. * 1937. Bottom diatoms from Olhon-gate of Baikal Lake, Siberia. Philippine Journal of Science 62 (3): 293-377 [March, published 21 June 1937]. 308. * 1937. Neogene Diatoms from eastern Shantung. Bulletin of the Geological Society of China 17(1^): 193-204. 309. * 1938. Diatoms from Philippines, I. Diatoms from drinking water, Balara, Rizal Province. Philippine Journal of Science 64 (3): 331. 287-298 [November, 1937, published 31 January 1938]. 310. * 1938. Subaerial diatoms from Shanghai. Philippine Journal of Science 64 (4): 443^51 [December, 1937, published 8 March 1938]. 311. * 1938. Fresh-water diatoms from the environs of Vladivostok. Philippine Journal of Science 65 (3): 251-261 [March, published 18 332. May 1938]. 312. * 1938. Subaerial diatoms from Pin-Chiang-Sheng Province, Manchoukuo. Philippine Journal of Science 65 (3): 263-281 [March, published 18 May 1938]. 313. * 1938. Diatoms from Kenon Lake, Transbaikalia, Siberia. Phil- 333. ippine Journal of Science 65 (4): 399-424 [April, published 30 June 1938]. 314. * 1938. Diatoms from Argun River, Hsing-An-Pei province, Manchoukuo. Philippine Journal of Science 66 (1): 43-74 [May, published 26 July 1938]. 315. * 1938. Diatoms from a peaty bog in Lianchiho River valley, eastern Siberia. Philippine Journal of Science 66 (2): 161-182 [June, 334. published 5 September 1938]. 316. * 1938. Diatoms from a mountain bog, Kaolingtze, Pinchiang- Sheng Province, Manchoukuo. Philippine Journal of Science 66 (3): 343-362 [July, published 3 October 1938]. 317. * 1938. Diatoms from Chengtu, Szechwan, western China. Phil- 335. ippine Journal of Science 66 (4): 479-496 [August, published 18 October 1938]. 318. * 1938. Diatoms collected by Mr. Yoshikazu Okada in Nippon. I. Mountain bog diatoms flora from Prov. Sinano. Journal of Japanese Botany 14 (2): 204-217 [February]. 319. * 1938. Notes on the algal flora of New Zealand. I. Fresh- water 336. diatoms from New Zealand. Philippine Journal of Science 67 (2): 1 67- 174 [October, published 23 November 1938]. 320. * 1938. Notes on the algal flora of New Zealand. II. Fresh- water algae from Napier. Philippine Journal of Science 67 (4): 411-419 [December, published 29 December 1938]. 32 1 . 1 940. On Chinese practises of tomato cultivation in environs of Harbin. The Railway Magazine Harbin 35: 61-65. (In Russian). 322. 1940. The sugar corn cultivation in Harbin. The Railway Maga- zine Harbin 36: 1 1 1-1 15. (In Russian). 323. 1940. On Chinese practises of cucumber cultivation in environs of Harbin. The Railway Magazine Harbin 37: 58-61 . (In Russian). 338. 324. Kuzmin V. & Skvortzov B.V. 1941. Bog and water plants of Manchu- ria, I. The genus Potamogeton. Philippine Journal of Science 74 (4): 395-409 [April, published 30 June 1941]. Skvortzov notes that the reprints were destroyed by fire during the last war in Manila (item 419, p. 411). Skvortzov B.V. 1943. (Redactore) Index Florae Harbinensis sive enumeratio plantarum circa Harbin sponte nascentium hucusque cognitarum. Fasc. I. Societas Investigatorum Przewalskiensis Harbin. & Baranov A. 1943. Diagnoses Plantarum Novarum et minus Cognitarum Mandshuriae. Societas Investigatorum Przewalskiensis Harbin. Baranov A. & Skvortzov B.V. 1945. Obzor vidov roda schavel' (Rumex L.) iz Severnoi Man'chzhurii. [Revisio specierum generis Rumex L. ex Manshuria boreal!.] Izvestiia Kharbinskogo Kraevedcheskogo Muzeia 1: 43-52. (In Russian). Skvortzov B.V. 1945. Rabota A. D. Voeikova v oblasti sadovodstva I akklimatizacii. [A. D. Voeikov's work in horticulture and acclimatiza- tion.] Izvestiia Kharbinskogo Kraevedcheskogo Muzeia 1. (In Russian). 1945. K istorii Obsestva Izuconia Man'chzhurskogo Kraa I Sozdannogo im Muzeia. [To the history of Manchuria Research Soci- ety and its museum.] Izvestiia Kharbinskogo Kraevedcheskogo Muzeia 1. (In Russian). * 1946. Novye I malo izvestnye vidy Algae, Flagellatae, Phycomicetae iz Azii, Ameriki, Afriki, a takzhe s ostrovov Aponii I Cejlona, olisannye v 1931^5 g.g. s. 18 tablicami risunkov. [Species novae et minus cognitae Algarum, Flagellatarum et Phycomicetarum Asiae, Africae, Americae et Japoniae nee non Ceylon anno 1931-1945 descripto et illustrate per tab. 1 - 1 8] . Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Proceedings of the Harbin Society Natural History and Ethnography) 2. Botany: 1-34. 1946. Vladimir Leont'evich Komarov. [V.L. Komarov (obitu- ary)]. Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Proceedings of the Harbin Society Natural History and Ethnography) 5. To The Memory of V.L. Komarov Member of the Academy of Science USSR: 5-8. 1947. Kul'tura Amerikanskogo muskatnogo vinograda v g. Kharbine. [American grape-vine cultivation in Harbin.] Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Pro- ceedings of the Harbin Society Natural History and Ethnography) 6. Agriculture: 7-12. 1947. K izucheniu vdiania holodioj zimy 1944-45 g.g. na morozostojkost' plodovyzh, agodnyzh I Irugikh rastenij g. Kharbina I ego okrestnostej. [To the study of the influense [sic] of the cold winter cold of 1944/1945 on the hardiness of the fruit-trees and other plants in Harbin. Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Proceedings of the Harbin Society Natural History and Ethnography) 6. Agriculture: 13-16. 1947. K aprobachii baklazhan, razvodimykh v Man'chzhurii. [Subspecies and Proles of the Egg plant (Solanum melongena L.) grown in Manchuria.] Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Proceedings of the Harbin Society Natural History and Ethnography) 6 Agriculture: 2 1-26. Kuzmin V. I. & Skvortzov B. V. 1947. Lekarstvennye rastenia otechestvennoi narodnoi i nauchnoi meditsiny v Man'chzhurii ikh primenie. [Russian medical herbs in Manchuria and their use]. Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Pro- ceedings of the Harbin Society Natural History and Ethnography) 6. Agriculture: 37-54. & 1949. Obzor vidov roda Bolotnits Heleocharis R. Br. sobrannykh v Severnoi Man'chzhurii. [Revisio specierum generis Heleocharis R. Br. (Cyperaceae) ad Manshuriam Borealum (China) Collectarum]. Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Proceedings of the Harbin Society Natural History and Ethnography) 9. Botany: 15-18. Skvortzov, B. V. 1 95 1 . Experiments in breeding ofJuglans manshurica Maxim, and Juglans Sieboldiana Maxim, var. cordilformis Makino in the Forest Experimental Station of the N.E. College of Agriculture at Harbin in April of 1950. Journal of the N.E. College of Agriculture in Harbin 2: 102-103. (In Russian). Baranov A. & Skvortzov B.V. 1954. Tipy rastitel'nosti Tigrovoi padi i Tigrovoi sopki (Sevemaia Man'chzhuria). [Vegetation types of the valley of Hoangniho River and the Takuok'ui Mt., North Manchuria]. 104 D.M. WILLIAMS & G. REID Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Acta Societatis Harbinensis pro Investigatione Naturae et Ethnographic) 12. Botany: 1-12. 339. & 1954. Opisaniia novykh i maloizvestnykh rastenii Man'chzhurii. Vypusk II. [Diagnoses plantarum novarum et minus cognitarum Mandshuriae]. Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov (Acta Societatis Harbinensis pro Investigatione Naturae et Ethnographic) 12. Botany: 27-38. (Item 326 is part I). 340. Skvortzov B. V., Gordeiev T., Kuzmin V., Baranov A. & Chang yui Liang. 1955. Spisok rastenii okrestnostei Kharbina. 2 izdanie, ispravlennoe I dopolnennoe. Index Florae Harbinensis sive enumeratio plantarum circa Harbin sponte nascentum hucusque cognitarum. Edito 2. Zapiski Kharbinskogo Obshchestva Estestvoispytatelei i Etnografov 14: 1^2. 341. , Noda, M., Wang, S. & Liou, S. 1955. Illustrated flora of ligneous plants of north-eastern China, [cited in Noda, 1960, 1971]. 342. Skvortzov, B. V. 1958. New and rare Flagellatae from Manchuria, eastern Asia. Philippine Journal of Science 86 (2): 139-202 [June 1957, published 28 July 1958]. 343. 1 959. Algae novae et minus cognitae Chinae Boreali-Orientalis. 1. Bulletin Herbarium of North East Forest Academy Harbin 1: 2731 [December]. 344. 1 960. Algae novae et minus cognitae Chinae Boreali-Orientalis. 2. Flagellatae prope oppidum Mukden in anno 1957 collectarum. Bulletin Herbarium of North East Forest Academy Harbin 2: 1-8 [March]. 345. 1961. Harbin Chrysophya[s/c], China Boreali-Orientalis. Bw//e- tin Herbarium of North East Forest Academy Harbin 3: 1-70 [December]. 346. Baranov A. & Skvortzov B. V. 1965. Plante novae et minus cognitae florae Chinae boreali orientalis. Quarterly Journal of Taiwan Mu- seum 18(1-2): 219-231 [June]. 347. Baranov A. & Skvortzov B. V. 1965. What is Allium macrostemon from Manchuria, China? Quarterly Journal of Taiwan Museum 18 (3- 4): [December]. 348. Skvortzov, B. V. 1966. T.P. Gordeiev's work in Machuria in A.I. Baranov (Ed) Collection of Materials to the biography of T.P. Gordeiev. 349. 1966. New and little known species of the genus Urceolus Meresch. (Peranemaceae, Euglenophyta) from N.-E. China and Brasil. Journal of Japanese Botany 41 (7): 203-213. 350. 1966. Species of the genus Heteronema (Duj.) nob. (Peranemaceae, Euglenophyta, Flagellata) collected in N.E. China, Eastern Asia and in Brasil, South America with 34 figures. Quarterly Journal of the Taiwan Museum 19 (1-2): 141-153 [June 1966]. 35 1 . Baranov A. & Skvortzov B. V. 1 966. Plante novae et minus cognitae florae Chinae boreali-orientalis (II). Quarterly Journal of Taiwan Museum 19 (1-2): 157-164 [June]. 352. Skvortzov, B. V. 1 966. Uber eine neue gattung Kolbena nov. gen., fam. Astasiaceae, Ord. Eugleninae, aus Brasilien. Svenska Botanisk Tidskrift 60(1): 8 1-84. 353. 1967. New and interesting species of Euglena Ehrenb. from the subtropics of Brasil. Nova Hedwigia 14: 379-386. 354. 1967. Notes on the Flagellata of Hongkong. I. Species of the genus Chologonium Ehr. (Chlorophyceae, Volvocales, Chlamydomonadaceae) from polluted waters in Hong Kong. Gardens ' Bulletin Singapore 22 (2): 187-189 [Published 27 December]. 355. 1967. Algological notes I. On new genera of colourless Flagellata genus Silvamonas gen. nov. of the Fam. Polytomellaceae Aragao Ord. Polyblepharidales, Vovlocinaceae [sic] from Sao Paulo, Brasil. Bulle- tin of Japanese Society ofPhycology 15 (1): 37-38 [April]. 356. 1 967. Algological notes II. The first records of species of genus Sphaerella Sommerfelt (Spharellaceae, Volvocales) from the sub- tropics of Brasil. Bulletin of Japanese Society of Phycology 15 (1): 38^0 [April]. 357. 1 967. Algological notes III. On species of Chlamydomonas Ehr. (Chlamydomonaceae, Volvocales) from Durban, Matal [sic], South Africa. Bulletin of Japanese Society of Phycology 15 (1): 40-44 [April]. 358. 1967. On new colourless Flagellata genus Huber-Pestaloz- ziamonas gen. nov. Fam. Peranemaceae, Euglenophyta. Bulletin of Japanese Society ofPhycology 15 (2): 76-81 [August]. 359. 1967. New and little known species of the genus Pteromonas Seligo (Phacotaceae, Volvocineae) from Harbin, Northern Manchuria, China. (Aug.) Bulletin of Japanese Society ofPhycology 15 (2): 82-95 [August]. 360. 1967. Generis novae flagellatarum in Chinae et Brasilia. Ceylon Journal of Science New Series: Biological Sciences 6 (2): 217-221 [February]. 361 . * 1967. Notes on Ceylon diatoms III. Ceylon Journal of Science New Series: Biological Sciences 6 (2): 221-224 [February]. 362. Skvortzov B. V. & Noda, M. 1967. Desmid flora from a Sphagnum swamp-lake in the crater of an ancient volcano Geershuschan of Wutalienchi area, northern Manchuria, China. Journal of Japanese Botany 42 (5): 136-250. (Abstract in Botanical Magazine, Tokyo 82 (968): 90, 1969). 363. & 1967. On species of a green flagellata of the genus Chlorogonium Ehr. Volvocineae, Chlorophyceae from Japan, Hong Kong and Brasil. Journal of Japanese Botany 42 (7): 193-200 [Ab- stract in Botanical Magazine, Tokyo 82 (968): 90, 1969]. 364. 1968. Flagellata species of genus Petalomonas Stein, Fam. Peranemaceae, Euglenophyta with quadrangular cells recorded in swamp water of Sao Paulo in mountain subtropical region of Brasil. Acta Botanica Venezuelica 3: 293-296 [December]. 365. Skvortzov B. V. & Noda, M. 1 968. Genus Stromia gen. nov. - A new colourless Flagellata from subtropics of Brasil, South America. Bulle- tin of Japanese Society of Phycology 16 (3): 143-144 [December]. (Abstract in Botanical Magazine, Tokyo 82 (978): 492, 1 969; the figure was mistakenly not printed according to Skvortzov (item 396: 414)). 366. & 1 968. On new and little known Flagellata from N.-E. Asia and South America. Bulletin Japanese Society ofPhycology 16: 145- 155. (Abstract in Botanical Magazine, Tokyo 82 (978): 491 , 1969). 367. 1968. Nota algologica I. Um novo genero de Flagelado verde de Sao Paulo: Rotundomastix genero Nov. fam. Heteromadtigaceae Korsch. Classe Volvocinaeae. Instituto de Engenharia Sanitaria Sursan: 1-9. 368. 1 968. Nota algologica. II. Um novo genero de flagelado verde de Sao Paulo: Palmercriamonas genero nov. Pyyrophyta. Instituto de Engenharia Sanitaria Sursan: 19. 369. 1968. Notas algologica. III. Nova familia Generos e especies de flagelados verdes de Sao Paulo Classe Protochrysidineae Skv. 1 96 1 . Chrysophyta. Parte-I. Instituto de Engenharia Sanitaria Sursan: 1- 17. 370. 1968. Genus novum Swirenkoiamonas (Chloromonadales) e Brasilia. Novosti Sistematiki Nizshikh Rastenii 1968: 63-64. 371. 1968. Genus novum Meyeriella (Cryptomonadales) e China. Novosti Sistematiki Nizshikh Rastenii 1968: 65-66. 372. 1968. New and little known Peridineae from northern Manchu- ria, China. Quarterly Journal of Taiwan Museum 21 (1-2): 79114 [June]. 373. 1968. New genera of green flagellatae recorded in N.-O. China (with 17 figs). [Generis novae flagellatarum viridis in N.-O. China inventae (cum fig. 17).] Revue Algologique 2: 121-130 [August]. 374. Bicudo, C. E. de M. & Skvortzov B. V. 1968. Contribution to the knowledge of Brazilian Dinophyceae - Immobile Anais da Sociedade Botanica do Brasil, XIX Congresso Nacional de Botanica: 31-39. 375. Skvortzov, B. V. 1968. On some species of Euglena Ehr. from Singa- pore (De Species Euglenae Ehr. ex Singapore). Gardens' Bulletin Singapore 22 (3): 447^50 [Published 29 June]. 376. 1968. On a new species of the genus Collodictyon Carter, a colourless flagellata new to the Hongkong flora (De specie et genere Collodictyon Carter nova ad flora Hongkongensis.) Gardens' Bulletin Singapore 22 (3): 451-454 [Published 29 June]. 377. 1968. New genera of primitive green flagellata from Hongkong and Sao Paulo, Brazil. Gardens' Bulletin Singapore 22 (3): 455^59 [Published 29 June]. 378. 1 968. On new genera of Cryptomonadinae recorded from north- eastern China. Journal of Japanese Botany 43 (1): 8-16 [Published January]. 379. Skvortzov B. V. & Noda, M. 1 968. On Brasilian and European species BORIS V. SKVORTZOV 105 of genus Vacuolaria. Journal of Japanese Botany 43 (3): 69-76 [March]. (Abstract in Botanical Magazine, Tokyo 82 (968): 90, 1969). 380. & 1968. On colourless flagellata of genus Entosiphon Stein, Fam. Peranemaceae, Euglenophyta collected in Sao Paulo, Brasil, South America. Journal of Japanese Botany 43 (6): 164-168 [June]. 381. & 1 968. Euglena species with one or two chromatophores recorded from South America and South Africa in 1 962-1 966. Journal of Japanese Botany 43 (8): 225-233 [August]. 382. & Bicudo, C. E. de M. 1 968. Isogamy in Furcilla stigmatophora (Chlamydomonadaceae Chlorophyceae). Sellowia 20: 45-49 [Au- gust]. 383. 1969. The list of new genera and type species of flagellates and algae published in 1925-1957. Part I. Acta Hydrobiologia, Hydrographica et Protistologia 34: 337-342. 384. 1969. The list of new genera and type species of flagellates and algae published in in 1959-1961. Part. II. Acta Hydrobiologia, Hydrographica et Protistologia 34: 343-347. 385. 1969. The list of new genera and type species of flagellates and algae published in 1967. Part III. Acta Hydrobiologia, Hydrographica et Protistologia 34: 348-352. 386. * 1969. Siberian and Chinese freshwater diatoms new to science. Botaniska Notiser 122: 375-379. 387. 1 969. On new brown or olive flagellata genus Protocryptochrysis gen. nov. Cryptomonadineae recorded in N.E. China. Quarterly Jour- nal of Taiwan Museum 22 (1-2): 223-239 [June 1969]. 388. 1969.NewandlittleknowngeneraofcolourlessflagellatesofFam. Astasiaceae, Euglenophyceae recorded in 1954-1968 from N.E. China and Brasil. Quarterly Journal of Taiwan Museum 22 (3-4): 223-239 [December 1969]. 389. Noda, M. & Skvortzov B. V. 1 969. New and rare desmids from Genho River of Great Chingan Mountains, autonomous region of Inner Mongolia, China. Science Reports ofNiigata University D, Biology 6: 65-86 [March]. 390. Skvortzov B. V. & Noda, M. 1969. On species of genus Entosiphon Stein, Fam. Peranemaceae, Euglenaceae from Sao Paulo, Brasil (2). Science Reports ofNiigata University D, Biology 6: 87-92 [March]. (Abstract in Botanical Magazine, Tokyo 83 (979): 39, 1970). 391. & 1969. On species of genus Cyclidiopsis Korsch. (Euglenaceae) from South America. Science Reports ofNiigata Uni- versity D, Biology 6: 93-95 [March]. (Abstract in Botanical Magazine, Tokyo 83 (979): 39, 1970). 392. & 1969. On new colourless Flagellata with refractile granules and cytoprokt of Class Chloromonadaceae from Subtropics of Brasil, South America. Science Reports of Niigata University D, Biology 6: 97-100 [March]. (Abstract in Botanical Magazine, Tokyo 83 (979): 39, 1970). 393. & 1969. On new genera of Gomesiamonadaceae fam. nov. (Pyrrophyta) from Brasil, South America. Science Reports ofNiigata University D, Biology 6: 101-105. (Abstract in Botanical Magazine, Tokyo 83 (979): 39, 1970). 394. & 1 969. Xanthophyta e America Borealis et Australis novi et minus cognitae. Journal of Japanese Botany 44 (4): 106-110 [April]. 395. & 1969. Xanthophyta novae et minus cognitae e Japoniae et Brasiliae. Journal of Japanese Botany 44 (7): 218-222 [July]. 396. 1970. Notes and papers printed during 1918-1966 on wild and useful plants, floras and vegetations of northern Manchuria eastern Mongolia and south China and mostly used separately. Bulletin of Science 1: 3-7. 397. Skvortzov B. V., Bicudo, C.E. de M. & Bicudo, R.M.T. 1970. First report of the occurence of Chloromonadophyceae in Brazil. Rickia 4: 93-98. 398. Bicudo, C. E. de M. & Skvortzov B. V. 1970. Contributions to the knowledge of Brazilian Dinophyceae - Free-living unarmoured gen- era Rickia 5: 1-22. 399. *Skvortzov B. V. & Noda, M. 1970. New and little-known varieties of Pinnularia viridis (Nitz.) Ehrenb. in Eastern and Arctic Europe and Asia. Science Reports of Niigata University D, Biology 7: 37^7. (Abstract in Botanical Magazine, Tokyo 83 (989): 391, 1970). 400. & 1970. On new species of Oedognium Link from Amur River district Eastern Siberia and from Northern Manchuria China. Science Reports ofNiigata University D, Biology 7: 49-55. (Abstract in Botanical Magazine, Tokyo 83 (979): 392, 1970). 401. & 1970. On species of genus Entosiphon Stein Fam. Peranemaceae Euglenophyta from Sao Paulo Brasil South America (3) Science Reports ofNiigata University D, Biology 7: 57-61 . (Abstract in Botanical Magazine, Tokyo 83 (979): 392, 1970). 402. * 1 97 1 . An algological note. On some new fresh-water diatoms from Soochow, Prov. Kiangsu, Middle China with 19 figures. Quar- terly Journal of Taiwan Museum 24 (1-2): 59-65 [June 1971]. 403. * 1971. Diatoms from Yenisei River and its tributaries, middle part of Siberia, Western Asia. Philippine Journal of Science 98 ( 1 ): 57- 113 [March 1969, published 14 July 1971]. 404. * & Noda, Mitsuzo. 1971. On Recent and Fossil fresh-water diatoms from Japan I. Science Reports ofNiigata University D, Biol- ogy 8: 1-11 [March]. (Abstract in Botanical Magazine, Tokyo 84 (993): 194, 1970). 405. * & 1971 . On Recent and fossil fresh-water diatoms from Japan II. Science Reports ofNiigata University D, Biology 8: 13-27 [March]. (Abstract in Botanical Magazine, Tokyo 84 (993): 194, 1970). 406. * & 1 97 1 . On Recent fresh-water and fossil diatoms floras of Korea. Science Reports ofNiigata University D, Biology 8: 29-34 [March]. 407. & 1 971. Colourless algae of the flagellate genus Monosiga from Victoria, Australia. Muelleria 2 (2): 147-148 [August]. 408. 1972. A study of a green unicellular alga of genus Botrydium Wallroth, Xanthophyta found in Subtropic of Brasil, South America. Quarterly Journal of Taiwan Museum 25 (1 & 2): 69-84 [June]. 409. & Noda, M. 1 972. On some species of genus Bodo (Her.) Alex. (Bodonaceae, Protomastigineae) from Brasil, Singapore and Japan ( 1 ). Science Reports ofNiigata University D, Biology 9: 17-24 [March]. 410. & 1972. On some species of genus Bodo (Her.) Alex. (Bodonaceae, Protomastigineae) from Brasil, Singapore and Japan (2). Science Reports ofNiigata University D, Biology 9: 25-30 [March]. 411. & 1972. On some species of genus Bodo (Her.) Alex. (Bodonaceae, Protomastigineae) from Brasil, Singapore and Japan (3). Science Reports ofNiigata University D, Biology 9: 31-35 [March]. 412. & 1972. On colourless flagellata of genus Monosiga S. Kent (Crasdedomonadaceae, Protomastigineae) from Brasil, Australia and Hong Kong. Science Reports ofNiigata University D, Biology 9: 37^3 [March]. 413. & 1972. On species of genus Monosiga S. Kent (Crasdedomonadaceae, Protomastigineae) from Japan, South Africa, Sweden and Brasil. Science Reports ofNiigata University D, Biology 9: 45^8 [March]. 414. & 1 973. On new species ofZygnema Agardh (Zygnemaceae) from Altai district of Siberia and N.-E. China (Manchuria). Science Reports ofNiigata University D, Biology 10: 37^0 [March]. 415. & 1973. On new species of Spirogyra Link from Western Siberia, Eastern Mongolia, Southern China and Manchuria. Science Reports ofNiigata University D, Biology 10: 41^*7 [March]. 416. & 1973. Notes on some species of genus Cercobodo Krassil. recorded in Japan in 1968-69. Science Reports of Niigata University D, Biology 10: 49-51 [March]. 417. 1974. On some colourless flagellates from Java and Brasil. Reinwardtia9(l): 177-182 [Published 31 December]. 418. 1974. Notes and papers printed from 1917-1970 (Bibliography). Quarterly Journal of Taiwan Museum 27 (3-4): 389^17 [December 1974]. 419. & Noda, Mitsuzo. 1974. Flagellates of clean and polluted waters II. New taxa of colourless flagellates with one flagellum - genus Conradinema gen. nov. and genus Peranemopsis Lackey (Peranemaceae, Euglenophyta). Science Reports ofNiigata University D, Biology 11: 29-63 [March]. 420. * 1975. Subaerial diatom flora from Hongkong, eastern Asia. Quarterly Journal of Taiwan Museum 28 (3 & 4): 401-430 [December 1975]. 421. & Noda, M. 1975. Flagellates of clean and polluted waters III. 106 D.M. WILLIAMS & G. REID New taxa of colourless species of genus Euglena Ehrenb. without a stigma. Science Reports of Niigata University D, Biology 12: 9-3 1 [March]. 422. 1975. Flagellates of clean and polluted waters IV. New taxa of colourless species of genus Euglena Ehrenb. with an eyespot. Prof. Mitsuzo Noda Commemorative Publication on his retirement from the Faculty of Science, Niigata University: 1-23. 423. 1975. Flagellates of clean and polluted waters V. A short descrip- tion of colourless Flagellates with one swimming flagellum - genus Mastigella (Frenzel) nov. (Rhizomastigaceae) reported during 1962- 1975 in many parts. Prof. Mitsuzo Noda Commemorative Publication on his retirement from the Faculty of Science, Niigata University: 25- 66. 424. 1975. Flagellates of clean and polluted waters VI. A short description of species of genus Rhizaspis Skuja of Rhizaspidaceae (Euglenophyta) found in subtropics of Brasil and also reported from Europe and South As. Prof. Mitsuzo Noda Commemorative Publica- tion on his retirement from the Faculty of Science, Niigata University: 67-75. 425. 1975. Flagellates of clean and polluted waters VII. A short description of unusual types of Flagellate-like organisms reported from Brasil, Hong Kong, Japan and in South Africa of Rhizaspidaceae of 4 new genera. Prof. Mitsuzo Noda Commemora- tive Publication on his retirement from the Faculty of Science, Niigata University: 77-107. 426. * 1976. Moss diatoms flora from River Can in the Northern part of Great Khingan Mountains, Inner Mongolia, China, with de- scription of a new genera [sic] Porosularia gen. nov. from Inner Mongolia. Northern Mahchuria [sic] and Southern China. First part. Quarterly Journal of Taiwan Museum 29 (1 & 2): 111-152 [June 1976]. 427. * 1976. Moss diatoms flora from River Can in the Northern part of Great Khingan Mts, China, with description of a new genera [sic] Porosularia gen. nov. from Northern and Southern China. The second part. Quarterly Journal of Taiwan Museum 29 (3-4): 397-439 [December 1976]. 428. 1 977. The flagellates of clear and polluted waters on new taxa of genus Cercobodo Krasskil., A colourless flagellata of Rhizomasti- gaceae, Pantostomatineae. Quarterly Journal of Taiwan Museum 30 (1-2): 89-121 [June 1977]. 429. & Noda, M. 1 98 1 . On new genera of green flagellates - genus Akiyamae gen. nov. and genus Tsumuraia gen. nov. (Chlamydo- monadaceae) recorded in north-eastern China (Manchuria). Seibutsu 4: 15-18. 430. & 1983. On new colourless flagellates like organisms of Protostasiaceae fam. nov. (Euglenophyzeae) recorded from Japan and Brasil. Bull. Niigita Coll. Pharm. 3: 45-52. 43 1 . & 1 983. New genera of green flagellates of Euglenophyta from N.-E. China and Brasil. Niigita Coll. Pharm. Seibutsu 6: 1-6. 432. & 1 983. On Euglena species with ovoid slightly metabolic cells from subtropics of Brazil. 3:45-52. Niigita Coll. Pharm. Seibutsu 6:7-11. 433. & 1985. On species of colourless flagellates of genus Codonosiga F. Stein found in mountain waters in subtropical zone of Brasil, South America. Seibutsu 7: 23-32. 434. & 1985. On species of colourless flagellates of genus Desmarella W.S. Kent (Craspedomonadaceae, Protomastiginiae) from Brazil and Japan. Seibutsu 7: 33-38. REFERENCES Anon. 1936. Ekspedisn 1935 goda v Man'chzhurii [The expeditions of the year 1935 about Manchuria]. Naturalist Man'chshurii (Naturalist of Manchuria) 1: 19. Baranov, A.I. 1 959. Russkie kraevedy v Man'chzhurii. (B. V. Skvortzov I ego nauchnaia rabota). [Russian regional specialists in Manchuria (B.V. Skvortzov and his scien- tific work). Sbornik 'Kharhinskie Kommercheskie Uchilischa ' [Collection of works 'Harbin Commercial School'] Fasc. 6: 47-49. San Francisco, California (Mimeographed). Chin, T.G. 1951. An annotated bibliography of Chinese diatoms from 1847 to 1946. Lingnam Science Journal 23: 151-158. Chung, H.H. 1929. The study of botany in Fukien. Lingnam Science Journal!: 121- 130. Flower, R.J & Williams D.M. 1999. The Natural History Museum 1999 meeting on the biodiversity of Lake Baikal and a workshop on the taxonomy of Lake Baikal diatoms: the combined reports. ECRC Report 62. Gee, N.G. 1926. A list of some Soochow and Ningpo diatoms. Lingnaam Agricultural Review 3: 151-155. Kociolek, J.P. & Spaulding, S.A. 1999. Freshwater diatom biogeography. Nova Hedwigia 71: 223-241. & Stoermer, E.F. 1988. Taxonomy and systematic position of the Gomphoneis quadripunctata species complex. Diatom Research 3: 95-108. Licent, E. 1936. Vingt deux annees d'exploration dans le Nord de la Chine, en Mandchourie, en Mongolie et au Bas-Tibet. (19141935). Publications du Musee Hoango ho-Pai ho Tien-Tsin 39: 1-41. Mann, D.G. 1999. The species concept in diatoms Phycologia 38: 437^95. Meyer, K. 1930. Einfuhrung in die Algenflora des Baicalsees. Byulleten ' Moskovskogo obshchestva ispytatelei prirody, odtel biologicheskii 39: 1 79-396. Noda, M. 1960. Botanical history in the north-east province of China (Manchuria). Journal of the Faculty of Science Niigata University II, Biology, Geology and Mineralogy 4(1): 15-25. 1971. Botanical history in the north-east province of China (Manchuria). Flora of the N.-E. Province (Manchuria) of China. Niigata. Patterson, D.J. 1994. Protozoa: Evolution and Systematics in K. Hausmann & N. Htilsmann (Eds) Progress in Protozoology, Proceedings of the IX International Congress of Protozoology, 1993: 1-14. Berlin. Pavlov, P.A. 1925. Sungagijskaa rechnaa biologicheskaa stancia [The Sungari River Biological Station]. Obshchestvo Izucheniya Man'chzhurskogo Kraya (Manchurian Research Society, Proceedings of the Sungari River Biological Station)E. 1 (1): 1-14. Sedelnikoff, A.N. 1905. Zaisen-see. Omsk. Selling, O.K. 1962. Robert Wilhelm Kolbe (10. Januar 1882 - 9. Marz 1960). Nova Hedwigia 4: 275-298. Skabichevskij, A.P. 1974. On the true authorship of taxons described in the work of B.W. Skvortzov and C.I. Meyer 'A contribution to the diatoms of Baikal lake'. Proceedings of the Sunraree River Biological Station, 1928, Vol. 1, No. 5. Byulleten ' Moskovskogo obshchestva ispytatelei prirody, odtel biologicheskii 79: 152-156. Sowerby, A. de C. 1930. The Naturalist in Manchuria. The cold-blooded vertebrates, invertebrates and flora of the Manchurian region, IV & V. Tientsin. Uedo, H. 1940. A catalogue of freshwater diatoms of Manchoukuo cited from Skvortzov's papers. Report of the Limnobiological Survey of Kwantung and Manchoukuo. Ueyama, S. & Kobayashi, H. 1983. Fossil diatoms from Kanomura (Wamura), Nagano Prefecture in comparsion with the Skvortzov's investigations using the same material. Bulletin Tokyo Gakugei University 4, 35: 71-94. Voigt, M. 1937. Fossil diatoms of China. The China Journal 27: 310-315. Williams, D.M., Huxley, R. & Ross, R. 1998. Unravelling Ehrenberg's names: applying the past to the present. In D.M. Williams & R. Huxley (Eds), Christian Gottfried Ehrenberg (1795-1876). The Linnean Society of London, Special Issue 1: 49-62. London. Young, C.C. 1936. On the cenozoic geology of Hu, Changho and Linchii districts (Shantung). Bulletin of the Geological Society of China 15: 171-187. 1937. New vertebrate horizons in China. Bulletin of the Geological Society of China 17: 269-288. Bull. nut. Hist, Mits. Land. (Bot.) 31(2): 107-1 17 Issued 29 November 2001 On the identity of Pleurosigma angulatum (Bacillariophyta) and related species GERALDINE REID Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD CONTENTS Introduction Materials & methods Diatom morphology Analysis Results Discussion Taxonomic treatment 1 . Pleurosigma angulatum (J.T. Quekett) W. Sm. 2. Pleurosigma quadratum W. Sm 3. Pleurosigma aestuarii (Breb.) W. Sm 4. Pleurosigma mamoranqi G. Reid sp. nov 5. Pleurosigma lysekilii G. Reid sp. nov. 6. Pleurosigma malmoensis G. Reid noni. nov. ... 7. Pleurosigma stidolphii Sterrenburg References ... 107 108 108 108 109 110 111 111 111 114 114 114 115 116 117 SYNOPSIS. The specific status of Pleurosigma angulatum (J.T. Quekett) W. Sm. is clarified. P. angulatum, P. quadratum W. Sm. and P. aestuarii (Breb.) W. Sm. are shown to be separate taxa. Recent research has subsumed the three taxa under the epithet angulatum on the basis that they may occur together and so may represent different stages in the life cycle of one species. The results of this study show this is clearly not the case. Two new species of Pleurosigma are described; P. mamoranqi sp. nov. from New Zealand and P. lysekilii sp. nov. from Sweden. One new name, P. malmoensis nom. nov., is given to the taxon previously called Pleurosigma minutum (Grun.) Cl. INTRODUCTION The genus Pleurosigma W. Sm. is a member of the family Pleurosigmataceae. Pleurosigma is a readily recognized genus owing to its large size, sigmoid outline or sigmoid raphe, and its arrange- ment of transverse and oblique striae. It has two or four ribbon-like plastids which undulate throughout the cell. Initially, species of Pleurosigma were distinguished largely on the basis of their shape (Kiitzing, 1833; Ehrenberg, 1838; Rabenhorst, 1853). Smith ( 1 852) used the new name Pleurosigma W. Sm. for sigmoid naviculoid diatoms; his reason for rejecting the earlier name Gyrosigma Hassall, which was in his view synonymous with Pleurosigma, was that it was an 'alliterative blunder' (Smith, 1856: 97). Smith (1852) introduced additional taxonomic criteria in this group of diatoms by separating species into two 'sections' on the basis of the arrangement of the striae. He also provided details of the stria density for each species and used this as the basis for discrimi- nating between them. Smith's (1852) two sections reflected the modern concepts of Gyrosigma and Pleurosigma, his first section being described as having 'Beads alternate, striae oblique', i.e. Pleurosigma (Round et al., 1990), whereas his second section had 'Beads opposite, striae transverse and longitudinal', i.e. Gyrosigma (Round etal., 1990). Peragallo (1891) investigated the angle of intersection of the striae in taxa of the family Pleurosigmataceae. By combining the angle of intersection and fineness of the striae, he was able to discriminate additional species and divide them into 11 groups; Pleurosigma was still included within Gyrosigma at this point. Cleve ( 1 894) later separated Gyrosigma from Pleurosigma using the orientation of the striae and assessed species limits within the genera by the shape and path of the raphe. Pleurosigma angulatum (J.T. Quekett) W. Sm. was conserved as the type of the genus Pleurosigma by Ross (during the 8th Inter- national Botanical Congress, Paris, 1954). P. angulatum has been treated in a variety of ways encompassing a large number of varietal forms (16 listed in Reid, 1998). It was first described by Quekett ( 1 848:438) as Navicula angulata from 'the Humber at Hull' . Recent work by Sterrenburg ( 1 99 1 a) and Ross & Sterrenburg ( 1 996) argued that P. angulatum be extended to encompass P. angulatum, P. quadratum W. Sm. and P. aestuarii (Breb.) W. Sm. This was based on the assumption that they are all part of the same species, just exhibitions of different stages in the 'vegetative cycle of a single organism; quadratum as early stage, angulatum representing mid- range and aestuarii as late stage' (Sterrenburg, 1 99 la: 564). Searches for Quekett's original material failed to locate any specimens and resulted in the proposal to conserve the name P. angulatum (J.T. Quekett) W. Sm. with a conserved type slide from the Wm. Smith collection (BM 23671) (Ross & Sterrenburg, 1996). In a detailed monographic study of the genus Pleurosigma, Reid The Natural History Museum, 2001 108 (1998) disagreed with Sterrenburg's findings. Her results showed that P. angulatum, P. quadratum and P. aestuarii should be recog- nized as separate taxa. This study extends Reid's (1998) findings using cladistic analyses to explore the relationships of the three taxa and their close relatives as indicated from Reid's results. MATERIALS & METHODS Slides were examined using a Reichert microscope. For photo- microscopy, specimens were examined on a Carl Zeiss microscope, with differential interference contrast, using plan apochromat objec- tives and tri colour green filter. Photos were taken using medium format Kodak Tmax 100 film. Cleaned specimens were strewn on aluminium stubs for SEM examination. The specimens were sputter coated with platinum and observed using a Hitachi S800 field electron microscope. All slides, negatives and SEM stubs of the specimens are housed in the herbarium of The Natural History Museum, London (BM). Diatom morphology Terminology for the siliceous parts of the diatom frustule follows that of Anon. (1975) and Ross et al. (1979). Characters that appeared constant within species were investi- gated using cladistic analysis. Character stability was established by investigating about ten populations from each taxon (examining more than 1 00 individuals from each population) for infra- and inter- specific variation. Individuals were cultured from each population and subjected to variations in temperature (9-30C), salinity (7^3 ppt) and light (8^45 umol m 2 sec~' ) to assess character stability under a multifactorial design. All characters were measured in both the 'wild' and the 'cultured' material. Only qualitative characters were used in the cladistic analysis. Quantitative characters were not used due to the inherent problems of coding continuous characters into discrete states for analysis (Pimentel & Riggins, 1987; Thiele, 1993; Reid & Sidwell, 2001). Many characters traditionally associated with diatom valve mor- phology are quantitative (e.g. length, breadth, stria ratios) and only used in species descriptions. The only characteristics of shape that can be used qualitatively are first, the broad definition of cells as arcuate (as in the outgroup Toxonidea Donkin, compared to sigmoid cells as in the remaining Pleurosigmataceae), and second, shape in girdle view (oblong in Pleurosigma and panduriform in Donkinid). Within the family Pleurosigmataceae the striae are arranged in one of two ways; transverse and oblique rows in Pleurosigma, Toxonidea and Donkinia, longitudinal and transverse rows in Gyrosigma and Donkinia. This character was first used to separate Gyrosigma and Pleurosigma into two distinct genera by Cleve ( 1 894). The areolae in the Pleurosigmataceae are loculate and occluded by a rica. The areolae open externally by a slit, the foramen. The internal areola opening in the Pleurosigmataceae can be divided into two types; either a single opening which is found in Gyrosigma, Toxonidea, some species of Pleurosigma (Fig. 3c) and Donkinia, or areolae divided by a siliceous bar, as found in some species of Pleurosigma and Donkinia (Fig. 5c). In some species of Pleurosigma the striae change orientation near the apex of the valve from the transverse/ oblique pattern to the transverse/longitudinal pattern (Fig. 3f)- This change was first noted and clearly drawn and discussed by Anthony (1870). Sterrenburg attributed this discovery to Hendey (1964), commenting that 'this had never been described or drawn in the previous century of observations' (Sterrenburg, 1991a:561). This is not the case, as it had been commented on frequently in the literature G. REID prior to Hendey, for example Brown (1914:327) regarded this as 'a good specific character', and Woodward ( 1 87 1 : 160) made detailed photographic illustrations of the character. The external central raphe fissures of Pleurosigma were first discussed as a taxonomic character by Cardinal et al. (1989). These are continuations of the external raphe slit opening, extending onto or across the central nodule, but not penetrating the valve. This character refers to the orientation of the central external raphe fissure endings (see Fig. 2g). Cardinal et al. (1989) used six character states in their study, only four of these are seen in the species under investigation here: either the central fissures are curved in opposite directions, similar directions, one centered and the other deflected, or the central fissures are overlapping. Central bars are siliceous thickenings deposited either side of the internal central nodule (Fig. 2f). Reid (1998) observed eight character states in her study, four of which are observed in the species under investigation here: central bars smooth and slender, with an indis- tinct hazy edge, with siliceous lateral extensions or thick and raised. A hyaline area may occur adjacent to these bars (this is an area which is not penetrated by areolae) and may follow the shape of these bars (Fig. 3h) or be transversely expanded (Fig. 5c) or absent (Fig. 2f). In some species of Pleurosigma the internal central area is raised to form a 'saddled' humped area. The internal central nodule may have an extra siliceous deposit in the centre in some species of Pleurosigma (Fig. 5c), a character referred to as 'central nodule raised' in the cladistic analysis. Analysis Table 1 lists the characters and the character states used in the analysis; character coding is presented in Table 2. The data set Table 1 Characters and character codes used in the cladistic analysis Character Character Description Character code Valve arcuate Valve sigmoid 1 1 Striae transverse and oblique Striae transverse and longitudinal 1 2 Raphe arcuate Raphe sigmoid 1 3 Areolae undivided Areolae divided 1 4 Striae same orientation at apex Striae change orientation at the apex 1 5 Central raphe fissures: Curved in same direction Curved in opposite directions 1 One centred the other deviated 2 Overlapping 3 6 Central area not saddled Central area saddled 1 7 Central bars: Smooth and slender With indistinct hazy edge 1 With siliceous lateral extensions 2 Thick and raised 3 8 Hyaline area: transversely expanded follows the central bars 1 absent 2 9 Central nodule raised Central nodule plain 1 10 Valve flat Valve vaulted 1 11 Girdle view oblong Girdle view paduriform 1 PLEUROSIGMA ANGULATUM AND RELATED SPECIES 109 Table 2 Data matrix used in the cladistic analysis 10 Toxonidea ( ) ( ) P. angulatum 1 2 1 2 P. quadratum 1 2 1 1 P. aestuarii 1 3 2 1 P. stidolphii 3 2 1 1 P. mamoranqi 1 3 P. lysekilii 9 9 9 P. malmoensis 9 9 9 P. rhombeum 1 1 1 (i D. minutum 1 1 I 9 3 1 1 1 1 D. latum [ 1 1 9 3 1 1 1 1 contains ten binary characters and two multistate characters, which were coded with reference to Toxonidea as an all zero outgroup (Nixon & Carpenter, 1993). Pleurosigma minutum Donkin (not to be confused with Pleurosigma minutum Grun.) is synonymous with Donkinia minutum (Donkin) Ralfs. As Donkinia is seemingly closely related to Pleurosigma, two species in the genus, D. minutum and D. latum E.J. Cox, were included in the analysis. P. rhombeum Grun. was included in the analysis because this is a taxon which is often misidentified as P. quadratum. Weighting is the importance applied to characters in analysis. When different characters are assumed to contribute to the same degree to the tree topology they are all given the same weight (uniform weighting), applied initially as the default in most current parsimony programs (e.g. Hennig86 - Farris, 1988; PAUP - Swofford, 1990). If the characters are assumed to contribute to different extents to the analysis they are given different weights (differential weighting). In the past, weighting has tended to be a highly subjective process with intuition being the only basis for giving some characters more weight than others in an analysis (e.g. Mayr, 1969). Weighting may take place prior to tree construction, in which case it is termed a priori weighting (Neff, 1986), which takes into account what is known about the characters, or after initial tree construction, in which case it takes into account the contribution the characters make to the resulting tree topology, a posteriori weighting (Neff, 1986). A priori weighting is not appropriate as it is impossible to know which characters are useful before an analysis. The first serious attempt to address the problem of how to weight characters was brought about by Farris (1969), but little use was made of it until he incorporated it into his computer program (Farris, 1988), as a differential a posteriori method. The method is based on the assumption that 'characters which have failed repeatedly to adjust to the expectation of hierarchic correlation are more likely to fail again in the future, and so they are less likely to predict accurately the distribution of as yet unobserved characters' (Goloboff, 1993o). In practice this is the same as excluding some characters and introducing the non-random replication of other characters. Farris's method (Farris, 1969, 1988) is iterative in that weights are applied to the most parsimonious trees for a given set of weights, and are then used in the reanalyses. It works on the basis that because the character consistency index (c) measures homoplasy but can never reach zero, by rescaling it with the retention index (ri) (which measures the amount of synapomorphy), characters with no synapomorphy can be disregarded in future analyses. Goloboff ( 1 99 1 a) shows that Farris's method of weighting still has problems as weights do not always increase with less homoplasy. Goloboff (1993ft) introduced a non-iterative method in which the weighting is based on a concave function of homoplasy, given as fit (f): f=(k+l)/(s+k+l-m) k = a constant of concavity, s= minimum number of steps a character can have on a particular tree, m= minimum number of steps a character can have on any tree. This is implemented using the computer program Pee- Wee (Goloboff 1993ft) to analyse the data matrix. It selects trees of highest weight as being the most parsimo- nious, as opposed to the shortest total length, as under Hennig86 (Farris, 1988). Some workers only advocate the use of weighting in the analysis if the initial unweighted data has failed to produce an adequate tree due to character conflict, or to choose between multiple trees (Turner & Zandee, 1995; Rodrigo, 1992). This approach is rejected here because weighting selects characters not cladograms. Many workers do not accept the concept of differential character weight- ing, arguing that every character should be attributed equal weight to create natural taxa because all characters are 'equally desirable' (Sneath & Sokal, 1973; Mann, 1982; Round et al., 1990). I shall use differential a posteriori weighting because it results in trees which are self-consistent with the data collected (Farris 1969, 1988; Goloboff, 1993, 1995;Platnicketal., 1991), in that not all characters contribute equal amounts of information to the analysis. They are given the weight they deserve after initial analysis and thus weight- ing is required in all analysis. Parsimonious trees for the data were found using Hennig86 (Farris, 1988) with the ie* command (implicit enumeration, which is guaranteed to find all the most parsimonious trees). Initially, all characters were given equal weight and were unordered. As it is possible that not all characters contribute to the same extent to the prediction of relationships (Farris, 1983; Goloboff, 1993a), a poste- riori differential weighting was investigated (as implemented in the program Pee-Wee (Goloboff, 1 993ft)). The option mult*50 was used to search for trees of highest fit, performing random addition sequences of 50 replications each. Replication was followed by tree bisection and branch-swapping. RESULTS Both Henning86 and Pee-Wee resolved the same tree (length 19, ci 89, ri 88, fit 75 (Fig. 1 )). The tree in Fig. 1 has seven nodes (labelled 0-6, with the most basal node of the tree). Pleurosigma angulatum and P. quadratum are resolved at node 4 supported by character 5 (central raphe fissure orientation) and character 7 (central bar type). Node 2 resolves P. aestuarii as sister taxon to these two taxa supported by character 4 (striae orientation at apex) and character 8 (hyaline area type). The resolution at node 3 (P. rhombeum, P. stidolpii, D. minutum and D. latum) is attributed to character 10 (valve vaulting). The resolution at node 5 (P. stidolpii, D. minutum and D. latum) is based on character 9 (central 110 G. REID [j=Toxonidea }==?. lysekilii P. malmoensis ^= p=P. mamoranqi li=P. aestuarii =2=| rj=P. angulatum U=4=!J=P. quadratum P. rhombeum P. stidolphii minutum latum Fig. 1 Tree resolved by Hennig86 and Pee- Wee. Numbers on braches indicate the nodes Table 3 Character weights Character Fit Character Fit / 6 / 1 10 7 10 2 / 8 7.5 3 / 9 7.5 4 10 10 10 5 10 11 10 nodule type) and at node 6 (D. minutum and D. latum) based on character 1 (striae orientation), character 8 (hyaline area type) and character 1 1 (girdle view shape). Weights attributed to the characters were high suggesting that they are relatively reliable (Table 3). Six characters received the maximum weight of 10, indicating that the characters were perform- ing well and were hierarchically correlated. Two received a value of 7.5, which also indicates a high level of correlation. Characters and 2 do not receive a weight because they only serve to separate the ingroup from the outgroup. Characters 3 and 6 do not receive a weight as they are autapomorphic. DISCUSSION Sterrenburg (199 la: 561) stated that There is no morphological discontinuity between Pleurosigma quadratum W. Sm., Pleurosigma angulatum sensu W. Smith and Pleurosigma aestuarii (Breb. ex Kiitz.) W. Smith and the species description of Pleurosigma angulatum is extended to account for this continuum.' However, detailed morphological studies of large numbers of specimens has shown that this is not the case and that they are separate taxa. The taxa differ in their hyaline areas; Pleurosigma angulatum and P. aestuarii do not have one (Figs 2f & 4c) whereas in P. quadratum the hyaline area follows the central bars (Fig. 3h). The taxa differ in their central bars, with P. quadratum having extra thickening at the centre of the bar (Fig. 3h) whereas P. angulatum has a more even deposit of silica (Fig. 2f) and P. aestuarii has smooth slender bars (Fig. 4b). P. quadratum is a much larger taxon than P. angulatum with a distinctive rhomboidal shape, compared to the more lanceo- late form of the latter species. P. aestuarii is much smaller with rostrate ends. P angulatum possesses a different pattern change at its apices to P. quadratum and P. aestuarii. On the outside curve of the raphe the pattern continues approximately twice as far as on the opposite side (Figs 2h-j), whereas in P. aestuarii and P. quadratum the change in stria orientation is equal on both sides of the raphe (Figs 3b, c, f & 4c). Due to these differences they are maintained as separate taxa. The basis for Sterrenburg's ( 1 99 la) summation is that the taxa are frequently found together. In this study, examination of many differ- ent populations showed that this is not the case, with their only occasionally being found together. While in nature many taxa frequently occur together, this is no basis for assuming that they are all to be regarded as one taxon; they may just have similar environ- mental requirements. Sterrenburg (199 la: 564) considers that there is 'a biological continuum between quadratum, angulatum and aestuarii'. Yet he admits that 'some samples may be almost pure aestuarii', continuing that 'While this may suggest the taxon's individuality, it does not necessarily imply it'. Reid (unpublished data) grew populations of Pleurosigma angulatum under different environmental conditions and monitored their morphometric changes. She found no evidence to support Sterrenburg's hypoth- esis. Sterrenburg ( 1 99 la: 563) states that 'central bars were found to be highly variable in several of the "states" defined and discussed in Cardinal et al. (1989).' This found central bars to be a stable character both within a population and between populations from different sites. As Sterrenburg is subsuming the taxa under one species this may account for the variability he is observing. Sterrenburg (199 la) also sees the three taxa as part of a size range continuum. If this is the case, why is there a complete size range for each species that can clearly be seen to belong to each nominate type? Examination of large populations from different locations reveals the range in variation, but each taxon can still be clearly recognized. Sterrenburg ( 1 99 1 a) suggested that Queckett ( 1 848) had illustrated two taxa because his illustrations show one taxon with 5 1 and one with 60 angle of stria intersection. However this does not necessar- ily imply the inclusion of two different taxa, because this range of angles can be found in Pleurosigma quadratum, which exhibits angles from 50-60. This is a continuous character, which cannot be fitted into discrete character states. Sterrenburg (199 la) refutes this possibility as 'Stria angle was found to be a stable parameter in this genus in Sterrenburg 1991 and the findings then indicated that 51 deg. falls outside the range of variation for Pleurosigma angulatum sensu W. Smith.' (Sterrenburg, 1991a:561). These observations were made on only two populations, one from the Wadden Sea, Holland and one from New Zealand (Sterrenburg 19916:371) so it is quite likely that the full range of variation was not observed, giving him only 57-62. However, the results of the systematic analysis do show these taxa to be closely related. Pleurosigma angulatum and P. quadratum are resolved in both the unweighted and the weighted analyses as sister taxa (Fig. 1 ) at node 4 supported by character 5 (central raphe fissure orientation) and character 7 (central bar type). P. aestuarii is shown to be their closest relative at node 2 supported by characters 4 (striae orientation at apex) and 8 (hyaline area type). The shift in orientation at the apex was discussed by Reid (1998) and has been shown to be a stable taxon-specific character within the genus Pleurosigma. This character is a very useful aid to the identification of this complex group as it is easily visible under the light microscope and is shown by only 15 members of the genus. It does not change under environ- mental conditions or during growth. No unmounted material was available for SEM investigation of Pleurosigma lysekilii Reid and P. malmoensis Reid, therefore characters 3 (areolae type), 8 (hyaline area type), and 9 (central PLEUROSIGMA ANGULATUM AND RELATED SPECIES 111 nodule type) are scored as '?' in the cladistic analysis, leaving them unresolved within the group at node of the tree (Fig. 1). Pleurosigma mamoranqi Reid differs from P. angulatum, P. quadratum and P. stidolphii Sterrenburg in having areolae crossed by a bar. It differs from P. angulatum and P. quadratum by its striae that do not change orientation near the apex. It has a large expanded hyaline area adjacent to the central bars. It differs from P. rhombeum by not having a saddled central area. This taxon does not fit any of the previously published descriptions of Pleurosigma, and as such is described as new. In the first part of his monograph Peragallo ( 1 89 1 : 1 2) considered Pleurosigma minutum to be a valid taxon and expands its distribution from Malmo, Sweden to include the Balearic Islands. However, in the second part of his monograph, Peragallo (1891:34) considers P. minutum as a doubtful or unknown species. He says 'Je n'ai pu 1'identifier avec certitude.' (Peragallo, 1891:34). The present study shows that P. minutum is a valid taxon that can be clearly distin- guished from other species of Pleurosigma. P. minutum is quite different to Donkinia minutum which has a highly vaulted valve with its raphe on a keel. It is panduriform in girdle view whereas P. minutum is rectangular. P. minutum has transverse and oblique striae whereas D. minutum has transverse and longitudinal striae. However, as the name P. minutum had been used prior to Grunow (P. minutum Donkin, 1858:24, 3/9) a new name, P. malmoensis, is proposed for this taxon. D. minutum was not shown to be closely related to P. malmoensis, being placed in a different part of the tree, and therefore they cannot be considered as the same taxon. TAXONOMIC TREATMENT 1 . Pleurosigma angulatum (J.T. Quekett) W. Sm. in Ann. Nat. Hist. 9: 7, pi. 1 fig. 8 (1852) Type: England, Hull, Humber Estuary (Lectotype: Belfast, 1996, Ross & Sterrenburg BM 23671!). Fig. 2 Navicula angulata J.T. Quekett in A practical treatise on the use of the microscope: 438, pi. 8 fig. 4-7 (1848). Pleurosigma angulatum f. minor Rabenh., Flora Europea Algarum: 234(1864). Pleurosigma angulatum var. robustum McCall, J. Linn. Soc., Botany 49: 265, 306 (1933). Valve rhomboidal sigmoid, length 120-280 um, breadth 30-35 urn (Figs 2a-d). Raphe sigmoid, central, becoming slightly eccentric near the apices (Figs 2a-d). External central raphe fissures with one centred and the other deflected to one side (Fig. 2g). Hyaline area absent (Fig. 2e, f). Striae 20 per 10 um, crossing at an angle of 54-60. Striae change orientation at the apex, the pattern change continues about twice the distance on the outer most curve of the raphe compared with the inner side (Figs 2h-j). Areolae undivided. Central bars of approxi- mately equal length with an indistinct outer edge (Fig. 2f). Ross & Sterrenburg ( 1 996) chose to conserve BM 2367 1 as the type of the species. They record it as 'Belfast, August 1849', and state that this slide is from a locality listed in Smith ( 1 853:65) 'Poole Bay, Aug. 1848; Belfast Bay, Aug. 1849; Coast of Sussex, April, May and Aug. 1852, W. Sm. Coast of Lancashire, Mr. Johnson. Rye, Mr. Jenner. Hull, Mr. R. Harrison. Coast of Norfolk, Mr. BrightwelV . This is somewhat misleading because in his earlier work, when he originally described P. angulatum, Smith (1852) gave the locality as Belfast Bay, Liverpool not Belfast (Ireland). The choice of locality is also somewhat strange as Quekett's (1848) original description is of a taxon found 'upon conferva in the Humber at Hull'. From the list in Smith's (1852) description ('Poole Bay; Belfast Bay, Liverpool; Coast of Sussex, &c.; Coast of Lancashire, Chr. Johnson, Esq.\ Rye, Mr. Jenner\ Hull, Mr. R. Harrisonl Norfolk, Thos. Brightwell Esq.l' (Smith 1852:7)) a Hull sample would have been a more judicious choice, for example BM 1 1 809 and BM 1 1 8 10 both from Hull and collected by R. Harrison. The slide (BM 2367 1 ) has unfortunately been damaged by someone circling the specimen of P. angulatum and this has caused a perforation in the coverslip which has led to the slide drying out. The taxon described by McCall ( 1 933) as Pleurosigma angulatum var. robustum (Fig. 2d) is subsumed under the name P. angulatum as no morphological differences can be found. MATERIAL EXAMINED UNITED KINGDOM. England. Hartlepool, n.d., Arnott 570, Greville Coll. (BM 96); sine loc., n.d., Wm. Smith 2, Greville Coll. (BM 204); Hull, 1856, G.N. s.n., Greville Coll. (BM 386); Blyth Harbour, November 1857, Donkin C. No. 1, Greville Coll. (BM 389); Harwich, n.d., Anon. Deby Coll. (BM 7251); Harwich, n.d., Anon. Deby Coll. (BM 7253); Hull, n.d., R. Harrison 117, Deby Coll. (BM 11809); Hull, n.d., R. Harrison 117, Deby Coll. (BM 11810); sine loc., n.d., W.S. 205, Deby Coll. (BM 1 333 1 ); Norfolk, n.d., F. Kitton 43, Deby Coll. (BM 13703); Poole Bay, August 1849, Anon., Wm. Smith Coll. (BM 23672); Pevensey, September 1851, Anon., Wm. Smith Coll. (BM 23674); Sussex, Ilford, April 1 852, Anon., Wm. Smith Coll. (BM 23675); Sussex, Lancing, August 1852, Ano/i.,Wm. Smith Coll. (BM 23676); Sussex, Seaford, December 1852, Anon., Wm. Smith Coll. (BM 23677); Pevensey, August 1853, Anon., Wm. Smith Coll. (BM 23679); Sussex, Newhaven, February 1854, Anon., Wm. Smith Coll. (BM 23681); Sussex, Southease, August 1854, Anon., Wm. Smith Coll. (BM 23682); Sussex, Shoreham, April 1854, Anon., Wm. Smith Coll. (BM 23683); Sussex, Shoreham, April 1 854, Anon., Wm. Smith Coll. (BM 23684); Essex, Walton, n.d., Comber 3 (BM 31459); Essex, Walton, n.d., Comber 2 (BM 31458); Den Marsh, May 1896, Comber s.n., (BM 31460); Den Marsh, May 1896, Corner s.n., (BM 3 1461); Kent, Broad Water, near chalk 'S.I 57, 326', n.d., Rylands s.n., (BM 48371); River Mersey, n.d., /. Hardmach 331, Rylands Coll. (BM 48441); River Mersey, n.d., /. Hardmach 331, Rylands Coll. (BM 48442); Hull, 1857, W 103, 331, Rylands Coll. (BM 48444); Humber, Hull, n.d., GN 103, Rylands Coll. (BM 49989); Humber, Hull, n.d., GN 103, Rylands Coll. (BM 49990); Humber, Hull, n.d., GN 103, Rylands Coll. (BM 49991); Humber, Hull, n.d., GN 103, Rylands Coll. (BM 49992); Lancaster, Salt pool, '(274)383', n.d., Rylands s.n., (BM 48593). Scotland: Fife, Tayport, Tents Muir, Brackish Burn, n.d., D.R. McCall s.n. (BM 8345 1 ); Tayport, mud scraping from brackish pool on links, 23 September 1916, ex A. W. Round s.n., , J.R. Carter Coll. (BM 93648). Northern Ireland: Belfast, Queens Island, n.d., Amott 655, Greville Coll. (BM 1648); Belfast, August 1849, Anon.,Wm. Smith Coll. (BM 23671); Carrickfergus, August 1849, Anon., Wm. Smith Coll. (BM 23673). IRELAND. Galway, July 1853, s.n., Wm. Smith Coll. (BM 23678). FRANCE, sine loc. 'Diatomees de France', n.d., 33/2,3 Anon., (BM - Adams Coll.). 2. Pleurosigma quadratum W. Sm., in A synopsis of the British Diatomaceae. 1: 65 pi. 20 fig. 204 (1853). Type (designated here): England, Sussex, August l85Q,Anon., BM 23669! (Isotype localities: Poole Bay; Coast of Sussex; Devonshire; Menai Straits; Folkestone). Fig. 3 Pleurosigma angulatum W. Sm., Ann. Nat. Hist. 9: 7, pi. 1 fig. 7 & 9(1852). Pleurosigma angulatum var. quadratum (W. Sm.) Van Heurck, Synopsis Diatomees Belgique: 115 (1885). Valves strongly rhombic, sigmoid, length 130-285 jam, breadth 50- 65 um (Fig. 3a, d). Raphe sigmoid eccentric towards the apices (Fig. 3a, d). Areolae undivided (Fig. 3c), transverse striae 19 per 10 um, Fig. 2 Pleurosigma angulatum. (a) BM 23671; (b) BM 23674; (c) BM 386; (d) P. angulatum var. robmtum (BM 83451); (e) light micrograph of central area (BM 23671); (f) SEM of internal central area, arrows indicating central bars; (g) SEM of external central fissures (arrows), one centred the other deflected; (h) light micrograph of valve apex showing change in striae orientation, arrows indicating the different distance of the striae change on either side of the raphe (BM 23674); (i) SEM of internal valve apex; (j) SEM of external valve apex. SEM micrographs from material of Wm. Smith Herbarium, Anon., Sussex, Lancing, August 1852. (Scale bars: a-e, h & j = 10 urn, f & i = 5 urn; g = 1 urn). PLEUROSIGMA ANGULATUM AND RELATED SPECIES 113 Fig. 3 Pleurosigma quadratum. (a) BM 23669; (b) SEM of external apex; (c) SEM of internal apex; (d) BM 23669; (e) SEM of external central area showing central raphe fissures; (f) light micrograph of valve apex showing change in striae orientation (BM 23669); (g) light micrograph of central area (BM 23669); (h) SEM of internal central area, arrow indicating hyaline area. SEM micrographs of Wm. Smith Herbarium material, Anon., Sussex, August 1850. (Scale bars: a, d, f, g = 10 urn; b, c, e, h = 5 urn). 114 G. REID oblique striae 18 per 10 um, intersecting at an angle of 50-60, changing orientation near the apex (Fig. 3b, c, f). External central raphe fissures one deviating to one side the other centred (Fig. 3e). Central bars of approximately equal length, thickened at the centre with an indistinct outer edge (Fig. 3g, h). Hyaline area follows central bars (Fig. 3h) in contrast to P. angulatum (Fig. 2f). MATERIAL EXAMINED UNITED KINGDOM. England: Harwich, n.d., Arnott s.n., Greville Coll. (BM 316); W. Smith 204, Greville Coll. (BM 219); Devon, Saltmarsh, n.d., Fig. 4 Pleurosigma aestuarii (a) SEM of valve; (b) SEM of internal central area; (c) SEM of internal valve apex, showing change in striae orientation; (d) SEM of external central area. Material from letter of de Brebisson 27 February 1849 to Walker Arnott. (Scale bars: a = 10 um; b, c = 2 urn; d = Sum). Gregory s.n., Greville Coll. (BM 2 129); Harwich, n.d., A. Gr. 814, Deby Coll. (BM 7108); Poole Bay, June 1 1 1849, Smith 204, Deby Coll. (BM 13320); Folkstone, November 1 852, J.R. Capron, Smith 204, Deby Coll. (BM 1 332 1 ); Norfolk, n.d., F. Kitton 43, Deby Coll. (BM 13703); test slide, sine loc., n.d., Anon., Roper Coll. (BM 20751); Sussex, August 1850, Anon., Wm. Smith Coll. (BM 23669); Hull, n.d., Dr M 330/1 , Rylands Coll. (BM 48384); Hull, n.d., Dr Ivl 330/2 Rylands Coll. (BM 48385); Northfleet (S. 750) 330, n.d., Anon., Rylands Coll. (BM 88383); Harwich, Walton Ferry, n.d., Anon. 814, Rylands Coll. (BM 50436). 3. Pleurosigma aestuarii (Breb.) W. Sm. in A Synopsis of the British Diatomaceae. 1: 65, pi. 31 fig. 275 (1853). Type: France, St Vaast, n.d., de Brebisson s.n., BM 100839! Fig. 4 Navicula aestuarii Breb. in Kiitzing, Species Algarum: 890 (1849). Gyrosigma aestuarii (Breb.) Griffith et Henfrey in The Micro- graphic Dictionary: 302, pi. 11 fig. 35 (1856). Valves lanceolate, slightly sigmoid, apices rostrate (Fig. 4a). Length 60-90 urn, breadth 12-20 urn. Raphe central becoming eccentric towards the apices, strongly sigmoid. Internal central bars smooth and of equal length (Fig. 4b). Hyaline area absent from besides central bars (Fig. 4b). External central raphe fissures overlapping (Fig. 4d). Areolae not divided by a silica bar (Fig. 4b). Striae change orientation at the apex (Fig. 4c). Transverse and oblique striae 1 8 per 10 urn intersecting at 60. MATERIAL EXAMINED FRANCE. St Vaast: n.d., Kiitzing 1567 (BM 18866); n.d., Kiitzing 1729 (BM 18867); n.d., de Brebisson s.n., (BM 100839); n.d., de Brebisson s.n., (BM 100840). 4. Pleurosigma mamoranqi G. Reid sp. nov. Type: New Zealand, South Island, Mamoranqi Bay, n.d., S.R. Stidolph s.n., BM8 1613! Fig. 5 Valva rhombicus, sigmoideo, 100-120 um longa, 15-19 urn lata. Raphe valde sigmoidea. Striae transapicales 26 per 10 um, striae obliquae 16 per 10 um. Poris habens transtrum. Transtra centralia laeves, in longitudine aequales, ovalis area hyalina. Valves rhomboidal, sigmoid, length 100-120 um, breadth 15-19 urn (Fig. 5a, h). Raphe strongly sigmoid and eccentric towards the apices (Fig. 5a, h). Areolae with a bar across them (Fig. 5c, e), transverse striae 26 per 10 um, oblique striae 16 per 10 um, intersect- ing at an angle of 56, striae do not change orientation at the apex (Fig. 5 d-g). Internal areolae of a different shape around the helictoglossa (Fig. 5e). External central raphe fissures overlapping (Fig. 5b, g). Internal central bars smooth, approximately equal in length (Fig. 5c). Hyaline area adjacent to the central bars oval (Fig. 5c). MATERIAL EXAMINED NEW ZEALAND. South Island. Mamoranqi Bay, n.d., S.R. Stidolph, BM81613. 5. Pleurosigma lysekilii G. Reid sp. nov. Type: Sweden, Lysekil, n.d., P.T. Cleve s.n., BM12896! (Cleve & Moller no. 142; Deby collection). Fig. 6 Pleurosigma angulatum f. minor Cleve in Cleve and Moller, Dia- toms (exsiccata) III: 4, no. 142 (1878) nom. nud. Valva lanceolatus ambitu ita leviter sigmoideo. Raphe centralis leviter sigmoidea. 160-262 um longa, 20-30 urn lata. Striae transapicales 17-19 per 10 um, striae obliquae 18-20 per 10 um. PLEUROSIGMA ANGULATUM AND RELATED SPECIES Fig. 5 Pleurosigma mamoranqi. (a) BM 81613; (b) SEM of external central area; (c) SEM internal central area, arrow indicating extra siliceous deposit in middle of central nodule; (d) SEM of external valve apex; (e) SEM of internal valve apex; (f) light microgaph of valve apex (BM 81613); (g) light micrograph of central area showing external central raphe fissures (BM 81613); (h) BM 81613. SEM micrographs from New Zealand, Momoranqi Bay, ex Stidolph 155. (Scale bars: a, f-h = 10 um; b, c = 5 jam; e = lum). Valves lanceolate, only slightly sigmoid at the apices, subacute (Fig. 6a). Raphe central sigmoid at the apices (Fig. 6a). Length 160- 262 jam, breadth 20-30 jam. Transverse striae 17-19 per 10 (am, oblique striae 18-20 per 10 (am, crossing at 5 8-61. No change in the orientation of the striae at the apex (Fig. 6b) in contrast to P. angulatum. Central area small (Fig. 6b). The name Pleurosigma angulatum f. minor Cleve and Moller cannot be applied to this taxon as it has prior use, Pleurosigma angulatum f. minor Rabenh. (1864). The taxon is not a form of angulatum in that it does not possess striae that change orientation at the apex. MATERIAL EXAMINED SWEDEN. Lysekil, n.d., P.T. Cleve s.n., Cleve & Moller no. 142, Deby Coll., (BM 12896); 'W.Goteborg, Bahus', n.d., P.T. Cleve s.n., Cleve & Moller no. 142, Wynne Baxter Coll.5342 (BM 59695). 6. Pleurosigma malmoensis G. Reid nom. nov. Type: Sweden, Malmo, n.d., M.O. Nordstedt s.n., BM 12902! (Cleve & Moller no. 136, Deby Collection). Fig. 7 Pleurosigma aestuarii var. minutum Grunow in Cleve and Moller Diatoms (exsiccata) III: 3, no 136 (1878). 116 G. REID Fig. 6 Pleurosigma lysekilii. (a) BM 12896; (b) valve apex; (c) central area. (Scale bars: a-c = 10 um). Pleurosigma minutum (Grunow) Cleve, Kong. Sven. Vet. Hand. 26: 41 (1894). Pleurosigma angulatum f. minutum (Grunow) De Toni, Syll. Alg.: 232(1891). Pleurosigma angulatum var. minutum (Grunow) McCall, J. Linn. Soc. Lon. Bot. 49(328): 266 (1933). Valves sigmoid, lanceolate, length 50-75 um, breadth 12-15 urn (Fig. 7). Raphe central, sigmoid (Fig. 7). Transverse striae 26-27 per lOum, oblique striae 28-29 per 10 um, crossing at an angle of 60. Central area small with smooth central bars of approximately equal length, there is no hyaline area adjacent to the central bars. External central fissures curved in the same directions (Fig. 7a). Striae do not change orientation at the apex (Fig. 7). MATERIAL EXAMINED SWEDEN. Malmo: n.d., M.O. Nordstedt s.n., Cleve & Moller. no. 136, Deby Coll. (BM 12902); n.d., M.O. Nordstedt s.n., Cleve & Moller no. 136 (BM - Adams Coll.). 7. Pleurosigma stidolphii Sterrenburg, Bot. Mar. 34: 568, figs 38- 46 (1991). Type: New Zealand, Otago Harbour, October 1961, A.J. Doig s.n., BM8 1608! Fig. 8 Valves very rhombic, sigmoid, tapering to acute apices. 100-270 um long; 30-60 um wide (Fig. 8a). Valve vaulted with raphe situated on a ridge (Fig. 8a). Internal central bars with siliceous lateral exten- sions. Central area small (Fig. 8c). Oblique striae 22-25 per 10 um; Fig. 7 Pleurosigma malmoensis BM 12902. (Scale bars: a-b = 10 jam). Fig. 8 Pleurosigma stidolphii BM 8 1 608. (Scale bars: a-c = PLEUROS1GMA ANGULATUM AND RELATED SPECIES 117 transverse striae 23-27 per 10 (am. Striae do not change orientation at the apex (Fig. 8b). Areolae undivided. External central raphe fissures overlapping. MATERIAL EXAMINED NEW ZEALAND. Otago Harbour, October 1961, AJ. Doig s.n., leg. S.R. Stidolph(BM81608). ACKNOWLEDGMENTS. I would like to thank Dave Williams and Eileen Cox for their help and comments on the paper and Peter York and Nick Hayes for help with photomicroscopy. REFERENCES Anon. 1975. Proposals for a standardization of diatom terminology and diagnoses. Beiheft Nova Hedwigia 53: 323-354. Anthony, J. 1870. On the structure of Pleurosigma angulatum and Pleurosigma quadratum. Monthly Microscopical Journal 4: 121-123. Brown, N.E. 1914. Some notes on the structure of diatoms. Journal of Quekett Microscopical Club, Series 2, 12(74): 317-338. Cardinal, A.C., Poulin, M. & Berard-Therriault, L. 1 989. New criteria for species characterization in the genera Donkinia, Gyrosigma, and Pleurosigma (Naviculaceae, Bacillariophyceae). Phycologia 28: 15-27. Cleve, P.T. 1894. Synopsis of naviculoid diatoms. Kongliga Svenska Vetenskaps- Akademiens Handlingar 26-27: 1-194, 1-219. Donkin, A.S. 1 858. On the marine Diatomaceae of Northumberland, with a description of eighteen new species. Transactions of the Microscopical Society 5: 12-34. Ehrenberg, C.G. 1838. Die Infusionsthierchen als vollkommende Organismen. Ein Blick in das tiefere organische Leben der Natur. Leipzig. Farris, J.S. 1969. A successive approximation approach to character weighting. Systematic Zoology 18: 374-385. 1983. The logical basis of phylogenetic analysis. 7-36. In: N. Platnick & V. Funk (Eds) Advances in Cladistics 2. Proceedings of the Second Willi Hennig Society. New York. 1988. Hennig86 version 1.5 manual; software and MSDOS program. New York. Goloboff, P.A. 1993a. Estimating character weights during tree search. Cladistics 9: 83-91. 1993b. PeeWee. MS-DOS program. Published by the author. 1995. Parsimony and weighting: a reply to Turner and Zandee. Cladistics 11: 91- 104. Hassall, A.H. 1 845 . A history of the British freshwater algae (including descriptions of the Diatomaceae and Desmidiaceae). London. Hendey, N.I. 1964. An introductory account of the smaller algae of the British coastal waters. Part V. Bacillariophyceae (Diatoms). Fisheries Investigations, Ministry of Agriculture, Fisheries and Food, series IV. London. Kiitzing, F.T. 1833. Synopsis Diatomacearum oder versuch einer systematischer Zusanmen stellung der Diatomeen. Linnaea, 8: 526-620, Taf 13-19. Berlin. 1849. Species Algarum. Lipsiae. Mann, D.G. 1982. An ontogenetic approach to diatom systematics. Proceedings of the 7th Diatom Symposium: 113-144. Mayr, E. 1969. Principles of systematic zoology. New York. McCall, D. 1 933. Diatoms (recent and fossil) of the Tay District. Journal of the Linnean Society of London, Botany 49: 219-308. Neff, N.A. 1986. A rational basis fora priori character weighting. Systematic Zoology 35: 110-123. Nixon, K.C. & Carpenter, J. M. 1993. On outgroups. Cladistics 9: 413^*26. Peragallo, H. 1891. Monographic du genre Pleurosigma et des genres allies. Le Diatomiste 4: 1-35. Pimentel, R.A. & Riggins, R. 1987. The nature of cladistic data. Cladistics 3: 201- 209. Platnick, N.I., Coddington, J.A., Forster, R. & Griswold, C.E. 1991. Spinneret morphology and the phylogeny of haplogyne spiders (Araneae, Aranemorphae). American Museum Nov. 3016: 1-73. Quekett, J.T. 1 848. A practical treatise on the use of the microscope including the different methods of preparing and examining animal, vegetable and mineral structures. Rabenhorst, L. 1853. Die Susswasser-Diatomaceen (Bacillarien) fur Freunde der Mikroskopie. Leipzig. Rabenhorst, L. 1864. Die Europea Algarum aquae dulcis et submarinae. Sectio I. Algas diatomaceas complectens, cum figuris generum ominium xylographice impressis. Apud Eduardum Kummerum, Lipsiae. Reid, G. 1998. The Taxonomy of Gyrosigma Hassall and Pleurosigma W. Smith. Unpublished PhD thesis. University of Bristol (1997). & Sidwell, K. 2001. Testing continuous characters in cladistic analysis. In: P. Forey & N. Macleod (Eds) Morphology, Shape and Phylogenetics. Special Volume of the Systematics Association. London. Rodrigo, A. 1992. Two optimality criteria for selecting subsets of most parsimonious trees. Systematic Biology 41: 33-40. Ross, R., Cox, E.J., Karayeva, N.I., Mann, D.G., Paddock, T.B.B., Simonsen, R. & Sims, P.A. 1979. An amended terminology for the siliceous components of the diatom cell. Beiheft Nova Hedwigia 64: 513-533. & Sterrenburg, F.A.S. 1 996. Proposal to Conserve or Reject. ( 1 205) Proposal to conserve the name Navicula angulata E.J. Quekett (Pleurosigma angulatum (E.J.Quekett) W.Sm.) (Bacillariophyta) with a conserved type. Taxon 45: 121-122. Round, F.E., Crawford, R.M. & Mann, D.G., 1990. The diatoms. Cambridge. Smith, W. 1852. Notes on the Diatomaceae with descriptions of the British species included in the genus Pleurosigma. Annals and Magazine of Natural History, 9: 1- 12. 1853. A synopsis of the British Diatomaceae; with remarks on their structure, functions and distributions; and instructions for collecting and preserving speci- mens. London. 1856. A synopsis of the British Diatomaceae; with remarks on their structure, functions and distributions; and instructions for collecting and preserving speci- mens. London. Sneath, P.H.A. & Sokal, R.R. 1973. Numerical Taxonomy. The principles and practices of numerical classification. San Francisco. Sterrenburg, F.A.S. 199 la. Studies on the genera Gyrosigma and Pleurosigma (Bacillariophyceae). The Typus Generis of Pleurosigma, some presumed varieties and imitative species. Botanica Marina 34: 561-573. 199 Ib. Studies on the genera Gyrosigma and Pleurosigma (Bacillariophyceae). Light microscopical criteria for taxonomy. Diatom Research 6: 367-389. Swofford, D.L. 1990. PAUP: Phylogenetic analysis using parsimony version 3.0. Champaign. Illinois. Thiele, K. 1993. The holy grail of the perfect character: the cladistic treatment of morphometric data. Cladistics 9: 275-304. Turner, H & Zandee, R. 1995. The behaviour of Goloboff s Tree fitness measure F. Cladistics 11: 57-72. Van Heurck, H. 1885. Synopsis des Diatomees de Belgique. Anvers. Woodward, J.J. 1871. On the structure of the Podura Scale and certain other test- objects, and of their representation by photo-micrography. Monthly Microscopical Journal 5: 149-162. INDEX TO NEW TAXA Hypericum sect. Elodeoida N. Robson 43 H. daliense N. Robson 83 sect. Graveolentia N. Robson 43 H. kingdonii N. Robson 74 sect. Hypericum H. monanthemum subsp. filicaule (Dyer) N. Robson 78 subsect. Erecta N. Robson 43 H. subcordatum (R. Keller) N. Robson 78 subsect. Hypericum series Senanensia N. Robson 43 Lianthus N. Robson 38 sect. Monanthema N. Robson 43 L. ellipticifolius (H.L. Li) N. Robson 38 sect. Sampsonia N. Robson 43 Pleurosigma lysekilii G. Reid 1 14 H. ascyron subsp. gebleri (Ledeb.) N. Robson 57 P. malmoensis G. Reid 115 H. ascyron subsp. pyramidatum (Aiton) N. Robson 58 P. mamoranqi G. Reid 1 14 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 24 No. 1 Pre-Linnaean references for the Macaronesian flora found in Leonard Plukenet's works and collections. J. Francisco-Ortega, A. Santos-Guerra and C.E. Jarvis. 1994. Pp. 1-34, 16 figs. Studies on the lichen genus Sticta (Schreber) Ach.: II. Typification of taxa from Swartz's Prodromus of 1788. D.J. Galloway. 1993. Pp. 35^18, 9 figs. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 4. Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F. Prud'homme van Reine and W.J. Woelkerling. 1994. Pp. 49-90 Ifig. Studies on the Cretan flora 3. Additions to the flora of Karpathos. N.J. Turland and L. Chilton. 1994. Pp. 91-100, 1 fig. 43.25 No. 2 Observations on the benthic marine algal flora of South Georgia: a floristic and ecological analysis. D.M. John, P.J.A. Pugh and I. Tittley. 1994. Pp. 101-1 14, 8 figs. Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical species (excluding Australia). D.J. 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Recent records of pteridophytes for Belize, Central America. D.A. Sutton, A. Hughes and B. Bulmer-Thomas. 2000. Pp. 81- 99. Collections of flowering plants by Francis Buchanan-Hamilton from Nepal, 1802-1803. J.R. Press and K.K. Shrestha. 2000. Pp. 101-130. 43.40 Volume 31 No. 1 New synonymy in some Asian species of Syrrhopodon (Calymperaceae: Musci). L.T. Ellis. 2001. Pp. l^t, 2 figs. Two new species of Pilea (Urticaceae) from Panama. A.K. Monro. 2001. Pp. 5-7, 1 fig. Synopsis of Mesoamerican Pilea (Urticaceae), including eighteen typifications and a key to the species. A.K. Monro. 2001. Pp. 9-25. The Japanese plant collection of Engelbert Kaempfer ( 1 65 1 - 1716) in the Sir Hans Sloane Herbarium at The Natural History Museum, London. P-A. Hinz. 2001. Pp. 27-34, 1 fig. Index to new taxa. P. 35. 43.40 CONTENTS 37 Studies in the genus Hypericum L. (Guttiferae) 4(1). Sections 7. Roscyna to 9. Hypericum sensu lato (part 1) N.K.B. Robson 89 A bibliography of the scientific work of Boris V. Skvortzov (1 896-1 980) with commentary on the publications concerning diatoms (Bacillariophyta) DM Williams and G. Reid 107 On the identity of Pleurosigma angulatum (Bacillariophyta) and related species G. Reid 119 Index to new taxa Bulletin of The Natural History Museum BOTANY SERIES Vol. 31 , No. 2, November 2001