LAMIALES NEWSLETTER

LAMIALES Issue number 4 February 1996 ISSN 1358-2305

EDITORIAL CONTENTS R.M. Harley & A. Paton Editorial 1 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK

The Lavender Bag 1 Welcome to the fourth Lamiales Universitaria, Coyoacan 04510, Newsletter. As usual, we still Mexico D.F. Mexico. Tel: Lamiaceae research in require articles for inclusion in the +5256224448. Fax: +525616 22 17. Hungary 1 next edition. If you would like to e-mail: [email protected] receive this or future Newsletters and T.P. Ramamoorthy, 412 Heart- Alien Salvia in Ethiopia 3 and are not already on our mailing wood Dr., Austin, TX 78745, USA. list, or wish to contribute an article, They are anxious to hear from any- Pollination ecology of please do not hesitate to contact us. one willing to help organise the con- Labiatae in Mediterranean 4 The editors’ e-mail addresses are: ference or who have ideas for sym- [email protected] or posium content. Studies on the genus Thymus 6 [email protected]. As reported in the last Newsletter the This edition of the Newsletter and Relationships of Subfamily Instituto de Quimica (UNAM, Mexi- the third edition (October 1994) will Pogostemonoideae 8 co City) have agreed to sponsor the shortly be available on the world Controversies over the next Lamiales conference. Due to wide web (http://www.rbgkew.org. Satureja complex 10 the current economic conditions in uk/science/lamiales). Mexico and to allow potential partici- This also gives a summary of what Obituary - Silvia Botta pants to plan ahead, it has been the Lamiales are and some of their de Miconi 11 decided to delay the conference until uses, details of Lamiales research at November 1998. It is hoped that the Kew and a list of currently recog- Bibliography of recent conference will be held in Oaxtepec, nised genera. Information concern- taxonomic publications 12 near Mexico City. The organisers ing the conference will also be car- will be: Baldomero Esquivel, Institu- ried here as planning proceeds. to de Quimica, Circuito Exterior, Cd. ❑

THE LAVENDER BAG THE ACTIVITY OF A LAMIACEAE Susyn Andrews RESEARCH TEAM IN Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK HUNGARY

Imre Máthé The Lavender Bag newsletter first in the genus. The Lavender Bag Institut of Ecology and Botany, appeared in May 1994 and has will appear in May and November Hungarian Academy of Sciences, been welcomed by Lavender and if you would like to receive H-1263, Vácrátót, Hungary enthusiasts all over the world. Its future issues, please send £2.50 aims are to facilitate information (£4.00 overseas via air mail) to the exchange, aid identification and Editor, Mrs Joan Head, 6 Church A significant part of Hungarian accurate labelling and provide a Gate, Clipston-the-Wolds, Key- Lamiaceae research is conducted by forum for amateur and professional worth, Nottingham, NG12 5PA, a joint research team of approxi- plantsmen and plantswomen who United Kingdom. mately 15 researchers at the have more than a passing interest Research Institute of Ecology and ➣

1 Botany of the Hungarian Academy botanical garden seed exchanges, biology of the species. The bio- The first results of these investiga- of Sciences (Vácrátót) and at the partly by gathering and/or cultivating mass and certain secondary tions will be published soon. A CENTRAL Institute of Pharmacognosy of the the native species (92 species are in metabolites of the species were AMERICAN WEEDY SALVIA IN ETHIOPIA A. Szent-Györgyi Medical Univer- Hungary). Nowadays the collection measured during the growing sea- The diversity of compounds pre- sent in many plants, forced us to sity (Szeged). This team is headed has 500 accessions of some 150 son (time-dependent variation). In Sebsebe Demissew deal with methodological prob- by the author of this paper. In addi- species in 30 genera. a similar way the populations of The National Herbarium, Addis Abeba University, P.O. Box 3434, Addis tion to the work of this team some various origin (space-dependent lems. We had to justify the repro- These populations give us the Abeba, Ethiopia. additional research is carried out in variation) were compared with each ducibility of our results and meth- opportunity for multidisciplinary co-operation with other Institutes other. Such studies have been per- ods, and we wanted to elaborate work. The continuous revision like the Institute of Pharmacognosy formed on Melissa officinalis L., quick and sensitive methods for In connection with a collaborative Leaves petiolate; Leaf-blade (reevaluation) of the taxa from a of the Semmelweis Medical Uni- Lavandula angustifolia Mill., Oci- routine investigations so that the research between the University of broadly ovate, 4-6 x 3.5-5 cm, morphological viewpoint is of out- versity (Budapest), the Medicinal mum basilicum L., Salvia officinalis comparative work could be more Addis Abeba, Ethiopia and the sparingly pubescent, acuminate at standing importance. As a result of Plant Research Institute L., S. pratensis L., S. nemorosa L., reliable. TLC Densitometry University of Illinois in Cham- the apex, truncate at the base, fine- this systematic work new taxa in (Budakalász) and with the Depart- S. nutans L., Marrubium species, proved to be a good tool beside paign Urbana, USA, I made field- ly and regularly crenate. Inflores- Hungary have been found. ment of Chemical Engineering of Hyssopus officinalis L., etc. The UV-VIS photometry, HPLC, trips in 1994-95 to the project sites cence simple or paniculate; verti- the Technical University Since the beginning of our work, purpose of this work was to select OPLC, GC, GC-MS measure- in Welo Region in north central cils 6-14 flowered. Calyx hispid, (Budapest). The research is spon- comparative studies have been car- the best strains for further breeding. ments. Ethiopia. In both trips, I came ribbed, 4-6 mm long, 2-lipped, sored partly by National Grants ried out, among them, the determi- At the same time information could Special processes were also tried for across and collected a very unusual upper lip + entire. Corolla blue, 5- (OTKA) and partly by companies. nation of the presence, or absence be gained about the expected diver- a more detailed evaluation of the herbaceous labiate, which I had not 7 mm long. Seeds black. The School of Pharmacy and Bio- of some chemical characteristics of sity of the species concerned. encountered before. After collect- chemistry of some common Lami- The plant is a native of Central medical Sciences, University of the Lamiaceae. The excellent work ing the specimen for the first time The occurrence and quantitative aceae. A series of investigations America. It has been introduced to Portsmouth, and University Insti- of Cantino & Sanders (1986) pro- in 1994, I tried to match it with changes of the N-methylated were carried out on Salvia offici- the United States and ?Canada. tutes in Transylvania (Romania) are vided the basis for subfamilial com- specimens at the National Herbari- amines characteristic of the family nalis, Lavandula angustifolia, According to available informa- among the collaborating foreign parison. In addition to essential um, Addis Abeba University are also investigated. Some com- Melissa officinalis for the compari- tion, the plant is only so far known research institutes. oils, rosmarinic acid, iridoids, etc. (ETH), but without success. In a pounds like trigonelline, choline son of their extracts gained by in the African Continent from Pre- we found that the freely occurring subsequent visit to the project area The theoretical basis of the investi- betain, stachydrin, etc. may be of supercritical fluid CO2 extraction, toria, South Africa, having been triterpene carbonic acids (oleanolic in 1995, I collected the plant in gations dates back to a general pro- important role in stress. The study and traditional water steam distilla- introduced in 1943 (Codd, 1985). and ursolic) may merit more other localities as well asking the ject, “Chemotaxonomic and Pro- of their variation may be of chemo- tion respectively. detailed study (Máthé et al 1993). local people if they had any infor- The species was probably intro- duction Biological Evaluation of taxonomic significance as well as The number of papers, short com- Though it has been known for a mation on the plant: local names, duced to Welo Region in north Plants, Native or Suitable for Intro- physiological one. munications, abstracts of presenta- long time that these compounds are local uses, etc. I was told by nearly central Ethiopia in the 1980’s. duction into Hungary”. This tions has surpassed one hundred present in the family, they seem to Some investigations were carried all informants that the plant was The Region was frequently affect- research was launched by the late since the beginning of this occur in much larger amounts, like out in connection with the pres- known in the area only for only ed by drought and famine followed Prof. I. Máthé Sen. more than three research. the essential oil content, in the ence of beta ecdysone in the four to six years and was spreading by massive humanitarian assis- decades ago. The goal of this pro- species of Nepetoideae rather than Lamiaceae. Ecdysteroids are pre- rapidly. I then thought that the tance, one element of which was ject is to evaluate plant taxa for cul- those of the Lamioideae. This phe- sent in Ajuga. We have tested plant may have been introduced. In the distribution of grains and cere- tivation in Hungary. Within it, the References. nomenon seemed to be interesting other genera and could not find a research visit in July 1995, to the als for food and for cultivation. study of the Lamiaceae was started Cantino P.D. Sanders R.W. (1986): enough to start intensive work on them in detectable quantities in Herbarium, Royal Botanic Gar- Since then the plant has gradually by three researchers in 1986. Subfamilial classification of Labi- the study of the variation of these some fifty species tried (including dens, Kew (K), my suspicion was established itself and at present it atae. Systematic Botany. 11: The object of this research is the compounds from various aspects. Salvia). This work is going on confirmed and the plant was iden- seems to be spreading vigorously, 163-185. comparative evaluation of various and many interesting chemotaxo- tified (with the help of Drs. Ray replacing native herbs in some The study of the pattern of inor- species and accessions of the same nomic data result. Harley and Alan Paton, Kew) as sites. The plant does not seem to ganic element accumulation (27 Máthé, I. Jr., Máthé, I., Máthé, Á., species from different origins grow- A relatively new aspect of our Salvia tiliifolia Vahl, a native of be eaten by grazers at all. Apart elements) also showed an interest- Dinka, M., Borhidi, A. & Miklóssy, ing under similar conditions. This approach is the isolation of sec- central America. from its uncontrolled expansion its ing picture. It turned out that a V.V. (1992). Studies of element includes chemical, biological pro- ondary metabolites from selected adverse effects, if there are any, certain correlation could be patterns in the family Lamiaceae. S. tiliifolia was described by Vahl duction, and taxonomic research. species. Detailed isolation remain to be seen. observed in the accumulation Acta Horticulturae 306: 268-275. (1794: 7) as S. tiliaefolia from The study of variation within the processes are performed in con- (Máthé et al 1992). The cluster Mexico. The species has been sub- same taxa helps find the best condi- nection with Salvia and Marrubi- Máthé I. Jr., Miklóssy V.V., Máthé analysis of the data revealed only sequently discussed in various Specimens Examined: tions for optimal yield or, as we call um for diterpenoids, Hyssopus, Á., Bernáth J., Oláh L., Blunden G., slender connections among genera publications: Correll and Johnston it, the optimum production of the Melissa and Lavandula for pheno- Patel A.V. (1993): Essential oil con- Welo: 5-6 km from Kombolcha at the subfamilial level. (1970: 1370); Standley and most significant secondary metabo- lic compounds like flavonoids and tent as chemotaxonomic marker for town towards Yegof State Forest Williams (1973: 298) and Codd lites. To achieve this end Lamiaceae Another level of our approach is the coumarines, Physostegia virgini- the genus Salvia with reference to along the edges of the main road, (1985: 100). populations were created in an study of the variation of populations ana (L.) Benth. for iridoids, etc. its variation in Salvia officinalis L. 2070-2150 m, 10 Dec. 1994, Seb- experimental field partly from seeds of the same species. This has been besides the essential oil content Acta Horticulturae 330: 123-132. Annual herb 40-70 cm tall. Stems sebe Demissew 4332(ETH); 4 km of various species, obtained via done by studying the production obtained by steam distillation. sparingly pilose, simple or branched. from Hayq towards Gobeya, edge ➣ ❑

2 3 of cultivated farm, 1930 m, 8 June Water Supply, Pretoria. Hoffmanns. & Link. The total polium. The group of solitary bees highly nectar rewarding flowers and comparatively few. On the other 1995, Sebsebe Demissew 4510 range of the family’s flowering time extended duration of flowering (55 hand, the harsh environment the Correll, D.S. & Johnston, M.C. comprised 95 species, making (ETH); Hayq Estifanos Church, 5 was 142 days, from calendar date 43 vs. 36 days of the Late flowering Late flowering species have to over- (1970). Manual of the Vascular 47.3% of all insect species visiting km from Hayq town under a big to 285. Flower life span was in group). In summary, they were come is compensated by an abun- plants of Texas. Texas Research Labiatae, out of which 15 were one Ficus tree, 1940 m, 11 June 1995, average 1.8 days (range 1.2Ð3.1). characterized by an increased floral dance of pollinator species. Yet, Foundation, Renner, Texas. plant specialists. The importance Sebsebe Demissew 4630 (ETH, K). The duration of flowering of a of Labiatae for specialist bees was attractiveness compared to the Late these differences are accompanied Standley, P.C. & Williams, L. O. species was negatively correlated due to the fact that Labiatae were flowering species and by a differen- by morphological adaptations of the (1973). Labiatae, Flora of with the time of flowering starting the main host plants at the end of tiation in flower prototypes within plants to overcome water stress: all References Guatemala Part 9, No.3. in Fiel- from January 1st. the flowering season, i.e. during the the group. By contrast, in Late Late flowering species are either diana Bot. 24(9): 237-317. flowering species, flower character extremely hairy or particularly fra- Nectar Ð Labiatae were by far the hot and rainless summer. Codd, L.E. (1985). Lamiaceae in differences were minimal, and the grant, whereas amongst those of the Vahl, M. (1794). Symbolae Botan- most nectar rewarding species in the As to the types of bees visiting O.E. Leistner (Editor). Flora of per flower nectar production low. Early group only two have these icae, part 3. phrygana, both in volume and sugar them, Labiatae formed a particular Southern Africa 4(1): 1-247. characteristics (viz. S. triloba, and content: half of the top ten nectar- group. Compared to non-labiates, Differences in floral morphology, Department of Agriculture and ❑ P. fruticosa). These differentiations producers of the phrygana were Labiatae were visited by the large- and floral rewards between the two in pollination profiles suggest that Labiatae. The differences became sized, and the long-tongued bees time guilds were accompanied by the real Mediterranean group of even higher if, instead of the per (viz. Anthophoridae, Apidae, and differences in the composition of Labiatae is the Late flowering one: flower nectar production of a Megachilidae), in particular by insect visitors. The most interest- LABIATAE: this is also supported by the fact that species, the per population nectar Megachilidae, while they were ing fact is that out of 201 pollinator A KEY FAMILY FOR WILD BEES AND THE the latter group is particularly visit- production was taken into consider- avoided by Andrenidae and Halicti- species of Labiatae (98 bees), only POLLINATION ed by Megachilidae, the most ation. These findings suggested the dae. The association of Labiatae 19 species (13 bees) were shared by ECOLOGY IN MEDITERRANEAN PHRYGANIC Mediterranean of all bee families in characterisation of the members of with Megachilidae, the most both Early and Late flowering COMMUNITIES this community. Labiatae as the consistent nectar Mediterranean of all 7 bee families groups. However, contrary to what producers of the Mediterranean occurring in phrygana, implies that one might have expected, the Managing phryganic Labiatae Theodora Petanidou ecosystems (cf. also Herrera, 1985). these two groups have interdepen- species most differentiated among for bees Ð Most Labiatae are peren- Catholic University of Leuven, Botanical Institute Ð Laboratory of Systemat- At community level this is very dently co-evolved in the Mediter- them and more nectar rewarding nials, especially those identified ics, Kardinaal Mercierlaan 92, B-3001 Heverlee (Leuven), Belgium. important because Labiatae consist ranean region. This is also demon- (i.e. Early flowering) were not visit- within the Mediterranean group, of a unique group that bees rely on strated by the fact that all late flow- ed by more visitors. The Early therefore they come into bloom for energy, and most importantly, ering Labiatae except T. polium flowering group was visited by a several years after a fire Ð a very for water supply in an area charac- constituted the stronger supporters total of 48 insect species, whereas regular element of the Mediter- terized by water shortage. of Megachilidae in the phrygana the Late flowering one by 172 ranean ecosystems. Because Labi- The major physiognomic character- atae are pollinated by solitary bees (i.e. the plants with the highest species. Moreover, the number of atae species are by far the consis- istic of the garrigue and the phry- (Herrera, 1987; Shmida & Dukas, Pollinators Ð All phryganic Labi- bridge value in facilitating insect species per plant species was tent nectar producers of phrygana, gana (i.e. the counterpart of garrigue 1990; Dafni, 1991). This paper atae species were found to be Megachilidae to survive over time; significantly higher in the Late and because they support a particu- in the East Mediterranean Basin), is summarises the results of a 12-year insect-pollinated, visited by a total cf. Petanidou & Ellis, 1995). flowering group even if T. larly diverse wild bee fauna, they the dominance of the Labiatae fami- study (1983Ð1994) at community of 201 insect species, primarily capitatus, supporting an extremely plan an essential key role in the ly. Thymus spp. are the most level on the pollination ecology of Hymenoptera and secondarily Why flowering in guilds? Ð Based high number of insect species, was conservation of the ecosystem as a prominent example, not only Labiatae in a phryganic ecosystem Diptera. The total number of polli- on seven flowering traits, viz. corol- excluded from the analysis. This whole. This implies that the vital because they give their characteris- near Athens, Greece (Petanidou, nation events amounted to 324. la depth, corolla size, height of the means that the species offering role that Labiatae play cannot be tic fragrance, but also because the 1991; Petanidou & Vokou, 1993; Forty-three insect species, about flower from the ground, flower copious floral rewards (i.e. Early substituted by other species, such genus is identified with the ecosys- Petanidou & Ellis, 1995; Petanidou half of them bees, were exclusively cover of the population, nectar flowering) were not those abundant- as annuals which come dominantly tem itself, locally known as thy- & Smets, 1995; Petanidou et al., supported by Labiatae. Thymus amount per flower, essential oil pro- ly serviced by pollinators (i.e. Late into bloom during the first post-fire marónas (Greece) or tomillares 1995). capitatus in particular, supported an duction, and flower colour, the 10 flowering). years. In other words, the too-fre- (Spain), from the words thymari or exceptionally high number of Labiatae of phrygana were distin- Flowering Ð Ten Labiatae species quent fires will certainly diminish tomillo meaning thyme. In W. flower visitors (123 species) and guished in two equally represented All the above findings imply that the occurred in the study area. In the the biological value of the phry- Greece phrygana may also be domi- 50% of the monotropous visitors groups: the Early flowering group, two groups of Labiatae are funda- order of flowering (i.e. median date gana, not only as to its floral ele- nated by another labiatae species, (i.e. those that are exclusive to one characterized by late winter-early mentally different as to their ecolo- of flowering) these were: Lamium ment, but as to the bee fauna as i.e. Phlomis fruticosa, therefore plant species) of the family. spring flowering, encompassing the gy and, consequently, as to their pol- amplexicaule L. ssp. amplexicaule, well. Again, in order to attain the locally called asphakónas, from first five species mentioned above; lination characteristics: the first Salvia verbenaca L., S. triloba L. Among the Hymenoptera visiting maximal diversity in these continu- aspháka, its common name. and the Late flowering group, com- group, by flowering early in the year fil., Phlomis fruticosa L., Prasium Labiatae, solitary bees were the prising the remainder, characterized faces milder environmental stresses ously changing Mediterranean The species of Labiatae are predom- majus L., Satureja thymbra L., most important group being active by late spring-summer flowering. (e.g. water shortage) but meets diffi- communities a mosaic of different- inantly bee-pollinated (van der Pijl, Stachys cretica L. ssp. cretica, Teu- throughout their flowering season, An epitome of the flowering fea- culties with pollinator availability; age-succession stages should be 1972; Stebbins, 1974; Faegri and crium polium ssp. capitatum (L.) while wasps formed a particular tures of the two groups revealed therefore, it maximises its attractive- maintained by incorporating tradi- van der Pijl, 1979). In the Mediter- Arcangeli, Ballota acetabulosa (L.) guild visiting the late flowering that the Early flowering species ness to optimise its competitive abil- tional methods of management, ranean region, in particular, Labi- Bentham, and Thymus capitatus (L.) Thymus capitatus and Teucrium were characterized by dissimilar ity at a time when pollinators are such as light grazing, controlled ➣

4 5 fires, and controlled bee-keeping Petanidou, Th. (1991). [Pollination in Mediterranean shrublands. Api- The west Mediterranean Region France and the north of Italy. This zarote (Canary Islands), T. serrula- especially during the first post-fire ecology in a phryganic ecosystem]. dologie 26: 39-52. seems to be the centre of origin of species has been cultivated since tus and T. schimperi in Ethiopia, T. years. In view of achieving this Thessaloniki: Ph.D. Thesis. (In Petanidou, Th. & Vokou, D. (1993). this genus. Jalas (1971) divided it Roman times and is well-known laevigatus in the southwest of the goal, creating phrygana nature Greek, with English summary). Pollination ecology of Labiatae in a into 8 sections: Mastichina, throughout the world as a pot-herb. Arabian Peninsula. Another group reserves is not only necessary but Micantes, Piperella, Pseudothym- of species are more or less herba- phryganic (East Mediterranean) Section Teucrioides was studied by more urgent than ever. Petanidou, Th. & Ellis, W.N. bra, Thymus, Teucrioides, Hypho- ceous and occur in the Mediter- ecosystem. American Journal of Hartvig (1987), and comprises the (1995). Interdependence of native dromi and Serpyllum. ranean mountains and the rest of Botany 80: 892-899. previously T. teucrioides and the bee faunas and floras in changing Eurasia as far as Japan and also The first 5 are only found on the newly described T. rechingeri and References Mediterranean communities. In: Pijl, van der L. (1972). Functional along the coasts of Greenland. The Iberian Peninsula, in the northwest T. leucospermus, found in the Matheson, A., Buchmann, S.L., considerations and observations of species of the last group seem to be Dafni, A. (1991). Advertisement, of Africa and the Macaronesian mountainous regions of Greece. O’Toole, C., Westrich, P. & the flowers of some Labiatae. younger in evolutionary terms and flower longevity, reward and nectar Region (Western Region). These 5 Williams, I.H. (eds) The conserva- Blumea 20: 92-103. At the moment I am working on the have probably been an actively protection in Labiatae. Acta Horti- sections are more or less well- tion of bees - Proceedings of a Sym- section Hyphodromi. I, like Jalas, evolving since the last glaciation, culturae 288: 340-346. Shmida, A. & Dukas, R. (1990). known (Morales, 1986, 1994). posium held by The Linnean Society recognize 3 subsections: Sub- when this group colonized the new Progressive reduction in the mean Faegri, K, & Pijl, van der L. (1979). of London & International Bee Section Mastichina comprises two bracteati, Serpyllastrum and Thym- lands free of ice. This group is also body size of solitary bees active dur- The principles of pollination ecolo- Research Association. Academic species found only on the Iberian bropsis. There are more than 60 taxonomically very difficult and ing the flowering season and its cor- gy. Pergamon, Oxford. Press. In press. Peninsula: T. mastichina or wild species within this section, that corresponds to the 3 last subsec- relation with the sizes of bee flowers Petanidou, Th., Ellis, W.N., Mar- marjoram and T. albicans. occur in the Mediterranean Region tions. Jalas (1971) divided this sec- of the mint family (Lamiaceae). Herrera, J. (1985). Nectar secre- garis, N.S. & Vokou, D. (1995). and the arid regions of Asia. tion into 7 subsections. tion patterns in southern Spanish Isreal Journal of Botany 39: 133-141. Section Micantes comprises the Constraints on flowering phenology herbaceous looking Ibero-macarone- Subsection Subbracteati includes Subsection Insulares comprises T. Mediterranean scrublands. Israel in a phryganic (East Mediterranean Journal of Botany 34: 47-58. Stebbins, G.L. (1974). Flowering sian T. caespititius, that occurs in the north African T. algeriensis, the willkommii, an endemic species that shrub) community. American Jour- Plants: Evolution above the species the northwest Atlantic Region of the central Spanish T. mastigophorus, occurs on the mountains of the Herrera, C.M. (1987). Components nal of Botany. In press. level. Harvard University Press, Iberian Peninsula and also in the south Italian T. spinulosus and provinces of Castellón and Tarrago- of pollinator “quality”: compara- Petanidou, Th. & Smets, E. (1995). Cambridge, Mass. Madeira and Azores, and the woody the east Mediterranean T. boissieri, na (East of Spain), T. richardii tive analysis of a diverse insect The potential of marginal lands for species from Morocco T. sature- T. dolopicus, T. cherlerioides and from the Balearic Islands, Maretti- ❑ assemblage. Oikos 50: 79-90. bees and apiculture: nectar secretion joides and T. riatarum. T. striatus. no island near Sicily and Section Piperella is made up of one Subsection Serpyllastrum includes Yugoslavia, the north African T. endemic species T. piperella from T. bracteatus, T. leptophyllus, T. dreatensis and T. guyonii, the Valencia, Murcia and Albacete lacaitae of the middle of Spain, T. Canary Island T. origanoides and provinces in eastern Spain. granatensis of the Southeast of the endemic species of northwest Spain, and T. fontqueri, that occurs Turkey T. bornmuellerii. Section Pseudothymbra is character- south of and near the Pyrenees; or ized by the presence of bracts and Subsection Pseudopiperellae com- STUDIES ON THE GENUS THYMUS L. the east Mediterranean T. bracteo- long flowers. In some cases the prises T. herba-barona of Mallorca sus, T. comptus and T. zygioides. (2n=28), Corse (2n=56) and Sardinia Ramón Morales corolla can be up to 2 cm. Of the 9 (2n=84) (Mayol & al., 1990) and T. Real Jardín Botánico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain species, 7 occur in the south of the Subsection Thymbropsis includes Iberian Peninsula (T. lotocephalus, the north African T. broussonetii, T. nitens from the South of France. T. villosus, T. longiflorus, T. mem- maroccanus, T. lanceolatus, T. Subsection Isolepides, with the branaceus, T. moroderi, T. funkii, T. numidicus, T. pallescens, the east European T. sibthorpii, T. The genus Thymus comprises about important place in the evolution of between species. All this makes tax- antoninae) and 2 are found in the Mediterranean T. laconicus, T. glabrescens, T. longedentatus and 150 species (Jalas & Kaleva, 1970) this genus and is responsible for the onomic studies in this genus a lot northwest of Africa (T. bleicheri- holosericeus and the Asian T. bovei, T. pannonicus. distributed throughout the arid, tem- other numbers. more difficult. anus, T. munbyanus). T. cariensis, T. cilicicus, T. decussa- Subsection Kotschyani includes the perate and cold regions of the Old tus, T. eigii, T. leucostomus, T. neu- The pollen of this genus is very As regards human use, thyme Section Thymus comprises 11 Ethiopian T. serrulatus and T. World north of the Equator and on the rophyllus, T. sipyleus and T. syria- homogeneous, usually hexacolpate species are very important and usual- species; 3 of these, T. hyemalis, T. schimperi, the Arabian T. laeviga- coasts of Greenland (Morales, 1989). cus. and with suprareticulate ornamenta- ly well-known as medicinal and aro- zygis and T. willdenowii are all in tus and a lot of Asian species. The known chromosome numbers tion; the size varies between 21-46 matic plants, used in pottery and also North Africa and the Iberian Penin- Section Serpyllum appears to be the Subsection Pseudomarginati are 2n=24, 26, 28, 30, 32, 42, 48, µm and P/E between 0.9 and 1.3 in cosmetics and perfumery in sula, 3 occur in the southwest of the oldest in the genus. More than 70 includes the European species T. 50, 52, 54, 56, 58, 60, 84 and 90, (Morales, 1986). almost all the places where they Iberian Peninsula (T. capitellatus, T. species belong to this section. They longicaulis, T. nervosus, T. ocheus, which correspond to the diploid, grow. Their essential oils are indus- camphoratus, T. carnosus) and 3 in occur throughout the area of the T. praecox, T. pulcherrimus, T. tetraploid and hexaploid levels. All chemical studies show the great trially obtained by distillation. They the southeast (T. orospedanus, T. genus, except in Madeira and the thracicus, T. stojanovii and the Probably, from a basic number x=7, homogeneity of this genus (Morales, are also used as ornamental plants baeticus, T. serpylloides). T. loscosii Azores. It is in this section that we Himalayan T. linearis (Jalas, 1973). the secondary basic numbers x=14 1986). Hybridization is very com- and a lot of cultivars have been is an endemic species from the Ebro find the biggest chromosomal vari- and x=15 originate. The most fre- mon where two or more species live obtained as a result of cultivation. River valley, and T. vulgaris lives in ation. There are also woody species Subsection Alternantes includes the quent numbers are 2n= 28, 30, 56 together. Usually there does not Thyme species have many popular the eastern half of the Iberian Penin- that grow in the mountains in arid European T. alpestris, T. alternans, and 60. Aneuploidy occupies an seem to be genetic incompatibility names in a lot of different languages. sula, the Mediterranean Region of areas e.g. T. origanoides in Lan- T. bihoriensis, T. comosus, T. ➣

6 7 froelichianus, T. oehmianus and T. Verbreitungstypen in der Gattung ción Serpyllum) y Thymbra en la hairy. Pogostemon was named for All species of Pogostemon, except auricularius (Blume) Hassk., P. pulegioides. Thymus L. (Labiatae). Feddes Península Ibérica. Ruizia 3: 1-324. its bearded stamens: pogon is Greek P. atropurpureus Benth., P. specio- hayneanus Benth., P. paniculatus Repert. 81(1-5): 93-106. for beard and pogonostemon means sus Benth., P. travancoricus Bed- (Willd.) Benth. and P. quadrifolius Subsection Serpyllum with the Euro- Morales, R. (1989). El género Thy- “with bearded stamen” (Stearn dome and P. reflexus Benth. which (Benth. in Wallich) Kuntze, in pean T. serpyllum and T. talijevii, Mayol, M., J. A. Rossello, M. Mus mus L. en la región mediterránea 1992). However close examination are glabrous, have the middle of the which living flowering material was and T. quinquecostatus of Japan. & R. Morales (1990). Thymus occidental (Lamiaceae). Biocosme herba-barona Loisel., novedad para Mésogéen 6(4): 205-211. of the taxa (Bhatti 1995) reveals filaments hairy. Press (1982), like examined, either in India or at Kew, I calculate that the Thymus includes important differences and calls into earlier workers, described the sta- it is clear that the filament hairs are España, en Mallorca. Anales Jard. Morales, R. (1994). El género Thy- more than 200 taxa. question the naturalness of Pogoste- minal hairs in the Pogostemon as situated unilaterally (Figure 2). The Bot. Madrid 47(2): 516. mus L. (Labiatae) en Africa. Anales monoideae. long and thread like in structure. density and structure of hairs on the As far as the chemistry of essential Morales, R. (1986). Taxonomía de Jard. Bot. Madrid 51(2): 205-236. However, close study of herbarium filaments make it difficult to observe oils is concerned, a lot of studies los géneros Thymus (excluida la sec- ❑ All four stamens are exserted from specimens and some living material the exact distribution of filament have been done and now we are cur- the corolla in all the taxa in ques- in India and at Kew shows that in hairs on most dried specimens. rently studying the composition of tion. However, in Anisomeles all species examined with hairy fil- the essential oils of 284 samples of although they are exserted from the The possession of moniliform fila- RELATIONSHIPS OF aments, 75 species in total, the hairs seven species around Madrid (Spain). corolla tube they do not exceed the ment hairs in Pogostemon contrasts SUBFAMILY POGOSTEMONOIDEAE are moniliform (Figure 1). The It was possible to analyze so many central lobe of the upper lip of the with the condition of other taxa in REVEALED BY STAMEN CHARACTERS possession of glabrous filaments is samples because it was not necessary corolla. Stamens emerge from the the Pogostemonoideae and the rest probably a derived feature in this to distil. With the new ATD (auto- upper part of the corolla tube, and of the Labiatae. Simple, uniseriate genus. The filaments of P. matic thermal desorption) method, G.R. Bhatti and M.J. Ingrouille are exserted between the sinus of and multicellular hairs have been brachystachyus Benth, and P. men- moreover only a small quantity (1-40 Biology Department, Birkbeck College, Malet Street, London upper and lower lobe of the corolla. recorded on the filaments in thoides Blume have been described mg) of plant was needed (Esteban & In Eurysolen gracilis Prain and Leu- Eurysolen gracilis and also in all Department of Botany, Shah Abdul Latif University Khairpur (MIRS) Sindh, as glabrous or nearly so (subnudis) al., 1993). Previous results showed cosceptrum canum Smith all fila- three species of Anisomeles exam- Pakistan. Fax 00 92 792 51914. in their type descriptions. Various that T. mastichina, T. vulgaris, T. ments are inserted under the upper ined (Figure 3). Stellate filament Floras have reported the same. lacaitae and T. praecox do not have lip of the corolla. In male flowers of hairs have been observed in Leu- However, P. brachystachyus has chemotypes and the composition of Colebrookea oppositifolia Smith sta- cosceptrum canum (Figure 4). A PhD. thesis on Pogostemon and Colebrookea Smith. In con- densely hairy filaments, and a few their essential oils was less variable. men filaments are exserted, but in Species of Comanthosphace and Desf. has recently been completed. trast to Pogostemon, these are all moniliform hairs have been seen on T. bracteatus did not have chemo- female flowers they are included in Rostrinucula have glabrous fila- Pogostemon with 80-90 species small genera with 1-6 species the filaments of P. menthoides. types, but its essential oils were very the corolla. ments. has sometimes been divided into each. Ryding (1993) recently sug- The hairs at the base of the fila- variable. T. pulegioides has 2 chemo- two genera, Pogostemon sensu gested that Comanthosphace, Leu- Each filament is terminated by a ments in Pogostemon, where pre- Endress (1994) has reported that types and T. zygis 4 chemotypes. stricto and Dysophylla Blume. A coseptrum and Rostrinucula be unilocular anther in all the taxa sent, are not moniliform but are several different angiosperm fami- number of other genera, all small removed from the Pogoste- except Anisomeles. Bilocular simple, unicellular or multicellular lies have moniliform hairs in their References and of uncertain affinity, have monoideae on the basis of nutlet anthers have been observed in the and villous, tomentose, strigose and flowers. For example they are Esteban, J.L., I. Martínez-Castro & been placed near Pogostemon anatomy. It has also been suggest- upper pair of filaments in A. indica setose. The presence or absence of widespread in the Commelinaceae J. Sanz (1993). Evaluation and either in the same tribe, Pogoste- ed (Cantino et al. 1992, Abu-Asab, (L.) Kuntze and A. malabarica (L.) these basal hairs has proved useful where they are found on the sta- optimization of the automatic ther- moneae (Benth. ex Endl.) Briq., or M.S. & Cantino, P.D. 1994) that R.Br. ex Sims, whereas, the lower in erecting a sub-generic classifica- mens of Rhoeo and Tradescantia. mal desorption method in the gas most recently in the same subfami- Anisomeles R.Br. has a close rela- pair of filaments are terminated by tion of Pogostemon, which will be Perhaps most interestingly monili- chromatographic determination of ly, Pogostemonoideae (Endl.) P.D. tionship with Pogostemon. unilocular anthers. All four fila- described elsewhere. form hairs are recorded for some Cantino, Harley and Wagstaff. plant volatile compounds. Journal of These taxa all have exserted sta- ments have a bilocular anther in A. members of the Verbenaceae. They are Rostrinucula Kudo, In the small number of species, P. Chromatography A, 657: 155-164. mens and in most taxa the anthers salviifolia R. Br. Endress (1994) included an illustra- Comanthosphace Moore, Leu- benghalensis (Burm. f.) Kuntze, P. are unilocular and the filaments are tion of Verbena peruviana which Hartvig, P. (1987). A taxonomical cosceptrum Smith, Eurysolen Prain revision of Thymus sect. Teucrioides shows moniliform hairs exactly like (Lamiaceae). Pl. Syst. Evol. 155: Fig. 1. Moniliform staminal hair of Fig. 2. Stamens of P. quadrifolius Fig. 3. Staminal hairs of Eurysolen Fig. 4. Staminal hairs of Leucoseptrum those of Pogostemon except they 197-213. Pogostemon quadrifolius. showing unilateral distribution of hairs gracilis. canum. are not borne on the filaments but at the sinus of the petals. Subfamily on filaments and unilocular anthers. Jalas, J. (1971). Notes on Thymus L. Pogostemonoideae might be better (Labiatae) in Europe. I. Supraspecif- restricted to Pogostemon. This will ic classification and nomenclature. be discussed in a paper recently Bot. J. Linn. Soc. 64(2): 199-215. submitted for publication. Jalas, J. (1973). Thymus subsect. Pseudomarginati in the Himalayas References and adjoining western mountain Bhatti, G.R. (1995). Systematics of ranges, and in Caucasia. Ann. Bot. Pogostemon Desf. (Labiatae) and Fennici 10: 104-122. related genera. Ph.D. Thesis, Birk- beck College, Univesity of London. Jalas, J. & K. Kaleva (1970). ➣ Supraspezifische Gliederung und

8 9 Cantino, P.D., Harley, R.M. and flowers. Cambridge University are good grounds for recognizing a tion site analysis of the Nepetoideae, Epling, C. & Játiva, C. (1966). A Naturk. Ser. A. 421: 1- 38. Wagstaff, S.J. (1992). Genera of Press. pp. 320-350. number of genera, which fall into descriptive key to the species of Sat- Wagstaff et al. (1995) produce evi- Stace, C.A. (1991). New Flora of Labiatae, status and classification. two distinct groups: Satureioid gen- ureja indigenous to North America. Press, J.R. (1982). Taxonomic stud- dence that Briquet’s treatment of the the British Isles. Cambridge Univer- In: R. Harley and T. Reynolds. era and Micromerioid genera. Brittonia 18: 244-248. ies in the Labiatae tribe Pogoste- group, submerging most genera sity Press. 675-677. Advances in Labiate Science. Royal moneae. Bull. Brit. Mus. (Nat. Unfortunately Doroszenko’s studies within Satureja, does not produce a Greuter, W., Satureja in Greuter, W. Botanic Gardens, Kew. pp. 511-522. Wagstaff, S.J., Olmstead, R.G. and Hist.). Bot. 10(1):1-89. did not extend to those genera with monophyletic group and several gen- & Raus, T. (editors) (1984). Med- Cantino, P.D. (1995). Parsimony Abu -Asab, M.S. & Cantino, P.D. only two fertile stamens, such as era need to be recognized. According Checklist Notulae 10. Willdenowia Ryding, O. (1993). Pericarp struc- analysis of cpDNA restriction site (1994). Systematic implications of Ziziphora in the Old World and a to their strict consensus tree, the 14(2): 302. ture and phylogeny if Lamiaceae variation in subfamily Nepetoideae pollen morphology in Subfamilies number of genera allied to Hedeoma merging of Satureja and Micromeria subfamily Pogostemonoideae. Pérez de Paz, P.L. (1994). Re the (Labiatae). American J. Bot. 82: Lamioideae and Pogostemonoideae in the Americas. In all respects apart would also require the inclusion of Nord. J. Bot. 14: 59-63. genus Micromeria Benth. (Labiatae) 886-892. (Labiatae). Ann. Missouri Bot. from stamen number these appear to such genera as Thymus, Thymbra Stearn, W.T. (1992). Botanical and nomenclatural inconsistency Gard. 81:653-686. be remarkably similar to Satureja and even Mentha, if a monophyletic Willemse, R.H. (1991). New combi- Latin. 2nd edition. David & Taxon. 43(2): 241-242. and its allies, and a detailed study group were to be maintained. nations and names for Macarone- Endress, P.K. (1994). Diversity and Charles, London. and cladistic analysis of the whole As a result of this, it seems very Seybold, S. (1988). Die Arten der sian Satureja taxa (Labiatae). Evolutionary Biology of tropical ❑ group is urgently needed. hasty to make any further purely Gattung Satureja L. (Labiatae) in Willdenowia 21: 81-85. Äthiopien. Stuttgarter Beitr. zur Doroszenko then goes on to recog- nomenclatural adjustments on the nize a total of 17 genera worldwide, grounds of convenience, until new CONTROVERSIES OVER THE SATUREJA two of which are new. Satureja evidence gives us surer support for COMPLEX sensu stricto is largely restricted to a firm basis for classification. With SILVIA MARGARITA BOTTA DE MICONI the Mediterranean region. 7 genera, the renewed interest in the family, 1942 - 1994. Ray Harley including Gardoquia and Xenopo- and with the evidence from molec- Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK ma, are restricted to the Americas, ular studies, this may now be close Sandy Atkins & Lulu Rico Arce and Micromeria, Calamintha, at hand. Herbarium, Royal Botanic Gardens Kew, Kew, Richmond, Surrey, Clinopodium, Acinos as well as the The outburst from Pedro Pérez de Calamintha Mill. (including sec- TW9 3 AE, UK. Irano-Turanian genus Cyclotrichi- Paz in Taxon (1994), over R.H. tions Clinopodium, Calomelissa References um are among the Old World gen- Willemse’s paper in which he sinks and Acinos.) Bentham, G. (1876). in Bentham, Silvia obtained her degree in Biolo- She undertook several study visits era recognized. Two new genera are all Macaronesian Micromeria into G. & Hooker, J.D. (editors). Genera gy in 1966 and became a teacher. At and collecting trips in South Ameri- Briquet (1896), however, placed all recognized from Africa. the genus Satureja, highlights one Plantarum 2: 1160-1223. Reeve & the same time, while teaching, she ca including Chile, Brazil and these into Satureja, recognizing 14 took several credits at the Faculty of Uruguay. In 1989 she was awarded of the main problems in taxonomy, The European genera have recently Co. London. sections within it. This treatment Science at the Universities of a scholarship to visit the Field where careful research and hesitan- been re-assessed by Stace (1991), has largely been followed by Epling Brenan, J.P.M. (1954). Plants col- Buenos Aires and La Plata. She Museum of Natural History in cy to publish due to insufficient who recognizes Satureja in the for the Americas (Epling & Játiva, lected by the Vernay Nyasaland began her research in 1971 at the Chicago, and from there she visited data, can be highjacked by hasty strict sense, but combines 1964, 1966), and also in Africa expedition of 1946. Mem. New Darwinion Insitute in Argentina, Austin, New York, Missouri and the assessment based on limited knowl- Calamintha, Acinos and Clinopodi- (Brenan, 1954; Seybold,1988) but York B.G. 9: 45-53. under the supervision of Arturo Smithsonian. edge. um under the latter, earlier name. not in Europe, where, until recently, Briquet, J. (1897). Labiatae. In A. Burkart, with whom she collaborat- In the recently published checklist of Micromeria, Clinopodium, Although Doroszenko’s treatment Engler & K. Prantl (editors). Die ed on the Flora Illustrada de Entre With the help of the Royal Soci- genera of Labiatae (Cantino et al., Calamintha and Acinos were usual- requires careful reassessment, he natürlichen Pflanzenfamilien. Teil Rios. In 1976, guided by Dr Angel L. ety, London she continued her 1992), the authors decided to retain ly treated as separate genera. was almost certainly correct in rec- 4, Abt. 3a. Leipzig. Cabrera, she began to specialise in revision of the tribe Verbeneae at Micromeria as a distinct genus, until Greuter, from a Mediterranean ognizing Old and New World the family Verbenaceae helped also Kew. She collaborated here with Cantino, P.D., Harley, R.M. & the results of a broader-based survey viewpoint, considered that the groups as generically distinct. by Nelida Troncoso. In 1985 she Dr Peter Brandham in the Jodrell Wagstaff, S.J. (1992). Genera of of the group, on a worldwide basis, division into Satureja and Among these, Gardoquia, which was awarded a scholarship to study Laboratories. Labiatae: Status and Classification. could be made known. Micromeria was an unnatural one, includes many large, shrubby in Paris with Dr Alicia Lourteig. species, primarily in the Andes, In R.M. Harley & T. Reynolds (edi- In 1985 she married Carlos Miconi, Bentham was responsible for the but also placed the other genera Here she was able to concentrate on with long, brightly coloured flowers tors.). Advances in Labiate Science: a sculptor. This was a very happy first detailed classification of the near Clinopodium into synonymy her work on the generic delimitation adapted to bird-pollination, certain- 511-522 (1992). period of her life, and Carlos Labiatae, modifying his system of the former, more on the grounds of Verbena, Glandularia and Junel- ly merits recognition. accompanied her on many of her several times. In his final classifi- of nomenclatural stability (Greuter Doroszenko, A. (1987). Taxonomic lia. She returned to Paris in 1991 to defend her thesis and she matriculat- trips away from home. cation (1876), he recognized a & Raus, 1984). While one may not wish to accept all studies on the Satureja complex. ed at the Université d’Orléans. number of genera that have, by A largely overlooked study by of Doroszenko’s conclusions, there Ph.D. thesis. Edinburgh University We first met Silvia in 1991 and some authors, been considered as Doroszenko (1987), entitled “Taxo- is little doubt that his painstaking and Royal Botanic Garden, Edin- Much of her work was taken up were immediately impressed with wholly or in part congeneric with nomic studies on the Satureja com- study went far towards providing a burgh. with editing and preparing many her enthusiasm for her work and her Satureja. These included Microme- plex”, and only available as an more natural arrangement of the Sat- Epling, C. & Játiva, C. (1964). floras especially Jujuy, Patagonia zest for life. She easily made friends ria (in which he placed the Ameri- unpublished thesis, concludes that ureja group, and a rationale for Revisión del género Satureja en and San Juan. She was the co-ordi- wherever she travelled, and her can Xenopoma Willd. as a section), on the basis of a number of charac- accepting a number of genera. In a America del Sur. Brittonia 16: 393- nator of the Verbenaceae for the warm and generous spirit will be Gardoquia Ruiz & Pavon, and ters, in particular leaf anatomy, there recently published cpDNA restric- 416. Floras of Paraguay and Argentina. sadly missed.

10 11 Tomas Barberan FA. Spanish CARYOPTERIS naceae) in Australia. J. Adelaide BIBLIOGRAPHY OF RECENT TAXONOMIC Labiatae: a source for natural Abu Asab MS, and others. Sys- Bot. Gard. 16: 1-16 (1995). PUBLICATIONS ON THE LAMIALES antioxidant compounds. Lamiales tematic implications of pollen mor- Newsl. no.2: 1-2 (1993). phology in Caryopteris (Labiatae). ELSHOLTZIA Syst. Bot. 18(3): 502-515 (1993). Dung NX, Chinh ND, Hai LD. ACHYROSPERMUM Cac ket qua nghien cu’u ve Chikuni A. A new species of CLERODENDRUM Hu’o’ng nhu xa (Elsholtzia wini- The following list of publications ANATOMY Morales R. Lamiaceae Guineae Achyrospermum Blume (Labiatae) Herman PPJ. Verbenaceae: a new tiana Craib., ho Lamiaceae) phan has been abstracted from the Kew Carlquist S. Wood anatomy of Aequatorialis nonnullae: 2. Font- from Malawi. Kew Bull. 50(3): species in the genus Clerodendrum. bo o’ Tay Nguyen. (Results of the Index of Taxonomic Literature Lamiaceae: a survey, with comments queria 39: 13-16 (1994). 605-610 (1995). Bothalia 25(1): 100-102 (1995). study on Elsholtzia winitiana (June 1993 to September 1995), on vascular and vasicentric trachei- Craib. distribution in Taynguyen.) and we are extremely grateful to ds. Aliso 13(2): 309-338 (1992). Phillipson P, Clement R, ANISOCHILUS Steane DA. Molecular systematics J. Biol. (Vietnam) 16(2): 13-15 the editors and compilers for their Rafamantanantsoa G. Madagas- Clement RA. Notes relating to the of Clerodendrum L. s.l. (Lami- (1994). assistance in preparing this bibli- Ryding O. The importance of peri- car: stop the car for another new flora of Bhutan: 18. New taxa and a aceae). Ph. D Thesis, University of ography. Where possible, articles carp structure in the classification of species! Lamiales Newsl. no.1: 5-7 new combination in the Labiatae. Oxford. 1995xi, 259p. Hu S, Fan C. (The scanning elec- are listed under genus or tribe, labiates. Lamiales Newsl. no.3: 1-3 (1992). Edinburgh J. Bot. 50(1): 33-38 tron microscope of pollen and fruit which are arranged alphabetically. (1994). (1993). CLINOPODIUM of four species “Xiangru”.) J. Publications which cover many Scott AJ, Bosser J, Harley RM. Morales Valverde R. Nota sobre Wuhan Bot. Res. 13(1): 95-96 CYTOLOGY genera are listed at the beginning 139. Labiees. In Bosser JM, and BALLOTA el genero Clinopodium L. (Labi- (1995) . Arista M, and others. Numeros under general subject headings. others, eds. Flore des Mas- Lancaster R. Plants that should be atae). An. Jard. Bot. Madrid 51(2): cromosomaticos de plantas occiden- All diacritical marks have been careignes: La Reunion, Maurice, better known: Ballota acetabulosa. 307-308 (1993 publ. 1994). Mi L, Lu A, Zhang L. (Studies on tales, 681-708. An. Jard. Bot. Madrid removed to facilitate editing. Rodrigues: 136. Myoporacees a Garden (London) 118(8): 362-363 the components of essential oils 51(2): 279-280 (1993 publ. 1994). 148. Hydnoracees. Mauritius: (1993). COMANTHOSPHACE from Elsholtzia spledens Nakai ex Efimova VA. Ob osobennostyakh Sugar Industry Research Institute Bhatti GR, Ingrouille MJ. The F Maekawa.) J. Wuhan Bot. Probatova NS, Sokolovskaya AP. prostranstvennoi evolyutsii semeist- 32p. BECIUM systematics of Comanthosphace S. Res.11(1): 94-96 (1993). Chisla khromosom nekotorykh vidov va Labiatae. In Tikhomirov VN, Balkwill K, and others. Specific Moore (Lamiaceae). Fontqueria 42: sosudistykh rastenii Rossiiskogo ed. Filogeniya i sistematika rastenii: Sikarwar RLS, Kaushik JP. Wall concepts in some Acanthaceae and in 1-10 (1995). Shimizu T, Imai K. (Occurrence DalnegoVostoka. (Chromosome mater.8 Mosk. soveshch. po filo- flora of Gwalior Fort, Madhya Becium (Labiatae). In Seyani JH, of Elsholtzia splendens Nakai ex F. numbers in some species of vascular genii rast. Moskva: Nauka, 1991 Pradesh. J. Econ. Taxon. Bot. 17(3): Chikuni AC, eds. Proceedings of the DICERANDRA Maekawa in Nagano Prefecture, plants from the Russian Far East.) pp.35-36. 627-633 (1993). 13th plenary meeting of AETFAT, Miller HA. A new species of central Japan.) J. Phytogeogr. Bot. Zhurn. 80(3): 85-88 (1995). Zomba, Malawi, 2-11 April, 1991: Dicerandra (Lamiaceae) from Flori- Taxon. 42(1): 39-41 (1994). Hedge IC. A plain man’s guide to Wu CY, Raven PH, eds. Flora of volume 1. Plants for people. Zomba: da. Phytologia 75(3): 185-189 Singh TP. Alterations in the basic generic groupings. Lamiales Newsl. China: volume 17. Verbenaceae National Herbarium and Botanic Gar- (1993). ENDOSTEMON chromosome numbers as a means of no.1: 1-3 (1992). through Solanaceae. Beijing: Sci- dens of Malawi, 1994 pp.347-358. Paton A, and others. A revision of speciation in Labiatae. Feddes ence Press; St. Louis, Mo.: Mis- DRACOCEPHALUM Endostemon (Labiatae). Kew Bull. Repert. 106(1-2): 39-47 (1995). 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Gard. 81(4): Schultze W, and others. Volatiles Jancic R. Anatomske osobine bil- Econ. Bot. 48(1): 60-64 (1994). flora: Nepeta parviflora M. Bieb.) ed species of Origanum. Kew 653-686 (1994). in flowers of balm (Melissa offici- jaka roda Mentha L. (Lamiaceae). Bot. Kozlem. 80(2): 99-102 (1993). Mag.11(3): 109-117 (1994).

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(Lami- divinorum (Labiatae). Sida 15(3): parative morphology in Schnabelia Ghazanfar SA. Novitates from the comb. nov. (Lamiaceae). Taiwania tematic implications of pollen mor- aceae): description de deux especes 349-366 (1993). of Chinese endemic plant and its flora of the Sultanate of Oman. Edin- 40(1): 13-19 (1995). phology in tribe (Labiatae). Syst. nouvelles du Bresil. (Contribution to systematic position: 2. Leaf archi- burgh J. Bot. 51(1): 59-63 (1994). Bot. 18(4): 563-574 (1993). the systematics of Salvia L. subg. Scholz S. Nuevos datos acerca de tecture in Schnabelia and its affinite PHYLLOSTEGIA Calosphace (Benth.) Benth.: descrip- Salvia herbanica Santos et Fernan- genera.) Acta Bot. Bor. Occid. Sin. PANZERINA Ryding O. Pericarp structure in the ROSMARINUS tion of two new species from Brazil.) dez (Lamiaceae). (New data on 15(2): 110-115 (1995). Krestovskaya TV. K nomenkla- tribe Prasieae (Lamiaceae: Ubera Jimenez JL, Hidalgo Fer- Bull. Mus. Nation. Hist. 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