Sion of Gypsophyla, Bolanthus, (Botanical Herbarium, Utrecht )
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Wentia 9 (1962) 1 —203 Sion of Gypsophyla, Bolanthus, Ankyropetalum and Phryna 1 (With Arabic summary) Y.I. Barkoudah (Botanical Museum and Herbarium, Utrecht) 2 (received March 6th, 1962) CONTENTS Introduction 2 GYPSOPHILA 4 A. General Part 4 1. Limits of the and revision of literature 4 genus 2. General morphology 9 the The root; the stem; the leaf; on the phyllotaxy of Gypsophila; found in inflorescence; remarks on the dichasial cymes Gypsophila-, the in the the aestiva- ontogeny of flower genus Gypsophila; calyx; tion of sepals; the corolla; aestivation of the corolla; the androecium; the and seeds. ovary; placentation; capsule 3. Cytology 26 4. Pollen morphology 28 5. Geography 29 6. Ecology 31 7. Uses 32 B. Taxonomic Part 33 1. Material 33 of the Diantheae 34 2. Diagnosis and key to the genera of the 35 3 Diagnosis and subdivisions genus 4. Key to the species 46 5. Species descriptions 55 BOLANTHUS 157 1. Diagnosis and delimitation from other genera 159 2. Geography 160 3. General morphology 160 other 162 4. Relation with genera 162 5. Key to the species 6. Species descriptions 163 ANKYROPETALUM 170 of the and discussion of the literature 171 1. Delimitation genus . 172 2. Key to the species 3. Species descriptions 173 PHRYNA 175 1. Diagnosis 176 x this a scholarship ) The author was enabled to carry out work by received from of the costs. Damascus University and by a special grant covering part publishing 2 ) Future address: Botanical Department, Damascus University, Damascus, Syria. 2 Y. I. BARKOUDAH 2. Taxonomic discussion 177 Species of uncertain status and names of uncertain application . 178 Excluded species ISO References 187 Index of exsiccatae 191 Index of names 198 INTRODUCTION Die ist Gattung schwierig zu umgrenzen, da sie sehr nahe Beziehungen zeigt zu Saponaria und Acanthophyllum. Pax und Hoffmann of Dr. This work was undertaken at the suggestion J. Lanjouw and Dr. F. P. Jonker. The citation heading this paragraph indicates that it the group of Caryophyllaceae with which deals, presents unusual taxonomic difficulties. At first, it was intended to restrict the revision to the in the of the work it genus Gypsophila. However, course was realized that the small . and genera Bolanthus, Ankyropetalum Phryna could not be left out of account as they had been regarded by some authors as subdivisions of Gypsophila and by others as near relatives of this For this revision of these genus. reason a complete genera too was included. The only previous revision of Gypsophila is that published by Williams (1889). His study, largely based on data derived from the literature, includes 76 species i.e. about Vsth of the number recognized those here. His views on the generic limits were strongly influenced by expressed by Bentham uiBentham and Hooker’s Genera Plantarum 1 (1862). Later authors did not follow him in this respect, and generally Boissier’s delimitation Pax and Hoffmann preferred (1867), so e.g. in the 2nd Edition of Engler und Prantl, Planzenfam. (1934). Pax had already accepted this delimitation in the first edition (1889). With the delimitation of the well its sub- regard to genus as as to divisions the author with Williams and present agrees largely (1889), less so with Boissier (1867). The sections Pseudoacanthophyllum Boiss. and included in Jordania Boiss., by Boissier Gypsophila, are transferred here to Acanthophyllum. The section Bolanthus Boiss., included in Gypsophila by Boissier and in Saponaria by Williams, is treated work with in the present as a separate genus; the same is done Ankyropetalum, which Williams included in Gypsophila. The sub- section Vittatae Williams (of the section Macrorrhizaea Boiss., which is regarded in this work as a subgenus of Gypsophila ), is transferred here to Saponaria. Gypsophila montana Balf. is transferred to Sapo- naria; iG. alsinoides Bunge to Arenaria. In Williams’ revision 76 (1889) Gypsophila species were recog- of these 8 excluded in the Of those nized; are present paper. species which were described afterwards, 31 are recognized here; with the 19 species described in this work this makes 50; the inclusion of GYPSOPHILA, BOLANTHUS, ANKYROPETALUM AND PHRYNA 3 these 50 made 2 species it necessary to distinguish new sections, 3 new subsections and 2 new series. In the revision therefore 126 present species of Gypsophila are Of these 19 are recognized. new; besides, 1 new subspecies and 2 new varieties are described. At the end follows a list of names of uncertain application. Out of the very many species of this group of genera, only nine, all of them native to Syria, were known to the author in the field. This the idea that it would be worth-while deficiency suggested to grow as many species as possible in the garden. Mr. E. A. Mennega, to whom the author is much indebted, succeeded in securing seed samples of 21 different species from other botanical gardens. Not all the seed samples which were received were correctly named; in misidentification fact, of Gypsophila species appeared to be more frequent in botanical gardens than in herbariums. Though the climatic conditions in the Netherlands are not favourable for growing Gypsophila species, the gardener of Cantonspark (Baarn) succeeded in bringing some of these species to flower. The section dealing with general morphology is based largely on material collected from these living plants. The author used this material also for studying the cytology of these However, this of the work species. part did not yield many in the results, first place because these plants were mostly well-known horticultural or else European species which had already been studied cytologically, and also because they represented only 2 sections of the " and because critical genus Gypsophila no species were among them. The author intends to continue in the future his investigations in this direction. It is that the revision will clear the hoped present help to up taxonomic of these The author position genera. realizes that some of the the relation of these with other problems concerning genera ~ members of the tribe Diantheae still remain to be solved. Cyto- taxonomic studies would presumably contribute valuable data. This the applies especially to genera Acanthophyllum, Bolanthus, Ankyro- and and which petalum Phryna to those Gypsophila species seem to these approach genera. Comparative ecological research too might help to understand the differentiation of the species. A study of this kind would be particularly interesting where closely related species are found to inhabit adjacent geographic areas. The author wishes his Acknowledgement. to express deep gratitude to Prof. Dr. J. Lanjouw, under whose supervision and in whose institute this work was carried out; to Prof. Dr. C. E. B. Bremekamp and to Dr. K. U. Kramer for their constructive criticism of the manuscript; to Dr. Elisabeth Mennega who corresponded with other herbariums in order to obtain the material on loan and who assisted him in his search for type specimens; to Dr. W. Punt for his assistance in the pollen morphological part. 4 Y. I. BARKOUDAH GYPSOPHILA A. GENERAL PART 1. Limits of the genus and revision of Literature in The name Gypsophila was mentioned for the first time Linne’s Diss. Chen.: 41 (1751). There, Linne described 7 species of Gypso- and phila, out of which one was afterwards removed to Arenaria another one to Tunica. Later, Linne described 2 other species, one of which was also a Tunica. That Linne failed to recognize the difference between Tunica and Gypsophila was due to the purely numerical basis of his system. Apparently, Linne did not consider Haller’s work, 1: 381 Stirp. Helv. (1742), where Tunica was regarded as a separate Scopoli Linne’s genus. (1772) revised work, and made the bound- aries between Gypsophila and Tunica sufficiently clear. Bentham in Bentham and Hooker (1862) made the difference quite pronounced the different subtribes of the tribe by referring two genera to two Sileneae; in the latter he distinguished three subtribes, viz. Diantheae (Tunica), Drypideae and Sileneae (Gypsophila), according to the seed and embryo shape. The mistake made by Desfontaink (1800), Sibthorp and Smith (1806), Seringe in DC. Prod. (1824), Bunge in Ledebour, FI. Ross. (1842), Grisebach (1843), Dufour J1860) and Janka (1867), by neglecting the differences between Tunica and Gypsophila, has very seldom been repeated in modern literature. The peltate seed with facial hilum, the straight embryo placed in the viz. with middle of the seed, the endosperm all around, i.e. also on the dorsal of side the seed, are characters which do not occur in any the habit and the Gypsophila species. Moreover, grass-like presence of characters stigmatic papilae all along the styles are typical Tunica which in are very rarely met Gypsophila. It is true that Gypsophila caricifolia Boiss. and Gypsophila graminifolia Bark, are in their habit more or less similar to Tunica, but fruits the flowers, and seeds of these two species are of the kind which is typical for Gypsophila. This suggests that these two in their lines those species developed vegetative organs on parallel to in which the vegetative parts of Tunica were differentiated; this how- there ever, does not mean that exists a near relation between these two genera. The difference between Gypsophila and Saponaria too was not clear to Linne, for he put Gypsophila porrigens (L.) Boiss. in Saponaria; it at an earlier date he had referred to Silene. Lamarck, FI. Fr. (1778) did not accept Gypsophila as a separate genus, and inserted its species in A. Braun that Saponaria. (1843) says though it is easy to separate from the other of the it is difficult Saponaria genera Lychnideae, very it from Bentham that to distinguish Gypsophila. (1862) says Gypso- and phila Saponaria are so intimately blended with each other that drawn no distinct boundary line can be between them, and some of the small-flowered species may according to him with almost equal right be referred to the one as to the other.