84 C. J. HUMPHRIES 



Synopsis 



The Mediterranean genus Anacyclus is revised. In all, nine species and three putative hybrids, grouped 

 into two sections, are recognized. The generic status and history of taxonomic treatments of Anacyclus 

 are discussed in relation to other genera of the Anthemideae. Phylogenetic relationships and the distribu- 

 tion of species are analysed according to the principles of Hennigian phylogenetic systematics. A key to 

 all species, subspecies, varieties and hybrids is given, each taxon is fully described, complete synonymies 

 are included and the relevant taxonomic characters are discussed in detail. Distribution maps for all 

 taxa and illustrations to the species are given, excluding only A. latealatus, a rare endemic from Turkey. 

 One new species and two new subspecies combinations are made, and the existence of three, previously 

 unrecognized, hybrids is elucidated. The account ends with a list of excluded taxa and a taxonomic index. 



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Introduction 



The genus Anacyclus belongs to the family Compositae tribe Anthemideae, which is restricted to 

 the Mediterranean region and is particularly well represented in the Maghreb countries. The 

 circumscription of the genus has been slightly reduced from that recognized by Jahandiez & 

 Maire (1934), to eliminate those species which belong to the Anthemis assemblage. 



The principal taxonomic problems in the genus are due mainly to the fact that not only are 

 several of the species extremely variable and closely related annual weeds with sympatric distribu- 

 tions, but the generic relationships are also not at all well known. This is one of two papers 

 dealing with the systematics and biology of Anacyclus (see also Humphries, in press, a). The 

 nomenclature and descriptive taxonomy are in need of critical revision, and so this has been 

 attempted in the present paper. In addition to the formal taxonomic treatment there are dis- 

 cussions of morphology, phytogeography and phylogeny, and to present the right context for 

 discussion of generic delimitation an historical account is also given. 



Materials and methods 



The revision is based partly on my own field studies, collections and cultivated material, which 

 are deposited at the British Museum (Natural History) (BM), and largely on herbarium material. 

 I was able to study five of the nine species in the field, four of which have been cultivated in the 

 greenhouses of Chelsea Physic Garden and used in experimental crossing studies (Humphries, 

 in press, a). I have studied material from the following herbaria (Index Herbariorum abbreviations 

 as in Holmgren & Keuken (1974): AV, B, BM, BR, C, E, FI, G, JE, K, L, LD, LE, M, MA, 

 MPU, P, RNG, S, W, WU, Z, ZT. 



The descriptions are based on both dried and living material, where available, and the variation 

 ranges cited attempt to cover the total variation exhibited by a particular species. Abnormal 

 values have been placed in parentheses either before or after the main range of variation. Flower- 

 ing periods, chromosome numbers, ecological data, locality lists and distribution maps have been 

 compiled almost entirely from specimens, and data from the literature has been included only 

 when substantiated by authentic material. One new species is recognized, two hybrids are des- 

 cribed, and several new combinations are made. 



Cross-sections of cypselas were made from my own collections, softened in water, embedded in 

 paraffin and ceresin wax, cut by microtome and stained in safranin combined with light green or 

 Clorazal Black E. 



The material examined is not normally listed after each species examined. A complete list of all 

 herbarium specimens seen has been placed in the library of the British Museum (Natural History). 

 Unlocalized material of any origin is omitted, except in the case of types and authentic historical 

 material. Formal citations are given for names and authentic records which extend knowledge of 

 ranges or taxonomy of the taxon in question. 



Descriptive terminology 



The descriptions and terms used in this work follow those outlined in Featherley (1954) and 

 Stearn (1966). The terminology for outlines and plane shapes adopted is that of the Systematics 



