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JABG02P001 Munir JOURNAL of the ADELAIDE BOTANIC GARDENS AN OPEN ACCESS JOURNAL FOR AUSTRALIAN SYSTEMATIC BOTANY flora.sa.gov.au/jabg Published by the STATE HERBARIUM OF SOUTH AUSTRALIA on behalf of the BOARD OF THE BOTANIC GARDENS AND STATE HERBARIUM © Board of the Botanic Gardens and State Herbarium, Adelaide, South Australia © Department of Environment, Water and Natural Resources, Government of South Australia All rights reserved State Herbarium of South Australia PO Box 2732 Kent Town SA 5071 Australia J. Adelaide Bot. Gard. 2(1): 1-138 (1979) A TAXONOMIC REVISION OF THE GENUS PITYRODIA (CHLOANTHACEAE)* Ahmad Abid Munir State Herbarium, Botanic Gardens, North Terrace, Adelaide, South Australia 5000 Abstract A taxonomic revision of the genus Pit.vrodia is provided. Dennisonia F. Muell, is reduced to synonymy. A key to the genera of the Chloanthaceae is provided. Forty-one species are recognized of which the following 16 new species and one variety are described: P. angustisepala, P. augustensis, P. byrnesii, P. chorisepala, P. gilruthiana, P. glabra, P. glutinosa, P. lanceo/ata, P. lanuginosa, P. megalophylla, P. ovata, P. puberula, P. pungens, P. quadrangulata, P. serrata, P. spenceri and P. exserta var. lanata. The new combination of P. ternifolia (Syn. Dennisonia tern(olia F. Muell.) is made and P. uncinata var. exserta Benth. is raised to the status of a species. The affinities and distribution are considered for the genus and each species. A key to the taxa is provided and a detailed description of each species is supplemented by a habit sketch of a flowering branch and analytical drawings of the flowers. Taxonomic History of the Genus The genus Pityrodia was described by Robert Brown (1810), with a single species, P. salvifolia, which he had collected himself in Queensland. It was referred to the Verbenaceae where it has been retained by the majority of botanists. Sprengel (1825) recorded this genus as a synonym of Premna L. which he placed in Linnaeus's "Didynamia Angiosperma" without reference to any family. Thesynonymous position for Pityrodia under the genus Premna was accepted only by Dietrich (1843) who referred it to the Verbenaceae. Poiret (1826) had, however, maintained Pityrodia asa distinct genus in the Verbenaceae. In 1828, Gaudichaud described a species of Pityrodia in a new genus Quoya, the type of which he had collected himself from Western Australia. It was also referred to the Verbenaceae where it was retained by Endlicher (1838, 1841), Meisner (1840), Spach (1840), Dietrich (1843), Walpers (1845), Lindley (1846), Schauer (1847), Bocquillon (1863) and Black (1870). Excepting Dietrich (1843), all the above named botanists maintained Pityrodia and Quoya as two distinct genera, thougha few of them listed the latter as a "genus dubium". Reichenbach (1828) also accepted Pityrodiaas a distinct genus which he referred to the tribe Verbeneae in the Verbenaceae, while treating Quo ya in the tribe ,Gesnereae of the Bignoniaceae. In 1830, Bartling transferred Pit yrodia to the tribe Viticeae (Verbenaceae). This new tribewas accepted for the genus by Lindley (1836), Spach (1840), Schauer (1847) and Bentham (1870). Subsequently, Endlicher (1838) placed it in the tribe Lantanae in the Verbenaceae, and this was followed by Meisner (1840), Endlicher (1841) and Walpers (1845). In 1839, Endlicher described two Pityrodia species in a further new genus Dasymalla, which he placed in the family Myoporaceae. This was followed by Meisner (1840), Spach *The present treatment of the genus Pityrodia concludes the series of taxonomic revisions in thefamily Chloanthaceae (= Dicrastylidaceae) (Munir, 1976-1978). Airy Shaw (1966, 1973) andsome others included in this family the albuminous seeded African genera Acharitea Benth., Cyclocheilon Oliv. and Nesogenes DC. After examining representatives of these genera, the present author excluded them from the Chloanthaceae for the following two main reasons (see also Munir, 1978e, pp. 412-416)': (1) The ovules in thesegenera are attached near the base of the ovary and (2) The two bracteoles subtending each flower are absent. A key to thegenera of the Chloanthaceae is provided on page 3. A. A. Munir J. Adelaide Bot. Gard. 2(1) (1979) (1840), Endlicher (1841), Dietrich (1843), Walpers (1845), Lindley (1846), de CandoIle (1847) and Black (1870). Excepting Dietrich (1843) and de CandoIle (1847), all the above authors also retainedPityrodiaandQuoyaas distinct genera. In 1859, F. Mueller described a new genusDennisoniawith a single species, the type of which he had collected himself from Northern Territory. It was referred to the Verbenaceae where it has been retained by the majority of botanists. During present studies, however,Dennisonia F. Muell. has been found to be congeneric withPityrodiaand is therefore recorded here as a new synonym of the latter. From 1859 onwards, F. Mueller regardedPityrodiaas a synonym ofChloanthesand described 10 new species under it. In 1864 he described one new species underQuo yabut he later transferred it to the genusChloanthes.A few of F. Mueller's species have since been reduced to synonymy. Turczaninow (1863) also described threePilyrodiaspecies one each underChloanthes, QuoyaandPityrodia. Twoof these species are now in synonymy. Bentham (1870) reinstated the genusPityrodiaas distinct fromChloanthes, and transferred to it allChloanthesspecies described by F. Mueller and one of the two described by Bartling (1845). He listed QuoyaGaud. andDasyniallaEndl. as synonyms ofPityrodia,and this has been accepted by the majority of botanists. Moreover, he referredthisgenustothepredominantlyAustraliansubtribeChloanthinae ("Chloantheae") of the tribe Viticeae in the Verbenaceae, and described a new variety, P.une matavar.e.vserta,which is now raised to the status of a species. In 1876, Bentham & Hooker upgraded the subtribe Chloanthinae to the tribe Chloantheae, without altering the circumscription of its genera. This tribe was accepted for the genus by Bailey (1883, 1890, 1901, 1913), Durand (1888), Post & Kuntze (1904) and Lemée (1943). Also in 1876, a further new genusDepretnesndia,in the Labiatae, was described by F. Mueller for a new species ofPityrodia.The new genus was later recognized by F. Mueller (1889) as synonym,ous withChloanthes. Briquet (1895) reclassified the Verbenaceae and upgraded the tribe Chloantheae to a subfamily Chloanthoideae. The latter consisted of three tribes: Achariteae, Chloantheae and Physopsideae withPitvrodiain the tribe Achariteae. This classification was adopted by Briquet (1896), Dalla forre and Harms (1904) and Melchior (1964). In 1904, Diels & E. Pritzel revised the Western Australian Verbenaceae comprising only Bentham & Hooker's tribe Chloantheae. They subdivided the tribe into two subtribes namely Lachnostachydinae and Chloanthinae, placingPityrodiain the latter. The genus was subdivided into four sections:Brachysolenia, Chloanthopsis, Depretnesndiaand Eupityrodia.These sections are found to be artificial and unnatural groups, and are, therefore, not maintained in this revision. Gardner (1931) and Junell (1934) retained Pityrodiain Briquet's subfamily Chloanthoideae, but within the subfamily, Junell referred it to Bentham & Hooker's tribe Chloantheae, and Gardner placed it in Diels & E. Pritzel's subtribe Chloanthinae without mention of the tribe Chloantheae. Gardner also adopted the sections proposed within this genus by Diels & E. Pritzel (1904). Hutchinson (1959) raised the status of Bentham & Hooker's tribe Chloantheae to the family Chloanthaceae, which differed from Verbenaceae (s.str.) chiefly in the albuminous seeds. The new family for the genus was accepted by Bullock (1959, 1960), Takhtajan (1959, 1969), Eichler (1965), Symon (1969), Gardner (1972) and Munir (1965, 1966, 1977, 1978a, 1978b, 1978c). Also in 1959, Moldenke published a résumé of the world Verbenaceae and referredPityrodiaand allied genera to the family Stilbaceae. Within this family, the genus was placed in the subfamily Chloanthoideae, tribe Achariteae. Airy Shaw (1965) referred all genera of Australian Verbenaceae (s.lat.) with albuminous seeds to the family Dicrastylidaceae Drumm.ex Harv. (nom.nud.), a name mentioned incidently by Harvey (1855) but not validated. The family name Dicrasty- lidaceae, however, has been adopted for the "Australian Verbenaceae" with albuminous seeds by Airy Shaw (1966, 1973), George (1967, 1972), Beard (1970), Maconochie & 2 J. Adelaide Bot. Gard. 2(1) (1979) Revision of Pn.rrodia Byrnes (1971), Moldenke (1971) and some others. In the present revision, Pityrodia is accepted as belonging to the family Chloanthaceae. Within this family, it is referred to the tribe Chloantheae, a position previously given to it by Briquet (1895) under Verbenaceae. Key to the genera of the Chloanthaceae As this is the last paper in the series of revisions of the genera of the Chloanthaceae,a key to the genera has been prepared. Non-Australian genera generally regardedas belonging to this family have been excluded (see footnote on p. 1). la.Flowers zygomorphic, heteromerous; corolla mostly more or less 2-lipped or unequally 5-lobed; stamens 4 (Tribe Chloantheae) 6 b. Flowers actinomorphic or nearly so, isornerous: corolla regular or nearly so, 4-8-lobed; stamens 4-8 (Tribe Physopsideae) 2 .2a. Perianth 5-8-merous 4 b. Perianth 4-merous 3 3a. Flowers in dense spikes; stamens included; style entire (not lobed) (Munir, 1978e: 578) . Physopsis b. Flowers in more or less capitate clusters; heads solitary or in corymbose panicles; stamens scarcely exserted;
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