CASTRIIANTHEMUM VOGT & OBERPRIELER, A NEW OF THE COMPOSITAE-

by ROBERT VOGT & CHRISTOPH OBERPRIELER*

Resumen VOGT, R. & CH. OBERPRIELER (1996). Castrilanthemum Vogt & Obeiprieler, un nuevo género de Compositae-Anthemideae. Anales Jard. Bot. Madrid 54:336-346 (en inglés). Se estudia en detalle Pyrethrum debeauxii, una especie anual endémica del sudeste de España, y se compara con los géneros cercanos. Esta especie debe adscribirse a un nuevo género, monotípico, Castrilanthemum Vogt & Oberprieler, que se describe aquí, a la vez que se propone la combinación C. debeauxii (Degen, Hervier & É. Rev.) Vogt & Oberprieler. Palabras clave: Spermatophyta, Compositae, Castrilanthemum, taxonomía, cladística, España.

Abstract VOGT, R. & CH. OBERPRIELER (1996). Castrilanthemum Vogt & Oberprieler, a new genus of the Compositae-Anthemideae. Anales Jard. Bot. Madrid 54:336-346. Pyrethrum debeauxii, a poorly-known annual endemic to SE-Spain, was studied in detail and compared with related genera. It is found to represent a new monotypic genus, Castrilanthemum Vogt & Oberprieler. The new combination C. debeauxii (Degen, Hervier & É. Rev.) Vogt & Oberprieler is proposed. Key words: Spermatophyta, Compositae, Castrilanthemum, , cladistic, Spain.

Pyrethrum debeauxii was described by uncertainty of its afnnities within this tribe A. von Degen and J. Hervier (in HERVIER, already dates back to Degen & Hervier (in 1905) on the basis of material collected HERVIER, 1905: 99) who write in the original in Andalusia at the beginning of this century diagnosis: "Planta nana, certe annua (...) facie by Élisée Reverchon, who distributed the Anthemidis cujusdam, a congeneribus specimens in his well-known exsiccata acheniis ecoronulatis valde diversa et in "Plantes d'Espagne" in 1904 and 1905. To the genere paradoxa". HEYWOOD (1954), in his present day this rare endemic of SE Spain has revisión of Tanacetum subsect. Leucanthe- been recollected only once, by J. Leal Pérez- mopsis Giroux -later on recognized at generic Chao in 1978. level as Leucanthemopsis (Giroux) Hey- The tribal placement of Pyrethrum wood- placed P. debeauxii in the vicinity of debeauxii in the Compositae-Anthemideae this taxon, mentioning it under his 'species was never a matter of doubt, whereas the dubiae'. However, a few years later GALIANO

* Botanischer Garlen und Botanisches Museum Berlin-Dahlem. Konigin-Luise-Str. 6-8. D-14191 Berlin (Germany). R. VOGT & CH. OBERPRIELER: CASTRILANTHEMUM 337

& HEYWOOD (1960:172) stated: "Esta especie ie* option was used to detect the most constituye probablemente una sección nueva parsimonious trees, and strict consensus trees del género , pero no hay were obtained using the nelsen command. material suficiente para una investigación completa", and therefore placed it with or cióse to Leucanthemum. Later treatments of RESULTS AND DISCUSSION the so called -complex (HEYWOOD & HUMPHRIES, 1977), e.g. in In their recent generic monograph of the Flora Europaea (HEYWOOD in TUTIN & al., Compositae-Anthemideae, BREMER & 1976) as well as in the recent monograph of HUMPHRIES (1993) subdivided the tribe into -Anthemideae by BREMER & 12 subtribes which they considered to HUMPHRIES (1993) and of the whole family by represent predominantly monophyletic BREMER (1994), do not consider P. debeauxii assemblages of genera, although no attempt due to the lack of material available to them at was made in that paper to test the monophyly that time. of the erected subtribes. Some have to be To ascertain the taxonomic position of considered as provisional groups. This is Pyrethrum debeauxii within the Compositae- mainly due to the obvious lack of characters Anthemideae, we carried out a comprehensive having any cladistically useful significance morphological study. Special emphasis was on a level higher than generic groups (e.g. placed on morphological and anatomical Leucanthemum-group, Achillea-Anacyclus- features of the fruit, due to their fundamental Leucocyclus-group, Cotula-gioup), causing importance in the generic delimitation within the collapse of the proposed cladogram for the both the tribe and the Chrysanthemum- interrelationships of the 12 subtribes as complex. mentioned by BREMER & HUMPHRIES (1993: 90). Another consequence seems to be that several of the proposed subtribes have a core MATERIAL AND METHODS formed by a monophyletic group of genera surrounded by a considerable number of The study is based on herbarium satellite genera for which relationships to the specimens housed at B, BP, M, P and W and core group or definite subtribal membership prívate collections of J. Leal Pérez-Chao and are not resolved. Thus, it is difficult to place R. Vogt. taxa not included by BREMER & HUMPHRIES, For comparative anatomical studies such as Pyrethrum debeauxii, in the right achenes were taken from herbarium subtribe. Nonetheless, we consider the specimens and soaked in a mixture (6 : 1) of combination of T-shaped hairs, epaleate 10% aqueous solution of dioctyl sodium receptacles, 10-ribbed achenes with myxo- sulfosuccinate and 96% ethanol for two genic cells, and anthers with triangular apical days (PETERSON & al., 1978). Following appendages as sufficient evidence to include dehydration through a 20% incremental series P. debeauxii in the Leucantheminae sensu to absolute ethanol with changes every 24 BREMER & HUMPHRIES (1993). hours, achenes were infiltrated with To determine the position of Pyrethrum Technovit 7100 (Heraeus-Kulzer) for four to debeauxii within the subtribe Leucanthe- six weeks. After embedding, median- minae we re-ran the data set provided by transverse sections of achenes were made BREMER & HUMPHRIES (1993), using the using a rotary microtome. Three to five central block of columns of the data matrix micron thick sections were stained in (BREMER & HUMPHRIES, 1993: 138) with toluidine blue and mounted in Vitroclud additional 34 scores for P. debeauxii (see (Langenbrinck). table 1). We used Lepidophorum as the Cladistic analyses were undertaken with outgroup. Hennig86, versión 1.5 (FARRIS, 1988). The The cladistic analysis resulted in 30 most 338 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

TABLEl

DATA MATRIX FOR PYRETHRUM DEBEAUXII USED IN THE CLADISTIC ANALYSES TOGETHER WITH DATA GIVEN FOR THE ÁNTHEMIDEAE- LEUCANTHEMINAE m BREMER & HUMPHRIES (1993: 138, tab. 21, middle columns) Character number 11 1 11111111111 11 1 1111 in BREMER & HUMPHRIES 1 4532 784107860573368221236474026 (1993:78-79) 1153342224444253714564060258131959 Pyrethrum debeauxii 1100010111000??00000??000000000000

1 = presence; 0 = absence; ? = missing data or not applicable parsimonious cladograms with a cladogram of 30), either P. debeauxii alone, a Pyre- length of 48 steps, a consistency index of thrum-Nivellea-cl&de, or a Pyrethrum 0.70, and a retention index of 0.75. Figure 1A debeauxii-Phalacrocarpum-clade is found shows the strict consensus tree. Figure IB and within a pectinate sequence of genera starting 1C show two of the 30 most parsimonious with Nipponanthemum and Leucanthemella cladograms. It is obvious that only the so- and ending with the Leucanthemum group called Leucanthemum-group, consisting of (fig. 1C). Leucanthemum, Rhodanthemum, Mauranthe- It is interesting to note that most of the mum, Chlamydophora, Chrysanthoglossum, disturbances leading to the partial collapse of Glossopappus, Coleostephus, and Plagius, is the consensus tree are caused by the position supported as a monophyletic assemblage of of Leucanthemopsis. In 20 of the 30 most genera by all of the cladograms, while the tree parsimonious trees this genus represents the structure beyond this subclade is unresolved sister taxon of the Leucanthemum-group, and the position of Pyrethrum debeauxii while in the remaining trees it forms a remains unclear. As pointed out by BREMER & monophyletic group with Hymenostemma HUMPHRIES (1993), VOGT (1991), and earlier and Prolongoa, due to the joint occurrence of by BRIQUET (1916), the members of the rather a flimsy pappus (character 165) in these three well defined Leucanthemum-group are all genera. BREMER & HUMPHRIES (1993) characterized by their specialized achene wall considered this character to be more with vallecular resin ducts and vascular important than differences in life habit and strands. This group of genera forms the core colour of involucral bract margins and of the subtribe Leucantheminae. consequently felt justified in accepting a Concerning the position of Pyrethrum cladogram one step longer than the most debeauxii, the 30 most parsimonious trees parsimonious one. If we accept their may be subdivided into the following two argument and give double weight to this scenarios: in most of the trees (18 out of 30) character, our analyses produce 10 most P. debeauxii forms a monophyletic group parsimonious trees (similar to the cladogram with Nivellea and Phalacrocarpum (as shown presented in fig. IB), all of them having a in fig. IB), partly being the sister taxon of Ni- Hymenostemma - Prolongoa - Leucanthe- vellea (based on its annual habit), partly the mopsis- and a Nivellea-Phalacrocarpum- sister taxon of Phalacrocarpum (based on its Pyrethrum defeeaMxn-subclade besides the involucral bracts with dark brown margins). Leucanthemum-group, with Phalacrocarpum This monophyletic group is characterized by as the sister group of Pyrethrum debeauxii in the apomorphic character state "pappus all cases. absent in ray and disc achenes" which occurs It becomes obvious that Pyrethrum debeau- in parallel in Leucanthemella also. In the xii possesses several characters of several remaining most parsimonious trees (12 out genera of the Leucantheminae which malees it R. VOGT & CH. OBERPRIELER: CASTRILANTHEMUM 339

Fig. l.-Strict consensus tree (A) and 2 of the 30 most parsimonious cladograms (B-C) of the Anthemideae- Leucantheminae after inclusión of Pyrethrum debeauxii. Solid bars indícate non-homoplastic synapomorphies; open bars indícate homoplastic synapomorphies with reversáis; double bars indícate parallelisms; crosses indícate reversáis. 340 ANALES JARDÍN BOTÁNICO DE MADRID, 54. 1996

Fig. 2,-Casírüanthemum debeauxii. Leal 263 (Herb. Vogt): A, stamens; B, apical appendages of the stamens; C, basal part of anthers and apical part of filamente; D, pollen grain; E, tip of style with the diverging branches. R. VOGT & CH. OBERPRIELER: CASTRILANTHEMUM 341

Fig. 'i.-Castrilanthemum debeauxii, Leal 263 (Herb. Vogt): A, achene; B, apical part of achene; C, basal detachment área of achene; D, transverse section of achene; E, detaii of transverse section of achene (me = myxogenic cell). 342 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

impossible to link this taxon to on of the al- medifixis, scariose fuscimarginata. Recep- ready existing genera. With its achenes taculum convexum, epaleaceum. Flores devoid of resin ducts it is clearly not a heteromorphi. Flores marginales ligulati, member of the Leucantkemum-growp which feminei, fértiles, albi; centrales infundi- forms the core of the Leucantheminae. It may bulares, hermaphroditi, fértiles, lutei sed be associated rather loosely with this group parte superiore purpurei. Stamina glabra, because of its 10-ribbed achenes with antherae basi rotundatae, filamento infra myxogenic cells (fig. 3). Most of the trees antheras túmida, connectivum supra obtained connect it with the two endemic antheram ovate dilatatum. Stylus teres, SW Mediterranean genera Nivellea and glaber, nectario basi cinctus, ramis 2 Phalacrocarpum, mostly because its ray and divaricatis, anguste oblongis, apice paulo disc achenes lack an apical corona. This penicillatis. Achaenia anguste obovoidea, character state is presumably apomorphic decemeostata, canalibus intercostalibus within the Leucantheminae and occurs in secretoriis carentia, cellulis myxogenis parallel in the Eurasian genus Leucanthe- epicostalibus instructa. Pappus nullus. mella. However, a relationship to the latter Chromosomatum numerus: 2n = 18 genus is unlikely because its achenes lack myxogenic cells. Pyrethrum debeauxii Typus: Castrilanthemum debeauxii differs from Nivellea in its possession of (Degen, Hervier & É. Rev.) Vogt & T-shaped hairs (fig. 4Bi) and a conical Oberprieler based on Pyrethrum debeauxii receptacle, and from Phalacrocarpum by its Degen, Hervier & É. Rev. annual habit and altérnate leaves. As BREMER & HUMPHRIES (1993) have pointed out, the Castrilanthemum debeauxii (Degen, subtribal position of Nivellea, and especially Hervier & É. Rev.) Vogt & Oberprieler, those of Phalacrocarpum and henee, also the comb. nov. position of Pyrethrum debeauxii, have to be = Pyrethrum debeauxii Degen, Hervier & considered as rather provisional. É. Rev. in Bull. Acad. Int. Géogr. Bot. 15: In line with efforts to circumscribe 99-100 (1905) [basion.] monophyletic units within the Leucanthe- Ind. loe: "In Hispaniae meridionalis minae, it is necessary to classify Pyrethrum provincia Jaén. In aridis calcareis montis debeauxii on the same taxonomic level as the Sierra de Castril alt. ce. 1800 mét. s. m. mense other units of this group. Therefore, the new majo 1903 rarissimum detexit el. É. Re- genus Castrilanthemum Vogt & Oberprieler verchon". is proposed. Lectotype (designated here): Élisée Castrilanthemum Vogt & Oberprieler, gen. Reverchon, Plantes d'Espagne 1904 nov.1 (Province de Jaén), n.° 1239, Pyrethrum debeauxii de Degen, Herv. Spec. nova!, Planta annua. Caules inferné foliati, pilis Sierra de Castril, lieux arides, sur le calcaire, medifixis dense obsiti. Folia petiolata, dense 1800 métres. Juin. Rare. (B!; isolectotypes in pilosa pilis medifixis, lamina pinnatisecta ad M!,P!). bipinnatisecta, segmentis ultimis brevibus, linearibus. Capitula solitaria pedunculata, -Pyrethrum debeauxii Degen & Hervier in radiata, heterogama. Involucri phylla É. Rev., Pl. Espagne 1904, n.° 1239 (1904), imbricata, triangulari-ovata vel oblongo- nom. nudum lanceolata, dorsaliter dense pilosa pilis Icones: HERVIER, 1905: 177.

1 Exsiccata: Élisée Reverchon - Plantes The ñame Castrilanthemum refere to the Sierra de Castril, a mountain range in SE Spain, where the new d'Espagne -1905, Province de Jaén, n.° 1239 genus occurs. (sub Pyrethrum debeauxii Degen & Herv., R. VOGT & CH. OBERPRIELER: CASTRILANTHEMUM 343

Fig. A.-Castrilanthemum debeauxii, Leal 263 (Herb. Vogt): A, habit; B, involucral bracts, from outermost to innermost, second bract from adaxial side; Bj, T-shaped hair from involucral bract; C, ray floret; D, tubular floret and achenes; E, stamens; F, style. 344 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

Fig. 5.-Castrilanthemum debeauxii, Leal 263 (Herb. Vogt): Outline of leaves, from base to peduncle, middle leaf with hair-covering.

Spec. nova.); Élisée Reverchon - Plantes d'Es- epaleaceous. Florets heteromorphic. Ray pagne - 1904 (Province de Jaén), n.° 1239 florets (fig. 4C) 7-15, female, fertile; limb (sub Pyrethrum debeauxii de Degen & Herv., white, revolute after anthesis, elliptical, Spec. nova!). broadly oblong or obovate, 7-12 mm long and 3.5-5 mm wide, apically 3-lobed, tubular part annual. Stems 4-10 cm high, solitary c. 1.5 mm long, laterally slightly winged. or branched at the base, erect or ascending- Tubular florets (fig. 4D), hermaphrodite, erect, green to glaucous, sulcate, reddish fertile, corolla 2.5-3 mm long, funnel-shaped, brown basally and on the ribs, covered with apically 5-lobed, yellowish with reddish T-shaped hairs, leafy in the lower part, lobes, abaxially slightly overlapping the peduncles 2-7 cm long, leañess. Leaves (fig. 5) altérnate (first leaves opposite and achenes. Anthers (figs. 2A-C, 4E) obtuse at often reddish brown), green to glaucous, the base, the filaments with a distinctly closely set in the lower part, broadly ovate enlarged collar, apical anther appendages to elliptical in outline, pinnatisect to ovate and blunt. Pollen tricolporate, spiny bipinnatisect with linear, minutely mucronate (fig. 2D). Style (figs. 2E, 4F) terete, slightly lobes, densely hairy with T-shaped hairs, swollen at the base, set in a conspicuous petiolate, petioles 3-7 mm long, slightly nectary, style branches truncate-penicillate at broadened at the base. Capitula terminal, the tips. Achenes (figs. 3A-E, 4D) narrowly long-pedunculate, solitary, 1.3-2.5 cm in obovoid, 2.5-3 mm long, 10-ribbed, with diameter, heterogamous and radiate. myxogenic cells along the ribs. Pappus Involucre hemispherical, 0.8-1.2 cm in absent. diameter (measured from pressed specimens). Chromosome number: 2n = 18. Involucral bracts (fig. 4B) imbrícate, in 4 rows, with broad brown to dark brown Flowering period: May-June. scarious margins, abaxially and marginally Distribution and habitat: Castrilanthe- densely hairy with T-shaped hairs (fig. 4B\); mum debeauxii is endemic to the NE córner of outermost bracts triangular; middle ones Andalusia in SE Spain (fig. 6). It is known ovate to oblong, 4-5 mm long and c. 1.5 mm from the Sierra de la Cabrilla in Jaén province wide, apically minutely mucronate; and the Sierra de Guillimona and the Sierra de innermost ones oblong or narrow obovate, Castril in Granada province. The plants grow rounded or mucronate, only distally with on dry stony slopes in scrub on limestone a brown margin. Receptacle convex, between 1700 and 1800 m. R. VOGT & CH. OBERPRIELER: CASTRILANTHEMUM 345

Fig. 6.-Distribution ofCastrilanthemum debeauxii in SE Spain according to material examined.

Cytology: The chromosome number was the only known localities for more than 70 reponed by LEAL PÉREZ-CHAO & al. (1980) years. It was collected again in 1978 by J. Leal who found 2« = 18, the most common number Pérez-Chao in the Sierra de Guillimona. All for diploids in the Compositae-Anthemideae. localities are localized in a small área at the More information concerning the formation eastern edge of Jaén and Granada provinces, of the karyotype is not available. demonstrating the restricted área and the scarcity of Castrilanthemum debeauxii. Remarles: Castrilanthemum debeauxii was discovered 1903 by É. Reverchon in the Lectotypification: Type locality is the Sierra de Castril in southern Spain (HERVIER, Sierra de Castril where Castrilanthemum 1905: 19) and forwarded for description to debeauxii was collected in May 1903 A. von Degen and J. Hervier who studied (HERVIER, 1905: 99). The plant material was Reverchon's extensive Andalusian plant distributed in 1904 by Reverchon as n.° 1239 collections. In 1905, Reverchon recollected of his exiccata "Plantes d'Espagne" under the species in the mountain ranges of the the ñame "Pyrethrum debeauxii de Degen, Sierra del Cuarto and the Sierra de la Cabrilla Herv. spec. nova!". Already in 1903 some (HERVIER, 1906: 207, 215). These remained unnumbered specimens of the collection were 346 ANALES JARDÍN BOTÁNICO DE MADRID, 54. 1996

distributed to O. Debeaux, A. von Degen and REFERENCES J. Hervier (HERVIER, 1905: 100) under the provisional ñame "Pyrethrum hispanicum BREMER, K. (1994). Asteraceae, Cladistics & Classification. Portland. Willk.?" The latter specimens, probably used BREMER, K. & C.J. HUMPHRIES (1993). Generic by Degen and Hervier for the original monograph of the Asteraceae-Anthemideae. Bull. diagnosis, have not been found in the herbaria Brit. Mus. (Nat. Hist.), Bot. 23:71-177. BRIQUET, J. (1916). Composées. In: J. Briquet & F. Ca- of the institutions which took over the prívate villier (eds.), Flore des Alpes Maritimes. Vol. 6(1). collections of these scientists. From the type Genéve, Bale & Lyon. material available in Berlin, Munich and FARRIS, J.S. (1988). Software & Manual. Hennig86 reference. Versión 1.5. New York. Paris, we designated as lectotype the GALIANO, E.F. & V.H. HEYWOOD (1960). Catálogo de specimen kept in the Botanical Museum plantas de la provincia de Jaén (mitad oriental). Jaén. Berlin-Dahlem (B). HERVIER, J. (1905). Excursions botaniques de M. Élisée Reverchon dans le massif de La Sagra et á Vélez- Rubio (Espagne) de 1899 a 1903. Bull. Acad. Int. Specimens seen Géogr.Bot. 15:1-32,57-72,89-120,157-178. SPAIN. GRANADA: Huesear, Sierra de Guillimona, HERVIER, J. (1906). Excursions botaniques de M. Élisée cerca del Cortijo de la Vidriera, matorrales en calizas, Reverchon dans le massif de La Sagra (Espagne) de 1700 m, 30SWH4010, 20-V-1978, /. Leal 263, Leal, 1904 á 1905. Bull. Acad. Int. Géogr. Bot. 16:201-221. HEYWOOD, V.H. (1954). A revisión of the Spanish Vogt. Flora Hispanica, Sierra de la Cabrilla, lieux andes, species of Tanacetum L. Subsect. Leucanthemopsis cale, 1800 m, VI-1905, É. Reverchon, M. JAÉN: Sierra de Giroux. Anales Inst. Bot. Cavanilles 12: 313-374. Castril, lieux arides, sur le calcaire, 1800 m, juin, HEYWOOD, V.H. & C.J. HUMPHRIES (1977). Anthemi- É. Reverchon, B, M, P. Sierra de Cabrilla, lieux arides, deae-systematic review: pp. 851-898. In: V.H. sur le calcaire, 1800 m, VI-1905, É. Reverchon, B, BP, Heywood, J.B. Harborne & B.L. Turner (eds.), The M, P, W. Sierra del Cuarto, lieux arides, sur le calcaire, Biology and Chemistry ofthe Compositae 2. London, 1800 m, VI-1904, É. Reverchon, BP. New York, San Francisco. LEAL PÉREZ-CHAO, J., A. ORTIZ VALBUENA, S. PAJARÓN SOTOMAYOR & M.L. RODRÍGUEZ PASCUAL (1980). ACKNOWLEDGEMENTS Números cromosómicos para la flora española, 155- 161. Lagascana 9:269-272. PETERSON, R.L., R.E. HERSEY & D.J. BRISSON (1978). We would like to thank Prof. Christopher Embedding softened herbarium material in Spurr's Humphries and Dr. Ilse Breitwieser for valuable resin for histological studies. Stain Technol. 53: 1-9. comments on the manuscript. The technical TUTIN, T.G., V.H. HEYWOOD, N.A. BURÓES, D.M. MOORE, assistance of Monika Lüchow and Jeannette D.H. VALENTINE, S.M. WALTERS & D.A. WEBB (eds.) Ueckert is gratefully acknowledged as well as the (1976). Flora Europaea. Vol. 4. Cambridge. VOGT, R. (1991). Die Gattung Leucanthemum Mill. drawing of the illustrations by Michael Rode- (Compositae-Anthemideae) auf der Iberischen wald. Halbinsel. Ruizia 10.