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Redalyc.California, a New Genus of Geraniaceae Endemic to The Anales del Jardín Botánico de Madrid ISSN: 0211-1322 [email protected] Consejo Superior de Investigaciones Científicas España Aldasoro, Juan José; Navarro, Carmen; Vargas, Pablo; Sáez, Llorenç; Aedo, Carlos California, a new genus of Geraniaceae endemic to the southwest of North America Anales del Jardín Botánico de Madrid, vol. 59, núm. 2, 2002, pp. 209-216 Consejo Superior de Investigaciones Científicas Madrid, España Available in: http://www.redalyc.org/articulo.oa?id=55659202 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative CALIFORNIA, A NEW GENUS OF GERANIACEAE ENDEMIC TO THE SOUTHWEST OF NORTH AMERICA by JUAN JOSÉ ALDASORO 1, CARMEN NAVARRO 2, PABLO VARGAS 1, LLORENÇ SÁEZ 3 & CARLOS AEDO 1 1 Real Jardín Botánico, CSIC. Plaza de Murillo, 2. E-28014 Madrid 2 Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense. E-28040 Madrid 3 Departamento de Botánica, Facultad de Biología, Universidad Autónoma de Bellaterra. E-08193 Barcelona Resumen ALDASORO, J.J., C. NAVARRO, P. VARGAS, LL. SÁEZ & C. AEDO (2002). California, un nuevo género de Geraniaceae endémico del sudoeste de Norteamérica. Anales Jard. Bot. Madrid 59(2): 209-216 (en inglés). Los datos morfológicos permiten distinguir, a nivel de género, Erodium macrophyllum Hook. & Arn. de las especies incluidas en Erodium y Monsonia (Geraniaceae). También los datos obtenidos de la secuencia de ADN cloroplástico (trnL-F) apoyan estas diferencias. Por lo tanto, proponemos un nuevo género monotípico, California Aldas., C. Navarro, P. Vargas, Ll. Sáez & Aedo, para que queden mejor reflejadas las relaciones filogenéticas en la clasificación de las Geraniaceae. Incluimos una diagnosis, ilustraciones, análisis comparativos de los caracteres diferenciales y una clave de géneros de Geraniaceae. Se propone una nueva combinación: California macrophylla (Hook. & Arn.) Aldas., C. Navarro, P. Vargas, Ll. Sáez & Aedo, comb. nov. [basión.: Erodium macrophyllum Hook. & Arn.]; y se designan dos lectótipos. Palabras clave: taxonomía, morfología, Geraniaceae, Erodium, Monsonia, nuevo género, Norteamérica. Abstract ALDASORO, J.J., C. NAVARRO, P. VARGAS, LL. SÁEZ & C. AEDO (2002). California, a new genus of Geraniaceae endemic to the southwest of North America. Anales Jard. Bot. Madrid 59(2): 209-216. Morphological data provide evidence for the separation of Erodium macrophyllum Hook. & Arn. in a new genus, different from either Erodium and Monsonia (Geraniaceae). Also cpDNA sequence data (trnL-F) support this view. Thus, we propose the recognition of a new monotypic genus, California Aldas., C. Navarro, P. Vargas, Ll. Sáez & Aedo, to better reflect phylogenetic relationships in Geraniaceae. We provide diagnosis, illustrations, comparative analysis of distinctive characters, and a key to genera of Geraniaceae. The new combination proposed is: California macrophylla (Hook. & Arn.) Aldas., C. Navarro, P. Vargas, Ll. Sáez & Aedo, comb. nov. [basion.: Erodium macrophyllum Hook. & Arn.]; and two lectotypes are designed. Key words: taxonomy, morphology, Geraniaceae, Erodium, Monsonia, new genus, North America. INTRODUCTION Geranium L., Monsonia L., Sarcocaulon (DC.) Sweet and Pelargonium L’Her. ex Geraniaceae is a world-wide family com- Aiton (HUTCHINSON, 1969; KNUTH, 1912). prising five genera: Erodium L’Her. ex Aiton, They have pentamerous flowers consisting 210 ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002 of five sepals; five petals; five antipetalous Erodium macrophyllum Hook. & Arn. has nectaries (Monsonia, Sarcocaulon, Erodium no staminodes, but has a quite distinctive and Geranium), but only one nectary in Pe- staminal structure: the filaments have a largonium; androecia with a variable number widely dilatated basal portion with two lateral of stamens (15 in Monsonia and Sarcocaulon, wing-like expansions apically rounded or 10 in Geranium, 2-7 in Pelargonium and 5 in almost auriculate (fig. 1c, d). The normal Erodium), and consistently a gynoecium with narrow apical part being aparently insert in 5 carpels in all genera (HUTCHINSON, 1969; the upper third of this basal structure (fig. 1d). KNUTH, 1912). Staminodes are only present in This peculiar staminal filament has a certain two genera: 5 in Erodium and a variable resemblance to each of the five groups of number in Pelargonium, from 3 to 8 (STRUCK stamens of M. brevirostrata (ALDASORO & & VAN DER WALT, 1996). Most Geraniaceae al., 2001). studies have emphasized the importance of Characters from leaves and fruits are much the androecium in family classifications, more variable within the Geraniaceae and because it displays the main reliable usually not used to recognize different genera, characters (KNUTH, 1912). but they serve to distinguish infrageneric taxa. Some of them also support the segregation of E. macrophyllum from Erodium. On the GENERIC DELIMITATION IN THE GERANIACEAE surface of Erodium mericarps there are stout bristles which may serve to facilitate burying, Generic delimitation of Erodium has never adherence and dispersal (fig. 1e; figs. 2A-D). seriously been questioned after it was At the base, Erodium species have a nearly segregated from Geranium as originally semicircular rim surrounding each bristle, circumscribed by Linnaeus (AITON, 1789; which is formed by fusion of papillae L’HERITIER, 1792). The major difference (fig. 2D). It serves to prevent the bristle from between Erodium and Geranium is the fertile returning and avoid unburying (EL-OQLAH, stamen number, ten forming two whorls in 1989; ALDASORO & al., 2001). The only Geranium and five in a whorl plus a second Erodium without these rims is E. macro- whorl of five staminodes in Erodium. In both phyllum (fig. 2C). In the Geraniaceae, similar cases, stamens and staminodes are free. rims are only found in some species of However, an exception is Geranium pusillum Monsonia (ALDASORO & al., 2001). The L., which lacks one whorl of fertile stamens mericarp apex of many species of Erodium (AEDO & al., 1998). The other Geraniaceae has two pits which are frequently filled by genera (Monsonia and Sarcocaulon) have a glands. These pits are in most Erodium whorl of 15 stamens fused in the base obliquely situated with respect to the (monadelphous) or fused in five groups of awn (fig. 2B), whereas they are almost three (pentadelphous), whereas Pelargonium perpendicularly disposed in E. macrophyllum has 2-7 monadelphous stamens. An exception (fig. 1e; fig. 2A). Monsonia has also these two is also found in Monsonia brevirostrata types of pits (oblique and perpendicularly Knuth, which has only five fertile stamens placed) depending on the species (ALDASORO (ALDASORO & al., 2001). The androecium & al., 2001). structure of Erodium could be related to that Finally, E. macrophyllum leaves are also of Geranium due to a likely substitution of a distinctive. They are rounded, with a cordate whorl of fertile stamens by staminodes. base and subpalmate veins (fig. 1a, b), while The protologue of Erodium macrophyllum all other species of Erodium have subpinnate Hook. & Arn. is somewhat misleading, not or pinnate veins. Monsonia has both allowing one to know the number of sterile subpalmate (or palmate) and pinnate veins, and fertile stamens: “This is a true Erodium, depending on the species. having the five sterile stamens in the flower...” As discussed above, E. macrophyllum (HOOKER & ARNOTT, 1838). Surprisingly, features resemble in some cases Erodium, in J.J. ALDASORO & AL.: CALIFORNIA, A NEW GENUS OF GERANIACEAE 211 Fig. 1.–California macrophylla: a, habit; b, basal leaf; c, open flower showing androecium and gynoecium; d, stamen; e, fruit and mericarp; f, mericarp (based on Heller 7831 P). [a, b by Ll. Sáez; c-f by J. Castillo]. 212 ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002 Fig. 2.–SEM micrographs of mericarps of California and Erodium: A, C. macrophylla mericarp apex (b, bristles; p, pit) (based on Heller 7831 P); B, E. laciniatum mericarp apex (based on Rigual s.n., MA 371870); C, C. macrophylla bristle base; D, E. atlanticum bristles base (based on Ball s.n., COI). J.J. ALDASORO & AL.: CALIFORNIA, A NEW GENUS OF GERANIACEAE 213 others Monsonia. This, together with a unique in Geranium sect. Batrachioidea W.D.J. androecium type (figs. 1c, d), leads us to Koch (AEDO & al., 1998); the androecium segregate it in a different genus, especially of M. brevirostrata (5 stamens) is a result of taking into account the androecium, the most reduction of stamen number (ALDASORO & reliable diagnostic character in Geraniaceae al., 2001). As far as we know, there is no taxonomy (KNUTH, 1912; YEO, 1990). information on the phylogenetic position of Moreover, a preliminary phylogeny of Pelargonium incarnatum. Geraniaceae using trnL-F sequences from representatives of the five genera plus California Aldas., C. Navarro, P. Vargas, E. macrophyllum allow us to hypothesize a Ll. Sáez & Aedo, gen. nov. relatively distant relationship between this Typus: California macrophylla (Hook. & species and Erodium. The two most- Arn.) Aldas. & al. [Erodium macrophyllum parsimonious trees resolve E. macrophyllum Hook. & Arn.] as sister to Erodium, even with low bootstrap Differt ab affini gen. Erodium staminodiis support (< 50 %), and indicate an ancient nullis, uniuscuiusque foveola versus aristam divergence between them. perpendiculariter disposita et
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