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Galium Aparine 56 CLIV. RUBIACEAE – RUBIEAE 8. Galium F. Ehrendorfer & F.X. Krendl [cf. Tutin & al. (eds.), Fl. Eur. 4: 12 (1976)]. Sin embargo, al realizar el actual tratamiento del género, no se ha localizado mate- rial alguno en los herbarios estudiados. A. tinctoria L., Sp. Pl.: 104 (1753). De esta especie solo se conoce material peninsular de procedencia muy dudosa, en un caso de las inmediaciones de La Maladeta (Huesca; Herbario General de M. Rivas Mateos, MAF 49870), y en otro, de Santander [MA 147889; cf. Laínz & al. in Bol. Inst. Estud. Asturianos, Supl. Ci. 6: 65 (1963)]. 8. Galium L. [nom. cons.]* [Gálium, -ii, n. – gr. gálion, -ou n.; lat. galion, -ii n. = en Dioscórides, una planta que recibió tal nom- bre porque hace cuajar la leche [gr. gála, gálaktos n.], con hojas semejantes a las de la aparínē –el amor de hortelano (Galium Aparine L., Rubiaceae)– y flores amarillas; se supone que se trata del cuajaleche (Galium verum L.); en Plinio, sería otro nombre de la galeopsis o galeobdolon, planta que tendría las hojas semejantes a las de la urtica –la ortiga (Urtica sp. pl., Urticaceae)– pero que, al ser restregadas, emitirían fuerte olor, a más de que posee flores purpúreas; se ha pensado que ahí se habla de Lamium purpureum L. (Labiatae) o de Scrophularia peregrina L. (Scrophulariaceae)] Sufrútices o hierbas perennes, a veces cespitosas o pulvinulares, a menudo estoloníferas o rizomatosas, con frecuencia trepadoras, o bien hierbas anuales. Tallos simples o ramificados, glabros y lisos o bien retrorso-escábridos, retrorso- escabriúsculos, papilosos o variadamente pelosos, con nudos por lo general con glándulas mazudas hacia la parte adaxial de las bases foliares. Hojas formando con las estípulas verticilos de 3-11(12), sésiles, atenuadas o apenas pecioladas, con 1 ó 3 nervios, glabras o con indumento variable, con márgenes lisos, retror- sos o antrorso-escábridos, escabriúsculos, papilosos o variadamente pelosos, a veces con glándulas epidérmicas hinchadas y alargadas hacia la parte distal del envés. Inflorescencia subumbeliforme, terminal, obovoide, con (2)3 ramas, o bien paniculiforme, obovoide, ovoide, oblonga o piramidal, multiflora, con ra- mas opuestas, menos frecuentemente en cima simple o compuesta y pauciflora, o bien cilíndrica, con flores o cimas geminadas axilares; inflorescencias parcia- les cimosas, en glomérulo, corimbiformes o umbeliformes, pedunculadas o sési- les, bracteadas, las de último orden con 2-3 flores, rara vez más. Brácteas en las inflorescencias de primer orden en verticilos de (1)2-11(12), foliáceas, disminu- yendo progresivamente de tamaño. Flores hermafroditas, rara vez también unise- xuales masculinas –plantas andromonoicas– o estériles, actinomorfas, homóga- mas o proterandras, tetrámeras –excepcionalmente trímeras, pentámeras o hexá- meras–, ± pediceladas, con menor frecuencia sésiles, sin bractéolas o, si existen, no se dan en todas las flores o cimas. Cáliz inexistente. Corola rotácea –a veces con lóbulos erectos–, acopada, craterimorfa, campanulada, tubulosa o infundibu- liforme, con (3)4(6) lóbulos agudos –rara vez obtusos o cuculados–, a veces api- culados, por lo general papilosos en la cara adaxial y en ocasiones con glándulas epidérmicas cerca del ápice de la cara abaxial, glabra o pelosa, blanca, con me- nor frecuencia amarillenta, verdosa, rosada, rojiza o violeta. Androceo con (3)4(5) estambres alternipétalos; filamentos glabros, insertos cerca de las escota- * A. Ortega Olivencia & J.A. Devesa CLIV. RUBIACEAE – RUBIEAE 57 8. Galium duras de la corola; anteras por lo general oblongas, dorsifijas o casi. Estilo 1, con 2 ramas generalmente subiguales, a veces dividido desde la base aparentando ser doble, muy excepcionalmente con 3 ramas, rodeado en la base por 2 nectarios carnosos anulares; estigmas globosos, capitados u oblongos. Fruto en esquizo- carpo, con (1)2 mericarpos. Mericarpos secos, oblongos, ± globosos, subrenifor- mes u ovoides, a veces cilíndricos y ± curvos al madurar o elipsoidales, glabros y con superficie lisa, rugosa, papilosa, granulosa, mamilífera o escabriúscula, o bien homogénea o heterogéneamente pelosos, con pelos setoso-uncinados, unci- nulados, híspidos, pubérulos, hirsutos o vilosos. x = 10, 11, 12. Observaciones.–El género comprende unas 400 especies [entre 600 y 700 según E.P. Phillips, Gen. S. Afr. Fl. Pl. ed. 2: 739 (1951)] distribuidas principal- mente por las regiones templadas. Aunque gran parte de sus especies son hierbas perennes que poseen estolo- nes o rizomas de interés para la multiplicación vegetativa, la reproducción se- xual constituye su principal mecanismo reproductor. En su gran mayoría son plantas hermafroditas, rara vez andromonoicas (v.gr., G. tricornutum y G. verru- cosum), presentando flores en ocasiones olorosas (es frecuente el olor a cumari- na, por ejemplo en G. odoratum, G. verum yG. uliginosum) y con mucha más frecuencia nectaríferas. La homogamia es frecuente (v.gr., G. tricornutum), pero también se conoce en el género la dicogamia, que se manifiesta bien por un movimiento de los es- tambres (al principio erectos y entonces estigmas convergentes, después curva- dos hacia atrás y estigmas divergentes, orientándose hacia abajo entre los lóbu- los de la corola), en el caso de las especies con corola rotácea o poco profunda, bien por un crecimiento del estilo a lo largo de la vida de la flor, en el caso de flores ± largamente tubulosas (v.gr., G. boissierianum, G. moralesianum). Sin embargo, no parece que dichos mecanismos impidan la autopolinización, sobre todo en presencia de los polinizadores. De manera puntual se ha indicado la existencia de cleistogamia y geitonogamia, sobre todo en especies gregarias, y se estima que la autogamia es el principal mecanismo de reproducción sexual en las especies anuales de flor pequeña. Tan solo se conoce un caso aislado de autoincompatibilidad (G. saxatile). Los polinizadores más frecuentes son dípteros (Bombylidae, Sirphidae, Mus- cidae, entre otros), lepidópteros, coleópteros y pequeños himenópteros, y entre los agentes dispersantes destacan los vertebrados en las especies con mericarpo peloso, sobre todo de tipo uncinado o uncinulado, conociéndose un caso de mir- mecocoria [G. circaezans Michx., Fl. Bor.-Amer. 1: 80 (1803)] de América. El género comprende algunas especies útiles como aromatizantes de bebidas y tabaco, sucedáneos del café (v.gr., G. aparine, en Irlanda), para la elaboración de quesos, fuente de colorantes (v.gr., G. verum, colorante rojo a partir de sus raí- ces), por su interés medicinal o incluso como ornamentales (v.gr., G. odoratum). En las descripciones, el grosor de los tallos se refiere al que poseen entre án- gulos consecutivos en el tercio basal. El término “hojas” hace referencia a los órganos foliáceos con independencia de su naturaleza (hojas o estípulas), y su número en cada verticilo se refiere siempre a los nudos de los tallos principales, salvo indicación en contra. El tamaño se ha medido en las hojas de los vertici- 58 CLIV. RUBIACEAE – RUBIEAE 8. Galium los medios y superiores de los tallos principales (nunca en los axilantes de la in- florescencia), y se corresponde con el de las piezas más largas. Respecto al in- dumento, de gran importancia taxonómica, cabe señalar que el correspondiente a los márgenes debe observarse en la cara abaxial. En relación con la inflorescencia general, se reconocen varios modelos bási- cos: a) paniculiforme, de forma piramidal, oblonga o ± cilíndrica, con un eje principal en el que se disponen las ramas, por lo general opuestas, partiendo de la axila de las brácteas de primer orden; cada rama, a su vez, está ± desnuda en su parte inferior (pedúnculo) y suele portar inflorescencias parciales paniculi- formes o cimosas, que son simples o más generalmente compuestas. Las inflo- rescencias de último orden son cimosas, por lo general laxas y pedunculadas –aunque a veces forman glomérulos–, con 2-3(4) flores, y cuando el número es mayor entonces adoptan forma umbeliforme o corimbiforme; b) umbeliforme o subumbeliforme, obovoide u ovoide, con 2-3(4) radios o ramas que parten del eje principal, desnudas en su parte inferior (pedúnculos) y portadoras de una cima compuesta y multiflora o rara vez simple; c) cima terminal simple, con 2 ó 3 flores, o compuesta y pauciflora; d) cilíndrica y oblonga, con un eje principal sobre el que se disponen (1)2 cimas axilares por nudo, opuestas, simples –a ve- ces reducidas a una sola flor o a una flor más una cima geminada– o compues- tas. En los tipos a y b pueden aparecer por debajo de la inflorescencia general una o varias ramas floríferas unilaterales a partir de sucesivos verticilos. Respecto a las flores, conviene señalar que el grosor de los pedicelos se ha medido hacia su parte media, y que el tamaño de aquéllas se basa en medidas to- madas de flores hidratadas o conservadas en alcohol, incluyendo en el tamaño de los lóbulos de la corola el apículo cuando existe. La longitud del estilo se refiere a flores postantésicas o viejas. En relación con las medidas de los mericarpos (lon- gitud máxima), conviene señalar que no incluyen el indumento cuando existe. Bibliografía.–M.C. BAILLET & P.M.É. TIMBAL-LAGRAVE in Mém. Acad. Sci. Toulouse ser. 5, 6: 217-251 (1862); J.I. BRIQUET & F.G. CAVILLIER in É. BURNAT, Fl. Alpes Marit. 5: 116-171 (1913-1915); A. DONNEAUX in Lejeunia 103: 1-74 (1981); F. EHRENDORFER in Oesterr. Bot. Z. 105: 212-228 (1958) [sect. Jubogalium]; in Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1, 169: 407-421 (1960) [sect. Leptogalium]; in Bot. J. Linn. Soc. 70: 2-6 (1975); F. EHRENDORFER & al. in Pl. Syst. Evol. 202: 121-135 (1996) [sect. Leptogalium y sect. Leiogalium]; F. EHRENDORFER & E. SCHÖNBECK-TEMESY in P.H DAVIS (ed.), Fl. Turkey 7: 767-849 (1982); A. JORDAN, Observ. Pl. Nouv. 3: 94-187 (1846); E. KLIPHUIS in Lagascalia 11: 229-244 (1983); in Mem. Soc. Brot. 27: 77-87 (1984); F. KRENDL in Oesterr. Bot. Z. 114: 508-549 (1967) [gr.
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