California, a new genus of geraniaceae endemic to the southwest of North America

CALIFORNIA, A NEW GENUS OF GERANIACEAE ENDEMIC TO THE SOUTHWEST OF NORTH AMERICA by JUAN JOSE ALDASORO', CARMEN NAVARRO2, PABLO VARGAS', LLORENC SAEZJ & CARLOS AEDO' 1 Real Ja din Bo inico, CSIC. Plaza de Mu illo, 2. E-28014 Mad id 2 Depai amen o de Biolog a Vege al II, Facul ad de Fa macia, Uni e sidad Complu ense. E-28040 Mad id 3 Depai amen o de Bo anica, Facul ad de Biolog a, Uni e sidad Au onoma de Bella e a. E-08193 Ba celona Resumen ALDASORO, J.J., C. NAVARRO, P. VARGAS, LL. SAEZ & C. AEDO (2002). Cali o nia, un nue o gene a de Ge aniaceae endemico del sudoes e de No eamenca. Anales la d. Bo . Mad id 59(2): 209-216 (en ingles). Los da os mo ol6gicos pe mi en dis ingui , a ni el de gene o, E odium mac ophyllum Hook. & Am. de las especies incluidas en E odium y Monsonia (Ge aniaceae). Tambien los da os ob enidos de la secuencia de ADN clo oplSs ico (»mL-F) apoyan es as di e encias. Po lo an o, p oponemos un nue o gene o mono pico, Cali o nia Aldas., C. Na a o, P. Va gas, Ll. Saez & Aedo, pa a que queden mejo e lejadas las elaciones ilogene icas en la clasi icacidn de las Ge aniaceae. Incluimos una diagnosis, ilus aciones, anilisis compa a i os de los ca ac e es di e enciales y una cla e de g6ne os de Ge aniaceae. Se p opone una nue a combinaci6n: Cali o nia mac ophylla (Hook. & Am.) Aldas., C. Na a o, P. Va gas, Ll. SSez & Aedo, comb, no . [basion.: E odium mac ophyllum Hook. & Am.]; y se designan dos lec d ipos. Palab as cla e: axonom a, mo ologia, Ge aniaceae, E odium, Monsonia, nue o gene o, No eameiica. Abs ac ALDASORO, J.J., C. NAVARRO, P. VARGAS, LL. SAEZ & C. AEDO (2002). Cali o nia, a new genus o Ge aniaceae endemic o he sou hwes o No h Ame ica. Anales la d. Bo . Mad id 59(2): 209-216. Mo phological da a p o ide e idence o he sepa a ion o E odium mac ophyllum Hook. & A n. in a new genus, di e en om ei he E odium and Monsonia (Ge aniaceae). Also cpDNA sequence da a ( nL-F) suppo his iew. Thus, we p opose he ecogni ion o a new mono ypic genus, Cali o nia Aldas., C. Na a o, P. Va gas, Ll. Saez & Aedo, o be e e lec phylogene ic ela ionships in Ge aniaceae. We p o ide diagnosis, illus a ions, compa a i e analysis o dis inc i e cha ac e s, and a key o gene a o Ge aniaceae. The new combina ion p oposed is: Cali o nia mac ophylla (Hook. & Am.) Aldas., C. Na a o, P. Va gas, Ll. Saez & Aedo, comb. no . [basion.: E odium mac ophyllum Hook. & Am.]; and wo lec o ypes a e designed. Key wo ds: axonomy, mo phology, Ge aniaceae, E odium, Monsonia, new genus, No h Ame ica. INTRODUCTION Ge anium L., Monsonia L,, Sa cocaulon (DC.) Swee and Pela gonium L'He . ex Ge aniaceae is a wo ld-wide amily com- Ai on (HUTCHINSON, 1969; KNUTH, 1912). p ising i e gene a: E odium L'He . ex Ai on, They ha e pen ame ous lowe s consis ing 210ANALES JARD N BOTANICO DE MADRID, 59(2) 2002 o i e sepals; i e pe als; i e an ipe alous nec a ies (Monsonia, Sa cocaulon, E odium and Ge anium), bu only one nec a y in Pe- la gonium; and oecia wi h a a iable numbe o s amens (15 in Monsonia and Sa cocaulon, 10 in Ge anium, 2-7 in Pela gonium and 5 in E odium), and consis en ly a gynoecium wi h 5 ca pels in all gene a (HUTCHINSON, 1969; KNUTH, 1912). S aminodes a e only p esen in wo gene a: 5 in E odium and a a iable numbe in Pela gonium, om 3 o 8 (STRUCK 6 VAN DER WALT, 1996). Mos Ge aniaceae s udies ha e emphasized he impo ance o he and oecium in amily classi ica ions, because i displays he main eliable cha ac e s (KNUTH, 1912). GENERIC DELIMITATION IN THE GERANIACEAE Gene ic delimi a ion o E odium has ne e se iously been ques ioned a e i was seg ega ed om Ge anium as o iginally ci cumsc ibed by Linnaeus (AITON, 1789; L'HERITIER, 1792). The majo di e ence be ween E odium and Ge anium is he e ile s amen numbe , en o ming wo who ls in Ge anium and i e in a who l plus a second who l o i e s aminodes in E odium. In bo h cases, s amens and s aminodes a e ee. Howe e , an excep ion is Ge anium pusillum L., which lacks one who l o e ile s amens (AEDO & al, 1998). The o he Ge aniaceae gene a {Monsonia and Sa cocaulon) ha e a who l o 15 s amens used in he base (monadelphous) o used in i e g oups o h ee (pen adelphous), whe eas Pela gonium has 2-7 monadelphous s amens. An excep ion is also ound in Monsonia b e i os a a Knu h, which has only i e e ile s amens (ALDASORO & al., 2001). The and oecium s uc u e o E odium could be ela ed o ha o Ge anium due o a likely subs i u ion o a who l o e ile s amens by s aminodes. The p o ologue o E odium mac ophyllum Hook. & A n. is somewha misleading, no allowing one o know he numbe o s e ile and e ile s amens: "This is a ue E odium, ha ing he i e s e ile s amens in he lowe ..." (HOOKER & ARNOTT, 1838). Su p isingly, E odium mac ophyllum Hook. & A n. has no s aminodes, bu has a qui e dis inc i e s aminal s uc u e: he ilamen s ha e a widely dila a ed basal po ion wi h wo la e al wing-like expansions apically ounded o almos au icula e ( ig. lc, d). The no mal na ow apical pa being apa en ly inse in he uppe hi d o his basal s uc u e ( ig. Id). This peculia s aminal ilamen has a ce ain esemblance o each o he i e g oups o s amens o M. b e i os a a (ALDASORO & al, 2001). Cha ac e s om lea es and ui s a e much mo e a iable wi hin he Ge aniaceae and usually no used o ecognize di e en gene a, bu hey se e o dis inguish in agene ic axa. Some o hem also suppo he seg ega ion o E. mac ophyllum om E odium. On he su ace o E odium me ica ps he e a e s ou b is les which may se e o acili a e bu ying, adhe ence and dispe sal ( ig. le; igs. 2A-D). A he base, E odium species ha e a nea ly semici cula im su ounding each b is le, which is o med by usion o papillae ( ig. 2D). I se es o p e en he b is le om e u ning and a oid unbu ying (EL-OQLAH, 1989; ALDASORO & al., 2001). The only E odium wi hou hese ims is E. mac o- phyllum ( ig. 2C). In he Ge aniaceae, simila ims a e only ound in some species o Monsonia (ALDASORO & al., 2001). The me ica p apex o many species o E odium has wo pi s which a e equen ly illed by glands. These pi s a e in mos E odium obliquely si ua ed wi h espec o he awn ( ig. 2B), whe eas hey a e almos pe pendicula ly disposed in E. mac ophyllum ( ig. le; ig. 2A). Monsonia has also hese wo ypes o pi s (oblique and pe pendicula ly placed) depending on he species (ALDASORO & a/., 2001). Finally, E. mac ophyllum lea es a e also dis inc i e. They a e ounded, wi h a co da e base and subpalma e eins ( ig. la, b), while all o he species o E odium ha e subpinna e o pinna e eins. Monsonia has bo h subpalma e (o palma e) and pinna e eins, depending on he species. As discussed abo e, E. mac ophyllum ea u es esemble in some cases E odium, in J.J. ALDASORO & AL.: CALIFORNIA, A NEW GENUS OFGERANIACEAE211 Fig. 1 -Cali o nia macwphylla: a, habi ; b, basal lea ; c, open lowe showing and oecium and gynoecium; d, s amen; e, ui and me ica p; , me ica p (based on Helle 7831 P). [a, b by LI. Saez; c- by J. Cas illo]. 212ANALBS JARDIN BOTANICO DE MADRID. 59(2) 2002 Fig. 2.-SEM mic og aphs o me ica ps o Cali o nia and E odium: A, C. mae ophylla me ica p apex (b, b is les; p, pi ) (based on Helle 7831 P); B, £. aciniulum me ica p apex (based on Rigual s.n., MA 371870); C, C. mac ophylla b is le base; D, K. a lan icum b is les base (based on Bull s.n, COI). J.J. ALDASORO & AL.: CALIFORNIA, A NEW GENUS OF GERANIACEAE213 o he s Monsonia. This, oge he wi h a unique and oecium ype ( igs, lc, d), leads us o seg ega e i in a di e en genus, especially aking in o accoun he and oecium, he mos eliable diagnos ic cha ac e in Ge aniaceae axonomy (KNUTH, 1912; YEO, 1990). Mo eo e , a p elimina y phylogeny o Ge aniaceae using nL-F sequences om ep esen a i es o he i e gene a plus E. mac ophyllum allow us o hypo hesize a ela i ely dis an ela ionship be ween his species and E odium. The wo mos - pa simonious ees esol e E. mac ophyllum as sis e o E odium, e en wi h low boo s ap suppo (< 50 %), and indica e an ancien di e gence be ween hem. KEY TO GERANIACEAE GENERA (see below o excep ions) in Ge anium sec . Ba achioidea W.D.J. Koch (AEDO & a/., 1998); he and oecium o M. b e i os a a (5 s amens) is a esul o educ ion o s amen numbe (ALDASORO & aL, 2001). As a as we know, he e is no in o ma ion on he phylogene ic posi ion o Pela gonium inca na um. Cali o nia Aldas., C. Na a o, P. Va gas, LI. S£ez & Aedo, gen. no . Typus: Cali o nia mac ophylla (Hook. & A n.) Aldas. & al. [E odium mac ophyllum Hook. & A n.] Di e ab q ini gen. E odium s aminodiis nullis, uniuscuiusque o eola e sus a is am pe pendicula i e disposi a e me ica pio um pilis bu sicula basali ca en ibus. No um genus, u pa e , nomen ahi e e is ubi habi a . This genus is simila o E odium, om 1. And oecium o 15 s amens 2 which can be dis inguished by he and oe- - And oecium o 2-10 s amens 3 cium, lack o s aminodes, me ica p pi pe - 2. Plan s succulen ; s ems co e ed wi h waxy pendicula ly disposed wi h espec o he awn, cu icle; longes pe ioles pe sis ing as spines b isde ims absem and lea es subpaima eiy Sa cocaulon eined - Plan s usually no succulen ; s ems no co e - „. ' . , . . ., . ed wi h waxy cu icle; longes pe ioles no We choose mls genenc ePl he 0 honou pe sis ing as spines Monsonia a Flons ic P o ince o Ame ica named 3. Flowe s wi h only a nec a y which is deeply Cali o nia, whe e his genus occu s, embedded in o he ecep acle ( a ely wi hou nec a y), gene ally zygomo phous; 2-7 mona- Cali o nia mac ophylla (Hook. & A n.) delphous s amens P la 8»n!»™ Aldas., C. Na a o, P. Va gas, LI. Saez & - Flowe s wi h i e nec a ies no deeply embedded A d h in o he ecep acle, ac inomo phous o sligh ly „,.' * ', ,, TI . „ » zygomo phous; 5 o 10 ee s amens 4 = E odium mac ophyllum Hook. & A n., 4. Flowe s wi h 10 e ile s amens Ge anium Bo - Beechey Voy.: 327 (1838), basion. - Flowe s wi h 5 e ile s amens 5 Ind. loc: "'Cali o nia, M . Douglas" 5. Flowe s wi h a who l o 5 s aminodes placed Typus: USA. Cali o nia, 1834, Douglas s.n. be ween he s amens; me ica ps wi h a semi- (lec o ype, he e designa ed, BM!; iso- ci cula im su ounding each b is le base lec o ype, GH) E odium - Flowe s wi hou s aminodes; me ica ps wi hou E odium cali o nicum G eene, Fl. F ancisc: a semici cula im su ounding each b is le base 99 (1891); E odium mac ophyllum a . Cali o nia cali o nicum (G eene) Jeps., Fl. W. Cali .: 247(1901) Some excep ions o Ge anium (G. pusil- Ind. loc: "Be keley Hills and eas wa d in he lum), Monsonia (M. b e i os a a), Pela go- M . Diablo Range" nium {P. inca na um) ha e no been included Typus: USA. Cali o nia, Be keley Hills, in he abo e key. P elimina y phylogenies 3 Ap . 1887, G eene s.n. (lec o ype, he e sugges ha G. pusillum and oecium (5 designa ed, NDG 28051; seeing colou s amens) is a de i a i e s a e om 10 s amens slide!) 214ANALES JARDIN BOTANICO DE MADRID, 59(2) 2002 Annual o biennial plan , sho ly caulescen o nea ly acaule. Roo s sho , sligh ly ami ied. S ems 10-40 cm long, ascending, hai y, wi h eglandula hai s 0.3-0.5 mm long and glandula hai s 0.4-0.6 mm long, all nea ly pa en o e o se, he glandula p oducing a eddish sec e ion. Lea es long pe iola e, ounded o co da e, c ena e-den a e, shallowly lobed ( a ely deeply lobed), wi h eglandula hai s 0.3-0.5 mm long and glandula hai s 0.5-0.6 mm long; sub- palma ely ne ed, wi h 1-2 pai s o seconda y ne es; basal lea es 11-43 x 10-44 mm ( a io lea wid h/lea leng h : 0.9-1.1); cauline lea es 10-33 x 8-32 mm; s ipules 3-4.1 mm long, o al-lanceola e, no memb anous, hai y, wi h glandula hai s 0.4-0.6 mm long along he ma gin and on he abaxial side. In lo escence 1-5 lowe ed, wi h peduncles a ising om he oo s ock; b ac s 2-5,3.2-5 x 1.3-1.6 mm, ee, acumina e, wi h a na ow sca iose ma gin, wi h eglandula hai s 0.2- 0.3 mm long on he abaxial side, glab ous adaxially; pedicels hai y, wi h eglandula hai s c. 0.2 mm long and glandula hai s 0.3- 0.6 mm long. Flowe s ac inomo phous, he maph odi e. Sepals 5-7.1 x 2.3-3.2 mm, awned, ac escen (11-14 x 4.3-5.6 mm in ui ), awn 0.8-1.5 mm long (0.9-1.3 long in ui ); abaxial side wi h c owded eglandula hai s 0.2-0.3 mm long and glandula hai s 0.5- 0.6 mm long. Pe als 6.2-8 x 2-3.2 mm, all simila , pink o whi e. S amen 5; ilamen 2.5- 3 mm long, wi h a glab ous, slende , ee pa (1.6-1.8 x 0.4-0.7 mm) inse ed on he uppe hi d o a dila a e basal po ion wi h wo la e al appendage-like wings; he basal po ion 2.3-2.6 x 1.3-1.4 mm, wi h he wo la e al wings apically au icula e, spa sely cilia e; an he s 0.8-0.9 mm long; pollen 35- 50 m in diam., e icula e, lumen o cells 3-5 m in diam.; wi hou sup a ec al p ocesses, icolpa e, yellow; s aminodes absen . Nec a ies 5, 0.4-0.5 x 0.5-0.6 mm, o al o ounded in ans e sal and longi udinal sec ions, adna e o he base o s aminal ilamen , colo unknown. Gynoecium 4.8- 5.6 mm long, densely hai y, wi h long glandula hai s; s igmas 0.8-1 mm long. F ui a schizoca p 44-54 mm in leng h; columella s ou ; awns no plumose (wi h sho hai s in he uppe pa ); me ica p leng h 9-12 mm (20-25 % o ui leng h); me ica p su ace wi hou papillae, wi h c owded b is les o a iable size, wi hou basal ims; apical pi p esen , pe pendicula ly si ua ed espec he awn, wi hou idges o u ows beside, wi h small glands inside, which a e also in he s a o awn. Seed 1 pe me ica p, 5-5.5 x 1.2- 1.4 mm, wi h ellip ic longi udinal sec ion; hilum leng h 2-2.4 mm (40-46 % o he seed leng h). Co yledon lea es o a e in ou line, wi h a co da e base. Ch omosome numbe . Unknown. ///K a OTW.TRELEASE(1888,pl. lO ig. 12); % 1. Phenology. Collec ed in lowe om Ma ch o he beginning o June. Ve nacula names. La ge-lea ed Fila ee (THOMAS, 1961) Dis ibu ion. Sou h O egon and Cali o nia (USA), and Baja Cali o nia (Mexico); also epo ed om A izona and S U ah (TAYLOR, 1993: 672); ig. 3. Habi a . Open habi a on iable clay soils; 50-1200 m. Rep esen a i e specimens examined USA. CALIFORNIA: Alameda Co., Al amon Pass, 37°44'N, 121°39'W, l-IV-1923, Ab ams 9946, CAS. Alameda Co., Co al Hollow, 37°32'N, 12 55'W, 7-IV-1940, Howe l 15393, CAS. Bu e Co., Chico, 39°44'N, 121°51'W, 1896, Aus in 984, P. Colusa Co., S ony o d Road, 39°22'N, 122°32'W, 23-IV-1926, Fe is 6406, CAS. Con a Cos a Co., An ioch, 38°0'N, 121°48'W, B andeque s.n., CAS. Con a Cos a Co., B en wood, 37°55'N, 121°41'W, 5-V-1892, Eas wood s.n., MO. Con a Cos a Co., nea Ki ke Pass, 37°57'N, 121°55'W, 10-V-1862, B ewe 1067, K. F esno Co., 14 mi W o Coalinga, 36°8'N, 120o21'W, 10-IV-1949, Wiggins 12040, CAS. Kem Co., 0.25 mi E o Whi e Ri e on G een ille oad, 35°8'N, 118°23'W, 30-111-1939, Fe is 9727, CAS. Ke n Co., Adobe Canyon, 33°31'N, 118°55'W, 26- V-1965, Twisselmann 10603, CAS. Ke n Co., Diablo Range, Co onwood Canyon, 35C5O'N, 118°42'W, 17-IV-1935, Wol 6441, CAS. Ke n Co., D ake Ridge, 35°8'N, 118°23'W, 28-IV-1952, Twisselmann 422, CAS. Ke n Co., Tehachapi, 35°7'N, llS^'W, 5-V-1905, Helle 7831, P, MO; ib dem, 13- V-1913, Eas wood 3251, CAS. Ke n Co., Temblo Summi , 35°3'N, 119°29'W, 20-IV-1966, Twisselmann 11974, CAS. Kings Co., Ree Ridge Canyon, 35°57'N, 120°13'W, 2-IV-1955, Twisselmann 1792, CAS. Kings Co., Ta Canyon, hills wes o A enal, 36° l'N, 120°8'W, 23-111-1940, Hoo e 4280, K. Los Angeles Co., Oso Canyon, 34°48'N, 118°43'W, 18-IV-1962, Twisselmann JJ. ALDASORO & AL.: CALIFORNIA, A NEW GENUS OF GERANIACEAE215 Fig. 3.-Dis ibu ion o Cali o nia mac ophylla based on examined specimens. 6959, CAS. Me ced Co., 37°10"N, 120°45'W, 23-IV- 1915, Eas wood 4428, CAS. Mon e ey Co., Pa k ield, 35°53'N, 120°25'W, 2-V-1937, Eas wood & Howell 4207, CAS. Mon e ey Co., San a Luc a M s., 36°3'N, 121°28'W, 24-V-1948, Unsdale s.n., CAS. Ri e side Co., Alice Mine, 34°3'N, 114°30'W, V-1807, Holl s.n., CAS. Sac amen o Co., Sac amen o alley, 38°35'N, 120°40'W, 1848, Ha ey 1675, K, W. San Beni o Co., 1 mi om Pe al a School nea Id ia, 36°25'N, 12 5'W, 28-IV-1940, Wiggins & Fe is 9391, CAS. San Beni o Co., 11 mi om He nandez, 36°22'N, 120°48'W, 29-IH- 1931, Howell 6026, CAS. San Beni o Co., 2.6 mi N o Id ia, 36°25'N, 121°5'W, 13-IV-1941, Wiggins & al. 41, CAS. San Beni o Co., nea San Beni o, 36°30'N, 121°4-W. 22-IV-1933, Howell 11084, CAS. San Beni o Co., Panoche, 36°35'N, 120°49'W, 3-V-1937, Eas wood & Howell 4286, CAS. San Diego Co., S o San Diego, 32°43'N. 117°8'W, 1875, Palme 5214, MO; ib dem, V- 1884, Cle elands.n., CAS; ib dem, 5-V-1905, Cle eland s.n., MO. San F ancisco Co., nea San F ancisco, 37°47-N, 122°24>W, 1868, Kellog & W. Ha o d 116, MO. San Joaqui'n Co., Hospi al Canyon, 37°37'N, 121°12'W, 18-IV-1938, Eas wood & Howell5089, CAS. San Joaqui'n Co., T acy, 37°44'N, 121°25'W, IV-1938, Bellne s.n., CAS. San Luis Obispo Co., 14 mi E o C es on, 35°31'N, 120°31'W, 21-IV-1949, Hoo e 7623, CAS. San Luis Obispo Co., Bi e wa e C eek, 35°6'N, 119°201W, 23-111-1965, Twisselmann 10496, CAS. San Luis Obispo Co., Choice Valley, 35O36'N, 120°9'W, 19-IV-1935, Wiggins 7959, CAS. San Luis Obispo Co., Cholame, 35"4yN, 120°17'W, 25-111-1935, Eas wood & Howell 2027, CAS. San Luis Obispo Co., C es on Road, 35°3 N, 120o31'W, 6-V-1952, Hoo e 8152, CAS. San Luis Obispo Co., Pain ed Rock, 35°8'N, 119°51'W, 21-IV-1941, Ripley & Ba neby 3283, CAS. San Luis Obispo Co., Pinole Sp ings, 35°29'N, 120°4' W, 7-V-1952, McMillan 117, CAS. San Luis Obispo Co., San Joaqu n Valley, 26-IV-1955, Twisselmann 1894, CAS. San Luis Obispo Co., Twisselmann Ranch, 35°39'N, 120°12'W, 16-IV-1954. Twisselmann 1066, CAS. San Ma eo Co., Pescade o, 37°36'N, 122°25'W, 7- IV-1896, Hudley s.n., CAS. Solano Co., Elmi a, 38o30'N, 121°36'W, 4-V-1886, Wallace s.n., CAS. Sonoma Co., Pe aluma, 38°15'N, 122°35'W. V-1880, Congdon s.n., CAS. S anislaus Co., A ec imba C eek, 37°33'N, 121°8'W, 24-IV-1937, Ewan 10220, CAS. S anislaus Co., hills wes o camp 76,37°33'N, 121°8'W, 16-VI-1862, B ewe 1280, K. Tehama Co., Vina, 39°55'N, 122°3'W. 29-IV-1917, S obe s.n., CAS. Yolo Co., Moon Ranch 7 mi W o Da is, 38°32'N, 121°44'W. 12-V-1955, C amp on 2773, CAS. OREGON: Jackson Co., nea Ashland, 42°11'N, 122°42'W, 15-IV-1889, Howell s.n., MO. MEXICO. BAIA CAUFORNIA: 20 m SW o Tia Juana, nea Rancho Cue as, 32°5'N, 116°50'W, 3-1V-1931, Wiggins 5123, CAS. A oyo de la Escope a, 30°42'N, 115°49'W, 3-VI-1975, Mo on 22375, CAS. ACKNOWLEDGEMENTS The au ho s hank M. Lainz o ende ing he La in diagnosis, and B. Baldwin and B. Ba olomew 216 ANALES JARD N BOTANICO DE MADRID, 59(2) 2002 o p o iding ma e ial sui able o DNA analysis. We also hank he cu a o s o BM, CAS, F, K, MO, NDG, P, W o he loan o specimens used in his s udy. This wo k was pa ly inanced by he Spanish Di ecci6n Gene al de In es igacidn Cien i ica y Tecnica (DGICYT) h ough he esea ch p ojec s PB96-0849, and REN200O-O818. REFERENCES AEDO, C, J.J. ALDASORO & C. NAVARRO (1998). Taxonomic e ision o Ge anium L., sec ions Di a ica a Rouy and Ba achioidea W.D.J. Koch (Ge aniaceae). Ann. Missou i Bo . Ga d. 85:594-630. AITON, W. (1789). Ho us Kewensis, ol. 2. London. ALDASORO, J.J., C. NAVARRO, P. VARGAS, & C. AEDO (2001). Ana omy, Mo phology, and Cladis ic Analysis o Monsonia L. (Ge aniaceae). Anales Ja d. Bo . Mad id 58:594-630. EL-OQLAH, E. (1989). A e ision o he genus E odium L'Heii ie in he Middle Eas . Feddes Repe . 100: 3-4. HOOKER, W.J. & G. ARNOTT (1838). The Bo any o Cap ain Beechey's Voyage. Hen y G. Bonn. London. HUTCHINSON, J. (1969). E olu ion and phy ogeny o lowe ing plan s. Academic P ess, London. KNUTH, R. (1912). Ge aniaceae. In: A. Engle (ed.), Das P lanzen eich. IV. 129 (He 53). Leizpig. L'HERITIER, C.L. (1792). Ge aniologia. Pa is, London, Vienna & S asbou g. STRUCK, M. & J.J.A. VAN DER WALT (1996). Flo al s uc u e and pollina ion in Pela gonium. The Biodi e si y o A ican Plan s. Kluwe Academic Publishe s. Ne he lands. TAYLOR, M.S. (1993). E odium. In: J.C. Hickman (ed.). The Jepson Manual. Uni . o Cali o nia P ess, Be keley, Los Angeles, London. THOMAS, J.H. (1961). Flo a o he San a C uz Moun ains o Cali o nia. S an o d Uni . P ess. S an o d, Cali o nia. TRELEASE, W. (1888). A s udy o No h Ame ican Ge aniaceae. Mem. Bos on Soc. Na . His . 4: 71-103, pi. 9-12. YEO, P. (1990). The classi ica ion o Ge aniaceae. In: P. Vo s e (ed.), P oceedings o he in e na ional Ge aniaceae symposium. S ellenbosch. Edi ado po Gines Lopez Gonzalez Acep ado pa a publicacidn: 10-XI-2001