Leaf morphology and anatomy of genus Androcymbium (Colchicaceae) in ~outhwestA h c a Absfmct N.,J. PEDROLA-MONWRT gl J. CAUJA&.CAS~LLS (2003). Lcaf morphology and anatomy of gcnus Androcymbiutn (Calehicaceae) in Swthwest Africa. Collecr. BOL {Burcelotur) MBMBRm, 26: 83-99. Morphologicai and matomical leaf studies wem made in 32 populations belonging to 17 laxa of the genus AndmcyntBirrm in Southwest Africa. The mo~hdogicalchatacters studied were conn md tunic chwacteristics, and number, distribution. shape, size. section. and color of leaves and braets. The micromorphological chmcters were the leaf indumcnt on both surface and margin, shape and size of epidemiic cells, type md arnount of stomata. Finally, the anatomical characters were mmphyll cell types, epidermic cel1 dxi, cellular wall types. size of central epidermic ctlls related to the other epidermic ceUs. and amount of idioblasis. Thc rtsuIts showed a great heterogencity in almst al1 characm rrnalyzed. Howevtr, macromorphological characteristics ltlated to color and shape of leaves and braets in the genus Andmcy~nbiun~ could be used as indicators of taxonomic afinities among species. Converse1y, thc micromorphological and anat o d a 1 chatilcteristics studied showed a great variation of types ihat does no1 agree with the interspecific mlationships cstablisbed from morpkologicai. allozymatic or cpDNA RFLP data in other research carried out within the genus. Key words: Andmcymbium, Colchicaceae, morphology, anatomy, -en M w ~ r v uN,.,J. P~ROLA-MONWRT & J. CAUJAP&CASTELLS (2003).Morfologla y anatomla dcl gbnero Andmcytnbium (CoIchicaceae} en Surafrica Occideninl. Collect. Bw. (Barcelona) 26: 83-99. Se realizaron estudios morfolbgicos y anatdmicos de las p m s vegetativas en 32 poblaciones pertenecientes a 17 taxones dcl gdnero Andmyrnbium en Surtífrica Occidental. LOScaractms mdoldgicos estudiados fueron Ias caraekrfsticas del owmo y be las tdnicas, y número, disulbuci6n. forma, tamafio, seccih, y color de las hojas y brácteas. Los caracteres micromorfoI6gicos estudiados fueron el indumento en la superficie y margen de la hoja. forma y m a n o dc las ctlulas epi&rmieas, y tipo y cantidad de estomas. Finalmente, los wactcres anat6micos fueron el tipo de cdlulu del mesofilo, tamaRo de las ot1uias epidérmicas, tipo de parcd celular, iamaíio de la9 células epiddmricascenaes comparado con el resto de céIulas de la lamina y cantidad de idioblastos. Los muItados mosiramn una p n httwogentidrid en la mayorla dE los caracteres analizados. A pesar de dla, algunas características macromorfol6gicasrelacionadas con el color y forma de las hojas y bráctcas en cl gdnero Androcymbium, podrían ser utilizadas como indicadores taxondrnicos de afinidades entre especies. Contrariamente, las caracterfs~casmicromarfol6gicas y anatdmieas estudiadas m m m n una gran variadad de formasque no st ajustan a las relaciones entre cspies establecidas previamente a panir de datos & morfologla, aloenzimas o WLPs del cpDNA. llardi Bothnic de Barcelona, cl Font i Quar s m . , 08038 Barcelona, Spain. E-mail: nuriamem@telefonica,ntt a Estacid Internacional de Biologia Mediterrhnia-Jardl Bothnic Marimurtra, Passeig Karl Faust 10, 17300 - Blancs, Girona, Apdo. de correos 112, Spain. J d n Botánico 'anario Viera y Clavijo, Apdo 14 de Tafira Alta, 35017- Las Prilmns de Gran Canaria, Spain. 84 COLLECTANEA BOTANICA (BARCELONA) 26. 2003 INTRODUCTION The species of Androcymbi/lm Willd. (Colchicaceae) are geophytes with no annual vegetative cycle that spend the unfavorable period buried as tunicate corms. The genus ineludes aboul40 speeies (ARNOLD & WETT, 1993; MOLLER-OOBLlES & MOLLER-OOBLlES, 1984, 1998; PEDROLA-MONFORT el al., 1999a, 1999b, in press.) wilh a disjunel diSlribulion in Afriea (lig. 1A). The six species in Nonh Africa are distributed throughout the Mediterranean basin and the Canary Islands. Despite their clase morphologicaJ similarity, ｾｒｄｎａＭｵｅｔａｍ el al. (1996) argued lhal several morphologica1 Jeaf eharaclers are useful for inler-speeilie differentiation. Qualitative (size, shape. color, and number of leaves) and anatomical characters (mesophyll cell types, epidermie ceH size, thickness of cell wall. and central epidermic cel! size related lO (he rest of the lamina) were particularly discriminative. The rest of the species in the genus are distributed in South Africa, mainly in the Westem region. These species display much higher variability levels than their Northem African congeners al lhe morphological (BAKER, 1974; KRAUSE, 1920; MOLLER-OoBLlES & MOLLEROoBLlES, 1998; MEMBRIVES, 2000), pollinie (MARTfN el al., 1993; MEMBRIVES, 2000; MEMBRIVES el al., 2002), karyologiea1 (MARGELI el al., 1998; MONTSERRAT el al., 2002), allozymatie (MEMBRIVES el al., 2001) and epONA ｓｌ ｅｔｓａｃＭｾｐａｊｕ ｃＨ el al., 1999) Jevels. The main objective of this work is to evaluate the variation in leaf and bract morphological and anatomical features of a comprehensive representation of laxa of the genus AI/{Irocymbiu11l distributed in Southwest Africa (fig. 18). 1 ＯｔｾＧＭｮ ,n n F ｾ " Ｌｾ , '" ｾ >-. \. .-'tr Ｌｾ " ｾ ........ 1'..(' -.. ¡..o... fn·EK' 30 ｾ ｾ W\l.I T .• ｾ ) ｬｓｾＧＢ ｾＬＮ Ｎｾ ｾ B " A POEl-ST ｾ 32 ｾｦＩ " """ ., ｾ ｾﾷ［ｴ Ｎｾ＼ａ Vi ｾ F éÍ ."'1""- .le- . 1\..- ,-ｾ V 20 22 Figure l.- A. Geographic.al distribution of genus Amlroc)'mbi/lm. B. Distribution of the Southweslem African populatiol1s studied. Abbreviations of populations Dore described in Table 1. N. MEMBRIVES. 1. PEDROLA-MONFORT & J. ｓｕｅｔｓａｃＭｾａｊｕ ｃ 85 MATERIAL AND METHODS We sampled 32 populations belonging to 17 taxa of the genus Androcymbi/ll1J (Table 1). These samples are currently in cultivation al lhe greenhouses of the "Estació Internacional de Biologia Mediterrania-Jardí Botanic Marimunra". Quantitative and qualitalive morphological data were obtained from the observation of adult plants in cultivation measured after lhe anlhesis. The terminology used for the morphological descriplions of ¡eaves and bracts follows HEYWOOD (1978). Micromorphological and anatomical characters were obtained from samples of the central pan of the lamina of the lirst leaL MacromorphologicaI characters We studied conn morphology, tunic characteristics and several vegetative characters in Icaves and bracts (number, distribution, shape, size, section. and color). Observations were made from material in cultivation and from herbarium sheets of specimens collected in the field. Micromorphological characters Leaves were heated in chloral hydrale lo separate lhe cutic1e and lhen dyed wilh saphranine. The studied characlers were the indument both on the surface and lhe margin of the leaf, the shape and size of epidermic cells, and the lype and amount of stomal:.l. The stomatic indices (STls) are the quotienl (expressed in percentage) belween the number of stomatic cells and lhe number of total cells (stomata plus epidermic cells) in a restricted leaf area. Problems with manipulation of lea ves in A. /¡ellssellimlllm and A. lumlleyi did not al10w us to obtain this kind of information for these species. Anatomical characters Leaves were fixed in Camoy's solution (JOHANSEN, 1970), and cut using a scalpel blade and a binocular magnifying glass. The anatomical characters studied were mesophy1l ce1l types, epidermic ceH sizes, ceHular wall {ypes, size of central epidermic ceJls relative to the other epidermic cells, and amount of idioblasts. REsULTS Corro The corm is rounded in most of the species studied, and dorsiventrally compressed in A. be///lI1l, A. poe/liall/ll1l, and A. wa/leri. The corro is wrapped by a leaf (called cataphyll) that becomes a tunic in the next cyele, with either a papery texture (in A. a/lSlrocllpense and other Southeast African species like A. decipie'ls, A. IOllgipes. and A. 'lQla/ense) or a coriaceous texture (the remaining species studied). The tunie surface is generally smooth. excepl for A. be//ul1l, A. poellianum, and A. walleri, where it is ragged. A basal elongation of the corm and tunics (the cresO is a diagnostie charaeteristic of the genus (lig. 2A). This feature is largely developed in A. cltspidatum (lig. 2B), and extends widely ayer the carm faces in A. bellum, A. poe/lianlll1l, and A. \Valte"; (lig. 2C), Morphology aud loaC dislribulion AH studied species shawed three leaves, rarely four (only in A. allslrocapellse. A. irrorallll1l and A. wa/leri). The ¡eaves are altemate (fig. 3A) and make up clase nades that generally form a basal roserte. A'ldrocymbium bellum, A. circ;'IOIllm, A. dregei and A. wa/leri develop an epigeous stem in cultivation. This variability in stem length is probably due to insuflicient sunlight in the greenhouses, and was al so deseribed previously in A. dregei (MOLLER-DoBLlES & MOLLER-DoBLlES, 1984). In almost all specios, .he leaves are 86 COLLECTANEA BQTANICA (BARCELONA) 26. 2003 Table l.- Populalions of Alldroc)'lIlbiul/1 analysed. Population. Cade and Locality. A. a/bol/ellse ScMnland subsp. clalllvjlliamellSe Pedrola. Membrives & 1. M. Monts.. ALBA· PK. 3219AA. Wuppenal: Clanwilliam to Wuppertal road. km 10. A. QlIstrocapellu U. Müll.·Doblies & D. Milll.-Doblies. AUST-GH. 3418AC. Simonstown: Cape or Good Hope. A. 1Iltstrocapellse U. MUII.-Doblies & D. MUII.-Doblies, AUST-WP. 34ISAD. Simonstown: Whale's Poinl. Cape Paiol Reserve. A. bel/IIIII Schltr. & K. Krause, BELL-VI. 2817DC. Yioolsdrirt: Slcinkopf lO Vioolsdrift rondo km 40. A. burclJellii Baker subsp. burchellii. BURC·HX. 3319BC. Worcester: from Worcester lO Towsrivier road. A. burchellii Baker subsp. pulcllrum Pedrola. Membrives. J. M. Monis. & Caujapé. PULe-CA. 31 19DA. Calvinia: Calvinia lO Ceres road. 7 km tumoff 10 Kreitzberg. A. bl/rchellii Baker subsp. pulcllrum Pedrola, Membrives. J. M. Monts. & Caujapé. PULC-NI. 3118AA. Calvinia: Wild nower reserve of Nieuwoudlville. A. capellse (L.) K,Krause. CAPE-HO. 3318AB. Cape Town: Ma1mesbury to Hopefie1d road, km 49. A. circillat/wI Baker. CIRC-NB. 29170B. Springbok: Springbok 10 Nababeep road. 100 m. A. circinawm Baker, CIRC-SB, 2917DB. Springbok: 3 km W of Springbok. A. cuspidawm Baker, CUSP-CA. 3119DA. Calvinia: Ca1vinia to Ceres road. 7 km lurnoff to Kreitzberg. A. cllspidatllm Baker. CUSP-MO, 332OCD. Monlagu: Near Montagu-Badsk100f. W of Ihe Gorgo. A. dregei c.rres!' DREG-PK, 3219AA Wupperta1: CJanwilliam lO Wuppenal road, km 28. A. egllilllocymbioll U. MUII.-Doblies & D. MUII.-Ooblies. EGHI-CI. 321808. Clanwilliam: Piketberg lO Citrusdal Pass. A. eghimocymbioll U. Mül1.-Doblies & O. MUlI.-Ooblies. EGHI-PK. 3219AA. Wuppertal: C1anwilliam to Wupperta1 road, km 28. A.llOllfamen5e Schinz. HANT-CA. 31 19DA. Ca1vinia: Calvinia lO Ceres road. 7 km lurnoffto Kreitzberg. A. /¡ell.fsen;allllm U. MUIl.-Ooblies & O. MUlI.·Ooblies, HENS-EK. 2817CC. Vioolsdrifl: Eksteenfontein 10 Modderfontein road. A. III11Itleyi Pedrola, Membrives. J. M. Monis. & Caujapé. HUNT-EKI, 2917AD. Springbok¡ Springbok to Port Nollolh road. 14 km 10 Eksleenfonlein. A. lumtleyi Pedrola, Membrives. J. M. Monts. & Caujapé. HUNT-EK3. 2917AO. Springbok: Springbok to Pon Nol1Olh road. 20 km 10 Eksteenfonlein. A. irraratllm Schltr. & K, Krause. IRRO-EK, 2917 AO. Springbok: Springbok lO Port Nolloth road. 6 km lO Eksteenfonlein. A. irrora//Im SchlLr. & K. Krause. IRRO-EK2. 2917AD. Springbok: Springbok lO Pon Nolloth, 15 km 10 Ekslccnfonlein. A. irroralllm $chltr. & K. Krausc. IRRO-EK6. 2817CC. Vioolsdrift: Eksleenfonlein 10 Modderfontein road. A. irroralllm Schltr. & K. Kmuse. IRRO-KA. 3018CB. Kamiesberg: Biuerfontein 10 Kliprand road. A. irroratllm Schltr. & K. Krause.IRRO-KW. 31 18BC. Vanrhynsdorp: Vredentallo Koekenaap road. 100 m lO train stalion. A. irrorarulIl Schltr. & K. Krause. IRRO-VP. 3119AC. Ca1vinia: Vanrhynspass. A. irrOrtltlll1l Schltr. & K. Krause. IRRO-VY, 3118AD.Vanrhynsdorp: VrendendaJ to Vanrhynsdorp road. A. poelriallllm U. Mü11.-Doblies & O. Müll.-Dob1ies. POEL-CO. 29170B. Springbok: $pringbok 10 Concordia road. A. poeltiallUIIl U. Müll.-Dob1ies & D. MUII.-Doblies. POEL-NB. 29170B. Springbok: Springbok to Nababeep road. 100 m. N. MEMBRIVES, J. PEDROLA-MONFORT & J. CAUJAI'É-CASTELLS 87 Table 1. Conl. A. poeltiallul1l U. MUII.-Doblies & D. Müll.-Doblies. POEL-ST. 2917DC. Springbok: Steinkopf to Springbok road. 5 km. A. villosum U. MUII.-Doblies & D. MUlI.-Doblies, VILL-EK. 2817CC. Vioolsdrift: I km S of Eksteenfonlein A. villosum U. Milll.·Doblies & D. Müll.-Doblies. VILL-ST. 29178C. Springbok: 3 km S of Steinkopf. A. walteri Pedrola. Membrives & 1. M. Monts.. WALT-ST. 2917DC. Springbok: Stcinkopf lO Springbok road, 5 km. A B Figure 2.- Shape of the corm cres! in Alldroc)'mbiufII. A. Basal erest. B. Basal crest largcly developed (A. c/lspidawm). C. Basal erest growing by the corm faces (A. bdllllll. A. poeltialllllll and A. wa/teri). I, B " Co o T A e Figure 3.- A. General plant Slrueture in Androc)'lIIbilll1/. B. Distichous distribution. C. Trislichous dislribulion. D. Helicoidal distribution (T= lunic; Ca= calaphyl1; I(x)= Icaves; b=bracl, f=flower). distributed distichausly (fig. 3B), like in masl genera af the Liliales (DAULOREN, 1985). However, (he distribution of ¡eaves in A. cuspidotllm is either tristichous (fig. 3C) or distichous with the first leaf in a transversal POSilioll. Leaf dislribution aroulld the stcm is helicoidal in A. dregei (fig. 3D) and in the Northem African circumscription of Ihe genus (PEDROLA-MoNFORT, 1993: ｓｾｒｄｎａＭｕｅｔａｍ el al., 1996). COLLECTANEA BQTANICA (BARCELONA) 26, 2003 88 The Icaves are generally linear-Ianceolate. However, lhey are acicular in A. dregei and A. hellsSeniall/llll, and ovate-Ianceolate in A. burchellii subsp. bllrcheJlii, A. b//rc!lellii subsp. pulc!lfllm, A. cuspidalllm, and A. irroralllm. The biometric characterislics of the first leaf are described in Table 2a. The ¡argesl leaves are those of A. auslrocapellse (mean value of 23.8 cm) and lhe shortesl (hose of A. cllspidawm (mean value of 4.1 cm). The narrQwest leaves belong to A. dregei and A. Ilellssell;allum (mean values of 0.1 and 0.2 cm, respeclively), and the widesl belong to A. bllrcheJJii subsp. burchellii, A. burchellii subsp. pulchrum, and A. capense (mean values of 2.3, 2.9 and 2.3 cm, respectively). Al! species sludied showed a fiat leaF apex, except for A. circinaflllll, A. hensseniwlI/11/, and A. villosum, which showed a circinate apex. Two general types of leaf section were observed (figs. 4, 5 and 6): fiat with a shallow depression in the region of the central nerve (A. burche/l;; subsp. bllrche/lii, A. bllrchel/ii subsp. pulchrum, A. capellse, A. cltspidatllm, A. hantamellse, A. irroratllm, A. poeltiallll1l1 and A. walterO, and V·shaped (A. albmlellse subsp. clallwilliamense, A. allstrocapellse, A. be/lum, A. dregei and A. eghimocymbioll). AlldrocymbiuJII circillatum, A. hellsscniallul1f, A. IlIlIItleyi and A. vil/OSI/JII showed an intermediate leaf seclion (fig. 5F; fig. 6F, 61, 6L). The leaves were either bright green (A. alballellse subsp. clamvill;amellse, A. (Il/strocapellse, A. burchel/ii subsp. burchel/ii, A. bllrchel/ii subsp. pulcllrum, A. capense, A. clIspidatum, A. egllimoc)'mbioll and A. irroratum) or green-glaucous (A. bel/llm, A. circinatum, A. dregei, A. hantamense, A. hellssell;amml, A. IlImtleyi, A. poeltiallllm, A. villosum and A. walterO. Table 2a.· Macromorphologicalleafcharacters in AIUJrocymbiul1I. Shape: A=Acicular; L=Linear; LA=Lanceolate: OV=Ovate. Measures are expressed in cm. Color: GL=Glauc; GR=Green.• = presence of red spots. Leaves Taxon Shape Length Width UW A. aJbclIlellse subsp. L-LA ( 10.0)13.3(15.5) (0.7)0.9(1.1) 14.8 V GR L-LA L aV-LA (13.5)23.8(35.6) (11.0)13.0(15.0) (12.5) 15.9(21.0) (0.5)1.1(2.1) (0.3)0.4(0.6) (1.6)2.3(3.7) 23.0 35.5 7.4 V V PI GR GL GR aV-LA (7.0) 13.0(23.0) (1.4)2.9(5.6) 5.2 PI GR LA L-LA aV-LA A L-LA LA A L-LA aV-LA L-LA L-LA L-LA (14.0)15.3(17.5) ( 14.0)19.3(24.5) (2.2)4.1(5.5) (6.5)7.9(9.2) (13.7)21.6(27.0) (11.0)15.2(20.5) (8.5) 15.0(21.0) (7.0)8.1(9.2) (5.0) 11.4(23.5) (9.0) 12.4( 15.5) (12.0)13.4(16.5) (11.0) 16.9(21.5) ( 1.7)2.3(3.0) (0.5)0.9(1.3) (1.1)1.7(2.2) (0.1)0.1(0.2) (0.4)0.7(1.0) (1.3)1.9(2.7) (0.2)0.2(0.2) (0.7)1.0(1.2) (0.5) 1.41(3.2) (0.4)0.7(1.3) (0.7)O.9( 1.0) (0.5)1.0(1.4) 6.8 24.3 2.5 71.3 35.4 8.5 84.0 8.8 9.3 19.9 15.8 18.3 PI PI-V PI V V PI PI-V PI-V PI PI PI-V PI GR GL' GR GL GR GL GL GL GR GL GL GL Section Colour cJamvi/Jiamellse A. ClI/Slrocapellse A. belJ/lm A. burcheJlii subsp. burchelJii A. burchellii subsp. pllJchml1l A. cClpellse A. circ;flallll1l A. clIsp;datum A. dregei A. eghil1loc)'lIIbioll A. IWlllamellse A. 1Jellssell;wIIIIII A. IlImtleyi A. irroralllm A. poeltiamtm A. Vi/lOSll1ll A. W(lller; . N. MEMBRIVES, J. PEDROLA-MONFORT & J. CAUJAPÉ-CASTELLS 89 , mm I mm • e D B I mm 0.1 mm E .. G f I mm 0.1 mm K J H Figure 4.- Mesophyll with differentiated cells. Section structure: A. A. burc/¡elfii subsp. burchellii. C. A. irrorl/lIllll. E. A. /ulIItamellse. G. A. el/pellse. H. A. WI.\·trocapell.\·e. J. A. dregei. Seclion: B. A. bllrcllellii subsp. burc/¡elfii. D. A. ¡rrora/lIl11. F. A. {UlI/llllllellse. l. A. mmrocapellse. K. A. dregei. Black circles in structure correspond to idioblasts. White circles in sections correspond to vascular vessels. COLLECTANEA BOTANICA (BARCELONA) 26. 2003 90 1 mm 0.1 mm 1 mm 0.1 mm . hunll • A ,, 1 mm ,·· 0.1 mm .. • ·· 1 mm .. · : C D · ·'. ·'. G '. H Figura 5.- Mesophyll with undifferentiated cells. Seclion Slructure: A. A. cl/.Jpidawm. C. A. E. A. IlIlI//ley;, G: A. \\Ia/teri. Section: B. A. clIspidallll1l. D. A. poel¡iclJllllll. F. A. !JOeltÜlIlfll1l. II/Ulrleyi. H. A. wo/teri. Black circ1es in struclure correspond to idioblasls. While circles in seclions correspond to vascular vessels. Alldrocymbill1l1 circ;,wIlIm (ealures red spots (approximately 1 mm in diameter) in leaves, bracts and lepal surface that appear jusI befare flowering. This characleristic is shared by (he new speciesA. complOllii U. MUIJ.-Doblies & D. MUlJ.-Doblies (MOLLER-DoBLlES & MOLLERDOIJL1ES. 1998). 80th leaf faces in Alldrocymbilllll species are greel1. except in A. burcl1ellii subsp. burchellii and in populations IRRü·KA and IRRO-VY of A. irroratwII (abbrevialions of populalions are in Table 1), where they are purplish on lhe abaxial face. The bracts usually differ from leaves in shape. color and leXlure. In most species. bracls were shorter and more rounded lhan leaves, and in sorne cases the color was different. Generally. the third leaf showed an intermediate shape between leaf and bract. Although most species of Androcymbillm surveyed showed elliptic or deltoid bracts, exceptions like acicular (A. dregel1, lanceolate (A. hellsse"ialllll1l and A. poeltiamllll), and orbicular (A. bllrchellii subsp. b/lrchellii and A. bllrchellii subsp. plllchrum) do occur (Table 2b). The bract measurements are given in Table 2b. The largest bracls occur in A. bllrchelW subsp. plllc!lrt/II/ and A. hmuame,¡se (mean values of 7.4 and 7.7 cm, respectively). and the shortesl in A. albaneflSe N. MEMBRIVES, 91 J. PEDROLA-MoNFORT & J. CAUJAPé-CASTELLS 0.1 0.1 mm mm I mm • • • e A '. D 0.1 mm "" . .. " 0.1 mm E 1 mm F ." 0.1 mm K l "" 1 mm Figure 6.- Mesophyll with undirrerenliated cells. Seclion structure: A. A. lllbllllt!IIJe subsp. c111lllvillillmellse. C. A. bel/lIl11. E. A. circiJlllllllll. G. A. eghillloc:ymbioll. 1. A. IU!I/.uellillllllm. K. A. vil/OSI/m. Section: B. A. alballel1Je subsp. c/clIlwi/liclI1leIlJe. D. A. belllllll. F. A. circiJllltlllll. H. A. eg"i/l/ocymbioll. J. A. 11e11.uelliclIllllll. L. A. vil/OSUJIl. Black circles in structure correspond lO idioblasls. While circles in seclions correspond lO v:lsculur vessels. COLLECrANEA BQTANICA (BARCELONA) 26, 2003 92 Table 2b.- Macromorphological braCI characlcrs in Alldrocymbilll1l. Shape: A=Acicular: DE=Dcltoid; E=Elliptic; LA=Lanceolatc; OR=Orbicular; OV=Ovate. Measures are expressed in CIll. Color: WH=White: GL=Glauc: RE=Reddish; GR=Green.• = presence of red SpOlS. Bracts Taxon A. alballellse subsp. clllllWilI iamellJe A. tlltSlrocapellse A. bellulIl A. bllrcltellii subsp. bllrche/lii A. bllrc"ellii subsp. A. A. A. A. fJlllchmm cllpeme circillatlll1l cflspidatfllll dregei A. eghimoc)'mbioll A. holltamellse A. lletlSSellialllllll A. JI/ml/e)'i A. invrall1m A. poelriall//11I A. vilfosllm A. Il'alreri Shape Lenglh Width UW Colour DE (1.7)2.7(4.0) (1.0) 1. 7(2.0) 1.71 GR DE E OR (2.5)3.8(5.5) (2.1 )3.3(6.2) (3.0)3.9(5.0) (2.5)3.6(4.0) (1.5)2.0(2.9) (3.0)4.3(5.2) 1.06 1.67 0.93 GR GL WH OV-OR (4.0)7.4(12.5) (4.5)6.3(9.0) 1.20 RE E E-DE DE A DE E-LA LA E-DE E-DE LA E-DE E-DE (4.0)4.7(5.5) (3.4)4.5(6.2) (2.4)4.1(6.5) (2.5)4.3(7.2) (2.5)4.1(8.0) (5.5)7.7(11) (3.4)5.7(7.5) (3.0)3.1(3.2) (2.0)3.8(5.0) (2.4)4.6(6.0) (3.5)5.5(7.5) (2.7)4.3(5.8) ( 1.7)2.3(3.4) (2.3)3.0(4.4) (1.1)2.4(4.0) (0.15)0.2(0.3) (1.7)2.8(4.0) (2.1 )3.8(6.0) (1.3) 1.7(2.3) (2.0)2.1(2.2) (0.8)2.2(3.5) (1.1) 1.8(2.8) (1.8)2.7(4.0) (2.1)3.1(4.2) 2.12 1.51 1.79 21.63 1.48 2.16 3.41 1.48 1.77 2.61 2.05 1.38 GR-WH GL' GR GR GR WH GL GL GR GL GL GL subsp. c1allWillial1lense (mean value of 2.7 cm). The narrowes( bracls belong to A. dregei (mean value of 0.2 cm) and the widest lo A. burchelfii subsp. pulchrulll (mean value of 6.3 cm). On lhe whole, lhe bracls were green or green-glaucous, bUl they can also be while (A. cllpellse, A. 11ll1l111me1lSe and A. burcJlellii subsp. burcJlellii) or reddish (A. burche/lii subsp. plllcJlmm). Micromorphological charaeters of surrace and lear margin By and large, lhe indumenl af lhe leaf margin was eilher smaalh ar papillase (fig. 7A). Unicellular hairs (which can be simple or bifurcate) were observed in A. vi/lOS1I1ll (rig. 78). Pluricellular hairs were shorl and sparse in A. bllrcllel!ii subsp. bllrchel/ii and A. bllrchellii subsp. pulcllrum (fig. 7C), and long and very frequent in A. capellse and A. ha1l1amellSe (fig. 7D). The leaf surface is glabrous in aH of the studied species excepl in A. vil/osum, which shows unicel1ular hairs (simple or bifurcated) on the abaxial face of lea ves and bracls. Tlle epidermis of the abaxial face of the lea ves showed rectangular (rig. 8A), rhomboidal (fig. 8B) ar palyganal eells (fig. Se). AlldrocYlllbilll1l ｩｬ･ｨｾ｢ subsp. bllrchellii and A. bllrchel/ii subsp. plllchmm had polygonal cells on both faces. This kind of cell was only observed on Ihe abaxial face in A. hmllamellse and A. irroratum (Table 3). The anliclinal walls of the epidermic cells were slraighl, except for those of A. clIspidatum, which were curved (fig. 80). The longest cells in the adaxial face belonged lo A. eg!limocymbioll, and N. J. MEMBRIVES, PEDROLA-MoNFORT & J. 93 CAUJAPÉ-CASn:LLS A e o 00 D B Figure 7.- Indument in leaf margino A. Papillose. B. Unicellular. simple and bifurcate hairs (A. vil/osI/m). C. Pluricellular short hairs. D. Pluricellular long hairs. 1( I I'1I '('!¿él I )' )m, \ J , /J! / " 111¡flJl/11 ", 1 1 JIIr '/ t !I(' (Ir} . /) / al; \ I 1IiI! . /!'(I ¡·/;/Jf11 ¡ I ! ",1(1 J I ¿¡JI (" [ / " JI!J (,r t" ,) i '-1 /1 ,'-}1.) 6 Figure 8.- Epidermic leaf cell shape. A. Rectangular cells. B. Rhomboidal cells. C. Polygonal cells. D. Rectangular cells with sinuose anticlinal walls. Black lines correspond lO 100 J.1111. COLLECTANEA BQTANICA (BARCELONA) 26. 2(0) 94 Table 3.- MicromorphologicaJ and anatomical leaf characterislics. Epidermic cells: meun±slandard deviation (...m). LAd and WAd = Epidermic cell length and widlh in adaxial race: SAd=Epidermic cell shape in adaxial race; Lab and Wab=Epidermic cells length and widlh in abaxial race: SAh= Epidermic cell shape in ahaxial race. RE=Reclangulor. RO=Romboidal. PO=Polygonal. leAd=Stomatic iodex in adaxial face (in %). leAh=Stomatic iodex in abaxial (ace (in %). Central epidermic cells of Ihe lamina: D=differenl size; S=similar size. M: Mesophyll (ND=undifferenliated; D=differentiated). Number of layers of palisade parenchyma in parenthesis. ID= Idyoblasts (F=Frequent. FF=Few frequenl). Numbers in parenthesis in 'Leaf margin' correspond lO Ihe cells forming the hairs.·: missing dala. Epidermic cells Taxon A. albollellse subsp. clall williamellSe A. Cl//Slrocapellst! A. bellllm A. b/lrchelli; subsp. bl/rchelli; A. bllrchellii subsp. pulchrllm A. capellse A. circillatllltl A. cllspidallllll A. clregei A. eghilllocymbiol/ A. IUIIlUlmellse A. hellSSelliamltll A. IlimIte)'; A. irroratum A. pOell;allllm A. I/illosl/III A. walleri Leaf mare:in LAd WAd SAd LAb Papillose 242.0.66.8 41.5.4.2 RE 310.5.42.4 Papillose Smoolh Hairy (4-6) 221.0.58.4 203.5.35.5 101.0.9.1 62.0.9.9 34.5.5.4 63.5.11.4 RO RE PO 289.0.69.1 165.5.45.8 214.0.20.2 Hairy (5-9) 70.5.12.4 78.0.6.7 PO 224.5.25.5 Hairy (8-10) Smoolh Smoolh Papillose Papi1lose Hairy (3-5) Smoolh Smooth Papillose Papillose Hairy (1) Papillose 174.0.35.8 232.5.60.0 183.5.24.9 214.5.70.1 285.5.42.7 178.5.30.9 34.0.2.9 27.5.3.5 41.5.5.2 27.0.2.1 53.5.4.2 58.5.4.5 RE RE RE RE RE RO RE 254.0.31.2 240.5.45.4 234.0.47.7 257.5.24.2 306.5.63.2 195.0.30.0 RO RE RE RE 217.0.42.5 228.5.85.4 198.5.41.7 157.0.19.4 - - 218.5.50.4 242.5.45.0 185.5.33.7 155.5.45.1 - 37.0.54 49.0.4.2 34.5.2.1 47.5.7.7 - - lhe shortesllo A. bllrcheJJii subsp. p"lchrum (Table 3). The widesl cells in lhe adaxial face belonged lo A. burchellii subsp. pl/lchrum. and the narrowesl cells to A. dregei. The longest cells in the abaxial face belonged lO A. alballellse subsp. clamvi/liamellse, and lhe shorlest lO A. walteri. The widesl cells in Ihe abaxial face belonged to A. halllamellse, and lhe narrowesl lO A. dregei. Androcymbium leaves are amphyslomatic wilh anomocylic stomata (D1LCHER, 1974), as in mosl families of the Liliales, except for Ihe Orchidaceae and Cypripediaceae (DAHLGREN el aL, 1985). The stomatic indices (STIs) ranked belween 20 in A. vi/losl/m and 39.6 in A. eghilllocymbioll in the adaxial face, and between 18.2 in A. bellulII and 40 in A. eghimocymbioll in lhe abaxial race (Table 3). Leaf anatomy The epidermis was formed by only one quadrangular or rounded cell layer. In a transversal CU1, the central epidermic cells on Ihe adaxial face were similar in size lO the res! of cells in A. allslrOCapellse, A. cuspidalllm, A. dregei and A. eghimocymbioll (fig. 4H, 4J; N. MEMBRIVES, J. PEDROLA-MoNFORT & J. ｓｌ ｅｔｓａｃＭｾｐａｊｕ ｃ 95 I Central cells WAb SAb leAd leAb 43.0±3.7 RE 35.7 38.6 45.5±6.9 39.0±4.5 52.5±3.5 RO RE RO 34. I 25.4 25.7 53.5±3.8 RO-RE 44.0.3.4 46.5±4.2 36.5±1.4 M ID D ND F 34.0 18.2 24.5 S S D D(2-3) ND D(4) F FF F 32.7 30.0 D D(2-4) F - RE-RO RE RE RE RO RO-PO RE 32.9 29.7 27.6 25.9 39.6 36.7 32.7 18.6 20.7 28.9 40.0 27.7 51.5±5.8 34.5±5.4 37.0±2. I 51.5±2.9 RO-PO RE RE RE-RO 36. I 25.5 20.0 31.1 31.6 18.8 20.0 28.4 D D S S S D D D D D D D D(I) ND ND D(I) ND D(5-7) ND ND D(2-3) ND ND ND F F FF FF F F FF F F FF F FF 25.5::t2.1 55.5±5.7 62.5±0.0 - - - - fig. 5A; fig. 6G). In the other species studied, the central cells were three lo four times larger Ihan lhe resl (fig. 4A, 4e, 4E, 40; fig. 5e, 5E. 50; fig. 6A, 6e. 6E. 61, 6K). The mesophyll was made up of 6-12 cell layers and corresponded lo 60-88% of Ihe lamina thickness. We were ablc lO recognize lwo mesophyll types according lo lheir having differenliated or undífferentiated cells (Table 3). The firsl type showed isodiamelric cells forming the spongy parenchyma, and prísmalic cells forming lhe palisade parenchyma (fig. 4). The palisade parenchyma in bolh faces of lhe lamina was observed in A. hanlllmense (fig. 4E). The palisade parenchyma on the adaxial face was shown in A. allstrocapellse. A. bllrchellii subsp. burchellii, A. burchellii subsp. pu/chrum, A. capellse. A. dregei and A. irl'orallll/l (fig. 4A, 4e, 40. 4H, 4J). The olher species showed a mesophyll wilh only isodiamelric cells (fig. 5 and 6). The density of leaf idioblasts (L c., specialised cel1s Ihat produce sulphurale mucopolysaccharides lhat are released belween Ihe parenchyma cells and are similar to Ihese in shape and size) was variable al lhe intm and inler·specific levels. In general, leaf idioblasts were less frequent in A. bellulII, A. c/lspidarll11/. A. dregei, A. hellSsenialllllll, A. poe/liallU11l and A. \Va/reri (Table 3). 96 COLLECTANEA BOT....NICA (BARCELONA) 26. 2003 DISCUSSION The presence of a papery eonn lunic could be considered a primitive lrait in AlldrocymbiulIl on the grounds thal it is shared with oLher genera of (he Colchicaceae like Colchicu11I or Omithogloss",,, (NORDENSTAM, 1982). These papery tunics have beco described in {he SouthweSlem African species A. austrocape"se, thal occurs along the Coasl [rom Cape Town lo Port Elizabeth, and a150 in species in Southeastem Afoca, like A. decipiens, A. IOllgipes and A. llataleflse. Following these observations, a papery texture could be associated wilh Ihe different amounl of rainfal1 in the areas where AlIdrocymbium is dislributed. AlldrocYlllbi/llll auslrocapense and the Southeastem African species inhabit a zone with abundant winter rainfall, whereas lhe rest of Soulhwestem African species studied (those which exhibit coriaceous lunics), occur in aclimate characlerized by high aridity. The distichous leaf distribulion wilh the first leaf in transversal posilion in the newly described species A. \VorSOllellse U. Milll.-Doblies & D. Milll.-Doblies (MOllERDOBLlES & MOLLER-DoBLlES, 1998) was the diagnostic characteristic used to differentiate it from A. clIspidlltltm, which is described with a tristichous leaf distribulion. However, we have observed bOlh leaf dislributions in the same populatiol1 of A. cltspidalllm in cultivation. In the face of this intra-populalion variabilily, lhe descriplion of A. worSOllense could be superfluous if olher discriminalive characterislics between lhese lWO species do nol exisl. A circinate apex was observed in A. circillatllm, A. villoswll and A. hensseniamun. The former t\\lO are c10sely related at the morphological (MOLLER-DoBLlES & MOLLERDOOllES, 1998; MEMORIVES, 2000), allozymalic (MEMORIVES el al., 2001), and cpDNA RFLPs levels ｓｌｅｔａｃＭｾｐｊｕＨ el aL, 1999). Conversely, A. hellssenit/lll/m does 110l bear a close relation to any of these two species either morphologically or allozyrnatically and it is included in a different seclion in the new propasal of classificalion of the genus (MEMBRIVES, 2000). Apex morphology could lherefore be considered a convergent characlerislic in lhese species. As opposed lO lhe morphological homogeneity reported in their Northem African congeners (GREUTER 1967; PEDROLA-MONFORT, 1993), Soulhwesl African speeies of Alldrocymbium exhibit differences in leaves and bracls lhat have been cOllsidered laxonomieally informative (SAKER, 1874; MOllER-DoollES & MOllER-DoBlIES, 1998). Specifically, our survey distinguishes three groups of species according to the shape and color of leaves and bracIs. One of lhem is formed by species wilh undifferenliated leaves and bracts (A. dregei and the Northem African species). A second group is lhal of species wilh bracls smaller and more rounded than leaves (A. albanense subsp. c/anwilliamellse, A. auslrocapense, A. bellllm, A. circillalllm, A. cllspidalllm, A. eghimocymbioll, A. hellsselliallum, A. IUllltleyi, A. irroratum, A. poeltialfum, A. villosuf1I and A. walteri). Finally. a third group is formed by species with bracls and lea ves different in shape and color (A. burcheJJii subsp. burcheJJii, A. burclleJJii subsp. pulcllrum, A. capense and A. IIa1ltamense). While admitting a few exceptions, Ihese groups show a close correspandence Wilh lhe phylogeoetic classificalion based on morphological dala proposed by MEMORIVES (2000). Anolher group according lO lhe gradual transformation from leaves to bracts was eSlablished by MOLLERDOBLlES & MOLLER-DoBLlES (1998) lo diagnose lhe new Subseclion Gradatocymbiu/1I io Iheir recent revision of Section Alltirocymbiul/l. This lrail was oot iocluded in the rnorphological c1adislic analysis, where only leaves and bracls, differentiated and undifferenliated in shape, were considered. The hairy indumenl 00 the leaf margins seems lo follow two different trends in the four species where it was observed. The firsl is characterized by long and frequent hairs and N. MEMBRIVES. J. PEDROLA-MONFORT & J. ｓｌｅｔＬＢＮｃＭｾｐｊｕａ 97 was observed in A. halllamellse and A. capellse. While these two species are c10sely related macromorphologically, they exhibit considerable differentiation at the al10zyrnalic level, with a genetic identity of 0.242 (MEMIlRIVES et al., 200 J). The second general trend features short and scanered hairs and was observed in A. bllrchelfii subsp. bllrcheJlii, and A. bllrcheJlii subsp. pulchrum. In sharp contrast with the heterogeneity obscrvcd in Ihe two species exhibiting the first kind of indumenl, these Iwo subspecies are very c10sely related al all studied levels: rnorphological. allozymatic (MEMIlRIVES et al.. 200 1). and cpDNA RFLPs ＨｃａｕｊｐｾＭｓｔｅｌ et al., 1999). WILSON (1998) described reclangular, rhomboidal and polygonal epidermic ceH shapes in species of the series Californicae of the genus Iris and suggested that a polygonal shape could be a consequence of the division of rhomboidal cells. If this was an explanation in AlldrocymbiwlI, lhen lhe stomatic indices (STI) in species with polygonal ce lis should be higher than in species wilh rectangular or rhomboidal cells. because the relationship between Ihe number of total celis as related to Ihe stomatic cells should be higher. Our results are at odds with Wilson's hypothesis, since we delected no significant differences (t;;;; 0.372; p;;;; 0.716) in the values of adaxial face STIs belween species wilh polygonal cells and species with rectangular or rhomboidal cells. The low morphological variability observed in macromorphological characterislics in the Northern African species of Alldrocymbi//1II conlrasts with the remarkable quali. tative and quantitative hcterogeneily in anatomical lraits (MATEU-ANDRÉS el aL. 1996). Firsl, mesophyll cells can be subdivided into two groups depending on whether the cclls are differentialed or undifferentiated. Secondly, the size of Ihe central epidermic cells is different in relalion lO Ihe rest of Ihe cells in the lamina. Analogous variability patlerns have been shown for Southwestern African species (Table 3). A,¡drocymbium cape1lse and A. dregei showed only one layer of prismalic cells of palisade parenchyma. These two species are nO( c10sely related at any ofthe different levels at which the genus has been sludied (MEMIlRIVES et aL, 2002; CAUJAPÉ·CASTELLS et al.. 1999). The other species with differenliated mesophyll cells showed between IWO and four cell layers, except for A. IWlIIamellse, which revealed Ihe entire mesophyll lo be composed of prismatic eells (from five to seven layers). Among these species, A. burchellii subsp. burchellii. and A. bllrchellii subsp. IJII/chrlllll, shared with A. irroralulII Ihe ovale leuf shape and thick texture, but are not c10sely relmed in Ihe analysis lo allozymic and cpDNA RFLPs. AmlrocYlllbil/m allslrocapense, with Iwo or three prismatic celllayers, is nol closely relaled to any of the olher species characterized by palisade parenchyma (Table 3). The idioblasts described in leaves, bracts and tepals in Amlrocymbium have been related lo the question of defence against herbivores (MATEU-ANDRÉi et al .. 1996). Similar secretive glandulae have not been reported in other genera of the Colchicaceae. Wilhin the monocots, only lhe Liliaceae produce organic sulphurs (BERNUARD. 1970) and Ihis Irend has been considered taxonomically significant (SAGHIR et aL, 1966). Although idioblasls can be considered an exclusive characteristic of Androcymbi/tm, the high intra and inler-spccific variability in their amount and distribulion hinders the use of this qualily as a spccies diagnostico On Ihe whole, Southern African species of the genus An(/rocymbiwlI show a remurkable heterogeneily in maeromorphological, micromorphological and analOmicul characlcristies analysed. Macromorphological characteristics related lo colour and shape of leaves and bracts could be used as taxonomic indicators of relationships among species in the genus. 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