Anais da Academia Brasileira de Ciências ISSN: 0001-3765 [email protected] Academia Brasileira de Ciências Brasil
DALTIN, ÂNGELA L.; ORIANI, ALINE; SCATENA, VERA L. Leaf and inflorescence axis anatomy of Brazilian species of Rapateoideae (Rapateaceae, Poales) Anais da Academia Brasileira de Ciências, vol. 87, núm. 1, marzo, 2015, pp. 157-171 Academia Brasileira de Ciências Rio de Janeiro, Brasil
Available in: http://www.redalyc.org/articulo.oa?id=32738838015
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 Anais da Academia Brasileira de Ciências (2015) 87(1): 157-171 (Annals of the Brazilian Academy of Sciences) Printed version ISSN 0001-3765 / Online version ISSN 1678-2690 htt://d.doi.or/10.1590/0001-376520152010071 www.scielo.br/aabc
Leaf and in orescence axis anatomy of Brazilian species of Rapateoideae (Rapateaceae, Poales)
ÂNGELA L. DALTIN, ALINE ORIANI and VERA L. SCATENA
eartamento de Botnica Instit to de Biociências niversidade stad al Pa lista Avenida 2 A 1515 Caia Postal 199 13506-900 io Claro SP Brasil
anscri receied n erar acceed r icain n
ABSTRACT The anatomy of leaves and in orescence axes of aans (2 s.) aaea (2 s.) Ceasen (1 s.) and cea (1 s.) ( aateoideae aateaceae) was st died to identify sef l characters for taxonomy. The cross-section shape of in orescence axis differentiates the genera, while the cross-section shape and structure of leaf midrib has a speci c value. The following characteristics are exclusive of aans : silica cells randomly distrib ted in the leaf eidermis licate chlorenchyma in the leaf blade presence of ber bundles in the mesophyll and in the in orescence axis parenchyma. aans is also distinguished by the number, type and distribution of vascular bundles in the in orescence axis. The genus aaea is characterized by the resence of stomata and silica cells only on the abaial eidermis of the leaves and chlorenchyma comosed of arm cells in the leaf blade. Characteristics with dianostic val e for Ceasen riedeians are: leaf eidermal cells with straiht to slihtly sin o s walls in frontal view, in orescence axes presenting a de ned cortex, ber bundles facing the larger vascular bundles and a stulous pith. The anatomical characteristics of the leaves and in orescence axes thus proved to be of taxonomic value in generic and speci c levels. They are also useful to differentiate Rapateoideae from other s bfamilies of aateaceae. Key words: anatomy Ceasen cea Poales aaea aateaceae aans .
INTRODUCTION Africa (a ire 1958 Stevenson et al. 1998). In aateaceae comrise aroimately 100 secies Brazil aroimately 39 secies and nine enera in 16-17 enera (Stevenson et al. 1998 ivnish are fo nd rowin rimarily in the Amazon reion et al. 200) and are a basal family in Poales alon (onteiro 201). hey are erennial herbs with a with Bromeliaceae and yhaceae (inder and short stem from which disticho s or sirodisticho s dall 2005 ivnish et al. 2010). he family has leaves and in orescence axes (scapes) originate a restricted distrib tion in So th America mostly (a ire 1958 Stevenson et al. 1998). in the ayana Shield with the ecetion of aateaceae are divided into three s bfamilies: ascaceas dinaei that occ rs in western aateoideae onotremoideae and Sao - fridericioideae (ivnish et al. 200). aateoideae Corresondence to: Aline Oriani -mail: alineorianiyahoo.com.br are the earliest diverent ones and comrise the
An Acad Bras Cienc (2015) 87 (1) 8 NA . AIN AIN OIANI and A . SCANA
enera aans aaea Ceasen in a ateaceae are anatomically characterized and cea (ivnish et al. 200 ). aans is by: aracytic stomata only on the abaial s rface the sister ro of aaea and Ceasen resence of silica cells in the e idermis meso hyll (ivnish et al. 200 ). hese three enera were the differentiated into alisade and s on y tiss e the rst to diverge and are remarkable for presenting last com osed of arm arenchyma and slime- dis ersion ran es that etend o t of the ayana rod cin trichomes in the leaf ails. Shield to the Amazon Basin (ivnish et al. 2000). The in orescence axes in Rapateaceae are he en s cea (made of fo r s ecies) characterized by: hypodermal bers as a continuous was described by a ire (1958) and later sheath or as strands a ro nd arenchyma synonymized as Ceasen by Steyermark com osed of s haeroidal or arm cells resence of (1988) d e to their mor holo ical similarities. henolic idioblasts and vasc lar b ndles nearer to owever Berry (200 ) in his s rvey on a ateaceae the centre enclosed by a common brous sheath of the enez elan ayana lora once a ain (Carl ist 1969). se arates the enera based on the mor holo y of iven the im ortance of identifyin anatomical seeds, leaves and in orescence axes. In the most characteristics with taonomic val e this a er aimed recent hylo enetic st dy of the family (ivnish et to describe the anatomy of leaves and in orescence al. 200 ) which resents s ra eneric taonomic aes of aans icr niaerais aaea chan es cea is also treated as a se arate en s. adsa ei Ceasen riedeians and cea sarrsa that was st died here is one of cea sarrsa (a ateoideae) to characterize three s ecies of the en s that occ rs in Brazil and the s bfamily and the enera contrib tin to the it is endemic (onteiro 201 ). nderstandin of infrafamiliar relationshi s. Ceasen is made up of eight species, ve MATERIALS AND METHODS of which are fo nd in Brazil and Ceasen riedeians that was st died here is also endemic The anatomy of leaves and in orescence axes was (a ire 1958 onteiro 201 ). aans analyzed in si Brazilian s ecies of a ateoideae. com rises two s ecies both occ rrin in Brazil he s ecies st died collectin areas and vo cher (a ire 1958 onteiro 201 ) and st died in this n mbers are resented in able I. he vo chers were work. aaea com rises abo t 20 s ecies (Stevenson de osited in the erbari m ioclarense (CB) et al. 1998) amon which 1 occ r in Brazil (onteiro at niversidade stad al Pa lista NSP. he 201 ) and two of them were also st died. collected material was xed in 50% FAA (37% In the literat re st dies on a ateaceae are formaldehyde, glacial acetic acid, 50% ethanol, scarce and incl de taonomy (a ire 1958 1:1:18 v/v) and subsequently stored in 70% ethanol 1965 Stevenson et al. 1998 Berry 200 ) ollen (ohansen 19 0) for the anatomical st dies. Only mor holo y (Carl ist 1961) anatomy of roots for cea sarrsa the anatomical st dy was leaves, stems and in orescence axes (Carlquist erformed on herborized material which was 1966 1969 errari et al. 201 ) embryolo y revio sly boiled in water and lycerin for tiss e (Venturelli and Bouman 1988), oral anatomy rehydration. e analyzed at least three individ als (Oriani and Scatena 2012) and hylo eny (ivnish of each s ecies collected from different o lations. et al. 2000 200 ). Cross lon it dinal and aradermal sections Accordin to Carl ist (1969) who analyzed in the median re ion of the leaves (blade and the anatomy of vegetative organs and in orescence sheath) and in orescence axes (scapes) were ais of 15 of the 17 enera of the family the leaves made by hand sin dis osable razor blades. he
An Acad Bras Cienc (2015) 87 (1) ANAO O APAOIA
TABLE I Species studied, collecting areas and voucher numbers.
Species Localization Voucher number Par e Nacional da Serra do Ci Ceasen riedeians rn. Scatena et al. 298 Santana do iacho inas erais Par e Nacional do ir cea sarrsa (Willd. ex Link) Maguire ombardi et al. 7579 Caracara oraima aaea adsa Abl. ala o ocantins Scatena et al. 2 Reserva Florestal Adolpho Ducke, aaea ei Pil . ombardi et al. 7611 anas Amazonas Par e Nacional do ir aans icr Ducke ombardi et al. 7580 Caracara oraima Reserva Florestal Adolpho Ducke, aans niaerais esv. ombardi et al. 7618 anas Amazonas sections were doble stained with basic fchsin All s ecies have e idermal cells containin silica and astra ble (oeser 1972) and monted on bodies that may be resent on one or both leaf srfaces. semi- ermanent slides with lycerin elatin. In frontal view these silica cells have a random he resence of henolic com onds was tested distribtion (i s. 1 and 2 - arrows) or are distribted with a eos eCl 3 ( ohansen 19 0). Cross and in lon itdinal rows (i s. 3 and - arrows). lon itdinal sections were also obtained by sin a In cross-section the e idermis is sin le-layered microtome (eica 22 5). or this r ose leaf with ronded or tablar cells (i s. 5-9) of similar and in orescence axis samples were dehydrated in size on both leaf sides (i s. 17-28). he e idermal an n-btyl alcohol series and embedded accordin cells have slightly thickened outer periclinal walls to eder and OBrien (1968) in (2-hydroyethyl)- (Figs. 6-9). Subepidermal ber strands (Fs) are methacrylate (eica istoresin mbeddin it). resent on both leaf sides (i s. 6 7 9 11 13-15). hese sections were stained with eriodic acid S ecies of aans also have ber bundles Schiffs rea ent (PAS) and tolidine ble (OBrien (b) scattered in the meso hyll (i s. 5 11 12 et al. 196 eder and OBrien 1968) and monted 29 and 30). In this ens the meso hyll resents on permanent slides using Entellan (Merck). a licate chlorenchyma (i s. 5 9 18 and 20) he reslts were docmented with a camera while in aaea the chlorenchyma is com osed (eica C 50) co led to a microsco e (eica of arm cells (i s. 7 22 and 2 ). Ceasen 000B) sin the AS software (eica riedeians (i . 26) and cea sarrsa (i s. A lication Site . .0.0). 8 and 28) are distin ished by havin a re lar chlorenchyma. At the midrib re ion of the blade RESULTS all s ecies have chlorenchyma with arm cells (i s. A 29-3 ). ith the ece tion of aaea adsa and he variable characteristics of the stdied s ecies are sarrsa all s ecies have idioblasts containin resented in able II. In frontal view the e idermal henolic com onds dis ersed in the meso hyll cells have sinos (i s. 1-3) or strai ht to sli htly (i s. 11 and 23 - arrows). sinos walls (i . ). he stomata are tetracytic he vasclar bndles are collateral and (i s. 3- ) and may occr on both sides or only on srronded by a doble sheath (i s. 18 20 23- the abaial leaf srface. hen resent on both sides 3 ). he oter sheath consists of thin-walled cells the stomata are more abndant on the abaial srface. while the inner sheath consists of thin or thick-
An Acad Bras Cienc (2015) 87 (1) NA . AIN AIN OIANI and A . SCANA
TABLE II Anatomical characteristics of the leaves of Rapateoideae (Rapateaceae) with taxonomic value. 1: Spathanthus unilateralis , 2: S. bicolor , 3: Rapatea paludosa , 4: R. ulei , 5: Cephalostemon riedelianus , 6: Duckea squarrosa (+ = present, - = absent).
Characteristics 3 4 idermal cells with sinos walls - idermal cells with straiht to slihtly sinos walls - - - - - Stomata on both srfaces - - Stomata only on the abaial srface - - - - idermal cells with silica bodies on both srfaces - - idermal cells with silica bodies only on the abaial srface - - - - Silica cells randomly distribted in the eidermis - - - - Silica cells distribted in lonitdinal rows in the eidermis - - iber bndles scattered in the mesohyll - - - - Plicate chlorenchyma - - - - Chlorenchyma with arm cells - - - - elar chlorenchyma - - - - asclar bndles sheath etensions facin both leaf srfaces - - - - - idrib lane-conve with nmeros vasclar bndles randomly arraned + - - - - - idrib crvilinear with nmeros vasclar bndles randomly arraned - - - - - idrib crvilinear with a central vasclar bndle and two smaller ones - - - - - idrib biconve with nmeros vasclar bndles randomly arraned - - - - - idrib concave-conve with a central vasclar bndle - - - - - idrib biconve with a central vasclar bndle and several smaller bndles arond - - - - - walled cells (i s. 18 20 23-27 31-33). Only (i. 3 ). Ceasen riedeians has a aaea adsa resents sheath etensions concave-conve midrib with only one central in the lar er vasclar bndles facin both leaf vasclar bndle (i. 31) while the midrib srfaces (i . 22). in cea sarrsa is biconve with one he cross-section sha e and strctre of central vasclar bndle srronded by several the midrib varies amon the s ecies. It is lane- smaller ones (i. 32). he vasclar bndles of conve in aans niaerais (i . 29) the midrib reion resent a consicos fiber and crvilinear in icr (i . 30) both with sheath on the hloem in all stdied secies nmeros vasclar bndles randomly arran ed. (is. 29-3 ). In aaea adsa the midrib is crvilinear he leaf marin is acte in most of the with one lar e vasclar bndle and two smaller secies (is. 19 21 23 and 25) with the ones (i . 33) and in . ei it is biconve with ecetion of aans icr (i. 17) and nmeros vasclar bndles randomly arran ed cea sarrsa (i. 27) which show obtse
An Acad Bras Cienc (2015) 87 (1) ANAO O APAOIA
Figures 1-10 - Anatomical characteristics of the leaves of aateoideae secies. (1- ) rontal view of the leaf blade eidermis. (1) aans icr adaial srface. (2) niaerais adaial srface. (3) aaea adsa abaial srface. ( ) Ceasen riedeians abaial srface. (5-7) Cross sections of the leaf blade. (5-6) niaerais . (7) adsa . (8) cea sarrsa . (9) icr . (10) etail of the trichomes resent on the adaial srface of the leaf sheath of icr . The arrows indicate the cells with silica bodies. (Fb = ber bundle; Fs = ber strand). Scale bar = 50µm (1, 3, 5, 7-9); 25µm (2, 4, 6, 10). marin. Sclerenchyma may occr st below chlorenchyma may be fond sch as in aans eidermis as in icr (i. 17) Ceasen niaerais (i. 19) aaea adsa (i. 21) riedeians (i. 25) and sarrsa (i. 27) or and ei (i. 23).
An Acad Bras Cienc (2015) 87 (1) NA . AIN AIN OIANI and A . SCANA
Figures 11-16 - Cross sections of the leaf sheath of aateoideae secies. (11) aans icr . (12) niaerais . (13) aaea ei . (1) adsa . (15 - 16) Ceasen riedeians . he arrow indicates a henolic idioblast. (b = ber bundle; Fs = ber strand). Scale bar = 100µm (11, 13-16); 50µm (12).
he anatomy of the leaf sheath is similar to mlticelllar niseriate trichomes on the ada ial the blade e cet for the tye of chlorenchyma srface (i. 10) that rodce mcilae. he which is comosed of arm cells in the leaf sheath marin of the sheath is formed by the e tension of (is. 11-15). In all stdied secies the sheath has the eidermis (i. 16).
An Acad Bras Cienc (2015) 87 (1) ANAO O APAOIA 163
Figures 17-28 - Cross sections of the leaf blade of aateoideae secies showin details of the marin and mesohyll. (17-18) aans icr . (19-20) niaerais . (21-22) aaea adsa (23-2) ei . (25- 26) Ceasen riedeians . (27-28) cea sarrsa . he arrow indicates a henolic idioblast. Scale bar = 50µm (17, 19, 21, 23, 25, 27); 70µm (18, 20, 22, 24, 26, 28).
An Acad Bras Cienc (2015) 87 (1) 164 NA . AIN AIN OIANI and A . SCANA
Figures 29-34 - Cross sections of the leaf midrib of aateoideae secies. (29) aans niaerais . (30) icr . (31) Ceasen riedeians . (32) cea sarrsa . (33) aaea adsa . (3) ei . (Fb = ber bundle; Fs = ber strand). Scale bar = 200µm (29- 30, 34); 50µm (31-32); 100µm (33).
An Acad Bras Cienc (2015) 87 (1) ANAO O APAOIA
Figures 35-46 - Cross sections of the in orescence axes of Rapateoideae species. (35) General view, (36) detail of the vasclar bndle and (37) detail of the eidermis of aans icr . (38) eneral view (39) detail of the vasclar bndle and (0-1) details of the eidermis of niaerais . (2) eneral view (3) detail of the oosite win () detail of the vasclar bndle and (5-6) details of the eidermis of aaea adsa . he arrows indicate the henolic idioblasts and the arrowheads indicate the epidermal cells with silica bodies. (Fb = ber bundle; Fs = ber strand). Scale bar = 500µm (35, 38); 100µm (36, 39, 44); 50µm (37, 40, 45); 25µm (41, 46); 200µm (42-43).
An Acad Bras Cienc (2015) 87 (1) NA . AIN AIN OIANI and A . SCANA
Figures 47-58 - Cross sections of the in orescence axes of Rapateoideae species. (47) General view, (48) detail of the trichome, (9) detail of the vasclar bndle and (50-51) details of the eidermis of aaea ei . (52) eneral view (53) detail of the vasclar bndle and (5) detail of the eidermis of Ceasen riedeians . (55) eneral view (56) detail of the vasclar bndle and (57-58) details of the eidermis of cea sarrsa . he arrows indicate the henolic idioblasts and the arrowheads indicate the epidermal cells with silica bodies. (Fs = ber strand; Vb = vascular bundle). Scale bar = 500µm (47); 50µm (48, 50, 54, 57); 100 µm (49, 53, 56); 25µm (51, 58); 200 µm (52, 55).
An Acad Bras Cienc (2015) 87 (1) ANAO O APAOIA 167
IN O SC NC AIS resents henolic idioblasts (arrows) sbeidermal he anatomical characteristics that differentiate ber strands (Fs) (Fig. 54) and ber bundles facing enera or secies are resented in able III. he the larer vasclar bndles formin the ribs (i. 53). cross-section shape of the in orescence axis varies All secies resent collateral vasclar bndles amon the enera. It is trianlar in aans srronded by a doble sheath: the oter sheath is (is. 35 and 38) ellitical in aaea (is. 2 and comosed of thin-walled cells and the inner sheath 7) and circlar in Ceasen riedeians (i. is composed of thick-walled cells (Figs. 36, 39, 52) and cea sarrsa (i. 55). In aaea the 9 53 and 56). In aans the vasclar in orescence axis has two wings, one opposite to bndles are simle (is. 36 and 39) while in the the other which are of different sizes in adsa remainin secies there are simle or comond (is. 2 and 3) and of the same size with vasclar vasclar bndles (is. 2 9 53 and 56). bndles in ei (i. 7). In C riedeians the arer vasclar bndles in nmber of 5 to 10 in orescence axis presents ribs at the region of the have a circlar distribtion in aaea (is. 2 larer vasclar bndles (is. 52 and 53). and 7) and in cea sarrsa (i. 55) while he eidermis is sinle-layered in all stdied smaller vasclar bndles are randomly distribted in species and has cells with slightly thickened outer the arenchyma. In aans (is. 35 and 38) ericlinal walls (is. 37 0 1 5 6 50 51 5 the larer vasclar bndles are more nmeros ( 10) 57 and 58) and tetracytic stomata. he eidermal cells and both smaller and larer ones are randomly may contain silica bodies (is. 0 1 5 6 50 51 distribted. In Ceasen riedeians the larer 57 and 58 - arrowheads) or henolic comonds (is. vasclar bndles in nmber of 5 to 10 alternate with 36 5 and 5). hey are aillose in aans the smaller ones formin the rin that delimits a niaerais (i. 0) and cea sarrsa (i. 57) parenchymatous, stulous pith (Fig. 52). and ronded in the remainin secies (is. 37 5 50 DISCUSSION and 5). lticelllar niseriate trichomes are resent in niaerais aaea ei and Ceasen Accordin to the reslts the leaves of the stdied riedeians . In ei these trichomes contain secies can be characterized by the resence of a henolic comonds (i. 8). sinle-layered eidermis with silica cells tetracytic In most of the stdied secies the corte is not stomata a relar licate or arm chlorenchyma de ned (Figs. 35, 38, 42, 47 and 55) and the vascular in the mesohyll and collateral doble-sheathed bndles are randomly distribted in a arenchyma vasclar bndles. he leaf sheath has mcilae- which is constitted by isodiametric and arm cells secretin trichomes on the ada ial srface a that delimit air canals (is. 35-39 9 50 55- chlorenchyma comosed of arm cells and marin 57). Subepidermal ber strands (Fs) and phenolic formed by the e tension of the eidermis. idioblasts (arrows) are also resent (is. 37 39 The in orescence axes are characterized 0 5 9 50 56 and 57). In addition to the by havin a sinle-layered eidermis with silica subepidermal ber strands, there are also ber cells tetracytic stomata a arenchyma comosed bndles (b) randomly scattered in the arenchyma of isodiametric or arm cells delimitin air canals of in orescence axis of aans secies (is. henolic idioblasts and collateral vasclar bndles. 37 and 39). Ceasen riedeians is the only hese featres were also observed in the leaves species that presents a de ned cortex, with vascular and in orescence axes of other representatives of bndles formin a rin (i. 52). In this secies the aateaceae (Carlist 1966 1969 errari et al. corte is comosed of sony chlorenchyma and 201) and may be sed to describe the family.
An Acad Bras Cienc (2015) 87 (1) 8 NA . AIN AIN OIANI and A . SCANA
TABLE III Anatomical characteristics of the in orescence axes of Rapateoideae (Rapateaceae) with taxonomic value. 1: Spathanthus unilateralis , 2: S. bicolor , 3: Rapatea paludosa , 4: R. ulei , 5: Cephalostemon riedelianus , 6: Duckea squarrosa (+ = present, - = absent).
Characteristics 3 4 rianlar shae in cross-section - - - - llitical shae in cross-section - - - - Circlar shae in cross-section - - - - neal wins withot vasclar bndles - - - - - ins of the same size with vasclar bndles - - - - - ibs in the reion of the larer vasclar bndles - - - - - richomes with henolic comonds - - - - - iber bndles scattered in the arenchyma - - - - De ned cortex - - - -- iber bndles facin the larer vasclar bndles - - - - - Simle vasclar bndles - - - - Simle and comond vasclar bndles - - Nmber of the larer vasclar bndles ranin from 5-10 - - Nmber of the larer vasclar bndles above 10 - - - - asclar bndles with a cylindrical distribtion formin a rin - - - - - istlos ith - - - - -
Accordin to Carlist (1969) the resence most of the secies stdied here the corte of the of aracytic stomata only on the aba ial srface of in orescence axis is not de ned and the vascular the leaves is a distinctive featre of aateaceae. bndles are randomly distribted in the arenchyma. he reslts obtained here however showed that he resence of silica bodies a common this characteristic varies amon the enera which characteristic of leaves and in orescence axes may inclde secies with stomata on both leaf of all stdied secies was also observed in other srfaces. Or reslts also showed that the stomata orans of aateaceae sch as stems bracts are tetracytic in areement with the reslts obtained and owers (Carlquist 1966, 1969, Stevenson by errari et al. (201) for secies of nrea et al. 1998 Oriani and Scatena 2012). Carlist eeis and aridericia therefore this is (1966) sed the resence of silica cells associated robably a characteristic of the family. with subepidermal ber strands in the stem to Carlist (1969) also reorted the resence characterize the sbfamily aateoideae. or the of central vasclar bndles enclosed by a leaves and in orescence axes, however, epidermal common brous sheath in the in orescence axes cells with silica bodies also occr in secies as a characteristic of the family. Never theless in that belon to different sbfamilies (Carlist
An Acad Bras Cienc (2015) 87 (1) ANAO O APAOIA
1969, Ferrari et al. 2014). The likely function of reviosly inclded in the a ateoideae (aire silica bodies is to redce the transiration and to 1958) corroboratin the actal circmscri tion of improve photosynthesis, through ef cient light the sbfamily ( ivnish et al. 200). distribtion in the arenchyma (Prychid et al. he data obtained in this stdy also show the 200). hey may also hel maintain the riidity of occrrence of characteristics that delimit the enera. stems and in orescence axes and reduce herbivory a an s s ecies have leaves with randomly on leaves (Prychid et al. 200). Silica bodies are distributed silica cells in the epidermis, ber resent in reresentatives of other families of bndles scattered in the meso hyll and a licated Poales as Poaceae Cyeraceae hrniaceae chlorenchyma. In these species, the in orescence Bromeliaceae Centroleidaceae cdeiocoleaceae aes are trianlar in cross-section and resent and oinvilleaceae (omlinson 1969 ahlren bers bundles randomly distributed in the et al. 1985 Silva and Alini 2003 Prychid et arenchyma and above ten sim le vasclar bndles al. 200 Arrda and Neves 2005 Proena and as dianostic characteristics. aaea s ecies Sao 2007). hey also occr in other commelinid have leaves with stomata and silica bodies only families (inclded in Arecales iniberales and on the abaial e idermis and with a chlorenchyma Commelinales) and may re resent a syna omor hy com osed of arm cells besides elli tical and wined of the clade with several secondary reversals in orescence axes. Ce a sen is distinished (Prychid et al. 200). by resentin leaves with e idermal cells with he mcilae-secretin trichomes in the straiht to slihtly sinos walls in frontal view leaf sheath were re orted for different enera of and ribbed in orescence axes with a de ned cortex, a ateaceae (Carl ist 1969 errari et al. 201) ber bundles facing the larger vascular bundles, and may re resent an a omor hy of the family. he and a stulous pith. fnction of the mcilae may be to rotect the leaves hen we com are the stdied enera it from solar irradiation and this featre may be related may be noticed that Ce a sen and cea to the environment where the family arose which share the reatest nmber of featres followed by was at the lowlands of the ayana Shield ( ivnish aaea and cea . he anatomical similarities et al. 2000 200). he mcilae rodction in leaves between Ce a sen and cea re ect their may even be related to different fnctions de endin mor holoical similarities already re orted by on the ty e of sbstance rodced. It may rotect other athors who even synonymized cea the leaves not only from intense liht bt also from as Ce a sen (Steyermark 1988). The high herbivores and athoens etreme tem eratres similarity between Ce a sen cea and and excessive water loss (Werker 2000). aaea also corroborates the hyloenetic analysis Or reslts show that the followin of the family in which Ce a sen and aaea characteristics are eclsive of a ateoideae: a ear as sister ro s ( ivnish et al. 2000 200). leaves and in orescence axes with epidermal cells Besides delimitin the enera some charac - with slightly thickened outer periclinal walls and teristics roved to be dianostic for s ecies sch subepidermal ber strands; vascular bundles of as the cross-section sha e and strctre of the leaf the leaf midrib region with a conspicuous ber midrib. he size and com osition of the wins resent sheath on the phloem; and in orescence axes in the in orescence axis of aaea differentiate the with henolic idioblasts in the e idermis and s ecies from each other. aaea a dsa is also arenchyma. hese featres are not resent in distinished by resentin vasclar bndle sheath s ecies of nrea ( errari et al. 201) a ens etensions facin both leaf srfaces.
An Acad Bras Cienc (2015) 87 (1) 170 NA . AIN AIN OIANI and A . SCANA
It may be em hasized that the anatomical com coros silicosos com distribio aleatria e characteristics identi ed in the leaves and parênquima cloro liano plicado no limbo foliar; e presença in orescence axes of Rapateoideae not only have de feixes de bras no meso lo e no parênquima do eixo taonomic vale bt may also have an ada tive da in orescência. a an s distin e-se tambm elo vale. he resence of silica bodies in both orans nmero tio e distribio dos feies vasclares no eio and mcilae- rodcin trichomes in the leaf da in orescência. O gênero aaea caracteriza-se ela sheath are eam les whose robable fnction has resena de estmatos e cllas eidrmicas com coros been discssed reviosly. he resence of air silicosos aenas na face abaial da folha e ela resena canals in the leaves and in orescence axes may also de parênquima cloro liano braciforme no limbo foliar. be related to environment. Since the s ecies row So caracteres com valor dia nstico ara Ce a sen predominantly in wet or ooded areas (Berry 2004, riede ian s : folha com cllas eidrmicas com aredes ivnish et al. 2000) the resence of air canals may retas a levemente sinosas em vista frontal e eio increase the aeration of these orans ths facilitatin da in orescência com córtex de nido, com cordões as echane (awase and hitmoyer 1980). esclerenim ticos sbeidrmicos voltados aos feies Arm cells delimiting air canals and ber bundles vasculares maiores e com medula stulosa. Dessa scattered in the meso hyll are also resent in the forma a artir de caractersticas anatmicas de folha leaves of some s ecies of Bromeliaceae (Proena e eixo da in orescência é possível distinguir gêneros e and Sao 2007 omlinson 1969) corroboratin the escies de aateaceae. As caractersticas levantadas hyloenetic roimity between these two families tambm ermitem diferenciar aateoideae das demais which have a basal osition in Poales (inder and sbfamlias de aateaceae. dall 2005 ivnish et al. 2010). Palavras-chave : anatomia Ce a sen cea
ACKNOWLEDGMENTS Poales aaea aateaceae a an s
The authors thank Conselho Nacional de REFERENCES Desenvolvimento Cientí co e Tecnológico (CNPq) A A CO AN N S . 2005 . Anatomia foliar de ( rant nmbers: 11582/2007-3 301692/2010-6 rieis iana Nees e rieis ciiai ia . oyama 71837/2011-3 and 11760/2012-3) and ndao (Cyeraceae) ss. Acta Bot Bras 19: 889-897. de Amaro Pesisa do stado de So Palo B P . 200 . aateaceae. In: Berry PE, Y atskievych K and olst B (ds) lora of the enezelan ayana (APSP) (rant nmbers: 2011/18275-0 and Poaceae- biaceae St ois: issori Botanical arden 2011/11536-3) for nancial support. We also thank Press SA . 13- 72. Beatriz de Aino ibeiro isboa for lo istical CA €IS S. 1961 . Pollen morholo y of aateaceae. Aliso 5: 39-66. sort in the Pare Nacional do ir . CA €IS S. 1966 . Anatomy of aateaceae: roots and stems. Phytomorholo y 16: 17-38. RESUMO CA €IS S. 1969 . aateaceae. In: Tomlinson PB (d) Anatomy of the onocotyledons III Commelinales- Estudou-se a anatomia de folha e eixo da in orescência de •in iberales ondon: Oford niversity Press n land a an s (2 s.) aaea (2 s.) Ce a sen . 130-1 5. (1 s.) e cea (1 s.) ( aateoideae aateaceae) A N C IO AN O P . 1985 . he families of the monocotyledons: strctre evoltion and bscando levantar caracteres teis ara a taonomia. A taonomy. Berlin: Srin er- erla . forma do eixo da in orescência em secção transversal N AN O‚B IN P . 1968 . Plant microtechnie: some diferencia os êneros enanto a forma e estrtra da rinciles and new methods. Amer Bot 55: 123-1 2. A I C SCANA AN O IANI A. 201 . eaf and nervura central do limbo foliar tem valor especí co. São in orescence peduncle anatomy: a contribution to the caracteres eclsivos de a an s : cllas eidrmicas taonomy of aateaceae. Plant Syst vol 300: 1579-1590.
An Acad Bras Cienc (2015) 87 (1) ANAO O APAOIA 171
I NIS A . 2010 . Assemblin to the tree of the OIANI A AN SCA NA . 2012 . he taonomic vale monocotyledons: lastome seence hylo eny and of the oral characters in Rapateaceae (Poales - evoltion of Poales. Ann O Bot ard 97: 58 -616. onocotyledons). Plant Syst vol 299: 291-303. I NIS ANS • A PA SON B B PO N A S AN SA O . 2007 . Anatomia foliar de P AN SS A . 2000 . oleclar evoltion bromlias ocorrentes em reas de cerrado do stado de adaptive radiation, and geographic diversi cation in the S o Palo Brasil. Acta Bot Bras 21: 657-673. amhiatlantic family aateaceae: vidence from nd PC I C A P AN O . 200 . Systematics seences and morholo y. voltion 5 : 1915-1937. and bioloy of silica bodies in monocotyledons. Bot ev I NIS I A C ANS A C PI S C 69: 377- 0. B P AN SS A . 200 . Ancient vicariance O S . 1972 . Die Nadel der Schwarzkiefer-Masenprodukt or recent lon-distance disersal Inferences abot und Keinstwert der Natur. Mikrokosmos 61: 33-36. hyloeny and soth american-african disnctions in SIA AN A INI . 2003 . Anatomia comarativa de aateaceae and Bromeliaceae based on nd seence folhas e cales de A n s scari s (l€e) hlm. data. Int Plant Sci 165: 35-5 . e A n s ssifolius (addi) hlm. (Poaceae). Braz O ANS N A . 19 0 . Plant microtechnique. New York: Bot 26: 185-192. craw ill. S NSON CO A AN BOO B. 1998 . aateaceae. AAS AN IO . 1980 . Aerenchyma In: Kubitzki K ( d) he families and enera of vasclar develoment in waterloed lants. Amer Bot 67: 18-22. lants - I onocotyledons Berlin: Sriner ermany AI B. 1958 . aateaceae. In: Maguire B and Wurdack . 15- 2 . ( ds) he botany of ayana ihland Part III. em S A A . 1988 . lora of the enezelan ayana - I. N Bot ard 10: 19- 9. Ann O Bot ard 75: 1565-1586. AI B. 1965 . aateaceae. In: aire B ( d) he OINSON PB . 1969 . Commelinales-•iniberales. In: botany of the ayana ihland Part I. em N Bot mET calf E cR ( d) Anatomy of the monocotyledons ard 12: 69-102. Oford: Clarendon Press SA . 193-29 . ON IO . 201 . aateaceae. In: ista de escies N I AN BOAN . 1988 . eveloment of ovle da lora do Brasil. ardim Botnico do io de aneiro. and seed in aateaceae. Bot inn Soc 97: 267-29 . http:// oradobrasil.jbrj.gov.br/jabot/ oradobrasil/FB205. . 2000 . richome diversity and develoment. In: Accessed 1 ay 201 . allahan and ray C ( ds) Plant trichomes ondon: IN P AN A P . 2005 . voltionary history of Academic Press nland . 1-30. Poales. Ann ev col vol Syst 36: 107-12 . OBI N P N AN CC . 196 . Polychromatic stainin of lant cell walls by tolidine ble O. Protolasma 59: 368-373.
An Acad Bras Cienc (2015) 87 (1)