Garcia et al 2012

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Revista Mexicana de Biodiversidad 83: 1036-1044, 2012 DOI: 10.7550/rmb.28124 Recibido: 20 octubre 2011; aceptado: 02 junio 2012 Stem and root anatomy of two species of Echinopsis (Trichocereeae: Cactaceae) Anatomía de la raíz y del tallo de dos especies de Echinopsis (Trichocereeae: Cactaceae) Joelma dos Santos Garcia 1 , Edna Scremin-Dias 1 DQG 3DWULFLD 6RI¿DWWL 2 1 Universidade Federal de Mato Grosso do Sul, CCBS, Departamento de Biologia, Programa de Pós Graduação em Biologia Vegetal Cidade Universitária, S/N, Caixa Postal 549, CEP 79.070.900 Campo Grande, MS, Brasil. 2 Universidade Federal do Paraná, SCB, Departamento de Botânica, Programa de Pós-Graduação em Botânica, Caixa Postal 19031, CEP 81.531.990 Curitiba, PR, Brasil. SVRI¿DWWLXISU#JPDLOFRP Abstract. This study characterizes and compares the stem and root anatomy of Echinopsis calochlora and E. rhodotricha (Cactaceae) occurring in the Central-Western Region of Brazil, in Mato Grosso do Sul State. Three individuals of each VSHFLHV ZHUH FROOHFWHG ¿[HG VWRUHG DQG SUHSDUHG IROORZLQJ XVXDO DQDWRP\ WHFKQLTXHV IRU VXEVHTXHQW REVHUYDWLRQ LQ light and scanning electronic microscopy. Echinopsis calochlora revealed uniseriated epidermis, while E. rhodotricha KDG SDWFKHV RI ELVVHULDWHG HSLGHUPLV DOO VSHFLHV VKRZHG WKLFN FXWLFOH SDUDOOHORF\WLF VWRPDWD DW WKH HSLGHUPLV OHYHO DQG D ZHOOGHYHORSHG K\SRGHUPLV &RUWLFDO DQG PHGXOODU\ EXQGOHV DUH SUHVHQW LQ WKH VWXGLHG VSHFLHV DV ZHOO DV PXFLODJH FHOOV LQ WKH FRUWH[ UHJLRQ 7KH VHFRQGDU\ SKORHP LV FRPSRVHG E\ VLHYH WXEH HOHPHQWV FRPSDQLRQ FHOOV D[LDO DQG UDGLDO SDUHQFK\PD 6FOHUHLGV ZHUH IRXQG DW WKH RXWHU UHJLRQV RI SKORHP LQ WKH URRWV 7KH VHFRQGDU\ [\OHP LV QRQ ¿EURXV LQ WKH VWHPV RI E. calochlora DQG ¿EURXV LQ WKH VWHPV RI E. rhodotricha DQG LQ WKH URRWV RI ERWK VSHFLHV 0DQ\ RI WKHVH FKDUDFWHULVWLFV DUH FRPPRQO\ IRXQG LQ &DFWDFHDH DQG UHSUHVHQW LPSRUWDQW DGDSWDWLRQV IRU VXUYLYDO LQ [HULF environments. .H\ ZRUGV FRUWLFDO EXQGOHV HSLGHUPLV QRQ ¿EURXV ZRRG ZRRG DQDWRP\ Resumen. Este estudio está enfocado a caracterizar y comparar la anatomía de tallos y raíces de Echinopsis calochlora y E. rhodotricha &DFWDFHDH TXH KDELWDQ HQ OD UHJLyQ FHQWURRHVWH GH %UDVLO HQ HO (VWDGR GH 0DWR *URVVR GR 6XO 6H UHFROHFWDURQ LQGLYLGXRV GH FDGD HVSHFLH ORV FXDOHV IXHURQ ¿MDGRV DOPDFHQDGRV \ SUHSDUDGRV VLJXLHQGR ODV WpFQLFDV FRPXQHV GH DQDWRPtD SDUD REVHUYDUORV HQ PLFURVFRStD GH OX] \ HOHFWUyQLFD GH EDUULGR Echinopsis calochlora PRVWUy HSLGHUPLV XQLVHULDGD PLHQWUDV TXH OD GH E. rhodotricha IXH ELVHULDGD WRGDV ODV HVSHFLHV SUHVHQWDURQ FXWtFXOD JUXHVD HVWRPDV SDUDOHORFtWLFRV D QLYHO GH OD HSLGHUPLV \ XQD KLSRGHUPLV ELHQ GHVDUUROODGD 6H SUHVHQWDURQ KDFHV YDVFXODUHV FRUWLFDOHV \ PHGXODUHV HQ ODV HVSHFLHV HVWXGLDGDV DVt FRPR FpOXODV PXFLODJLQRVDV HQ OD UHJLyQ FRUWLFDO (O ÀRHPD VHFXQGDULR HVWi FRPSXHVWR GH HOHPHQWRV GH WXER FULERVR FpOXODV DFRPSDxDQWHV \ SDUpQTXLPD D[LDO \ UDGLDO 6H HQFRQWUDURQ HVFOHUHLGDV HQ ODV UHJLRQHV H[WHUQDV GHO ÀRHPD HQ ODV UDtFHV (O [LOHPD VHFXQGDULR HV QR¿EURVR HQ los tallos de E. calochlora \ ¿EURVR HQ ORV GH E. rhodotricha \ HQ ODV UDtFHV GH DPEDV HVSHFLHV 0XFKDV GH HVWDV FDUDFWHUtVWLFDV HVWiQ SUHVHQWHV FRP~QPHQWH HQ HVSHFLHV GH &DFWDFHDH OR TXH UHSUHVHQWD DGDSWDFLRQHV LPSRUWDQWHV SDUD OD VXSHUYLYHQFLD HQ DPELHQWHV [pULFRV 3DODEUDV FODYH KDFHV FRUWLFDOHV HSLGHUPLV PDGHUD QR¿EURVD DQDWRPtD GH OD PDGHUD *LEVRQ DQG 1REHO 6RI¿DWWL DQG $QJ\DORVV\ 7HUUD]DV DQG $ULDV $UUXGD HW DO considering the representativeness of the family, and the SUREOHPV LQ FLUFXPVFULELQJ JURXSV HVSHFLDOO\ ZLWKLQ WULEHV DV ZHOO DV HQDEOLQJ D EHWWHU XQGHUVWDQGLQJ RI WKH JURXS HYROXWLRQ 7HUUD]DV DQG $ULDV DQG WKHLU adaptations for survival in arid environments (Mauseth, (JJOL PHQWLRQHG WKH ODFN RI LQIRUPDWLRQ RQ Cactaceae in Brazil in the Brazilian southwestern region, HODERUDWLQJ D FKHFNOLVW RI QDWLYH VSHFLHV IRU WKH VWDWHV RI 0DWR *URVVR DQG 0DWR *URVVR GR 6XO )URP WKRVH Introduction 7KH VXEIDPLO\ &DFWRLGHDH &DFWDFHDH FRPSULVHV WKH ODUJHVW QXPEHU RI VSHFLHV LV WKH PRVW GLYHUVL¿HG LQ WHUPV RI OLIHIRUPV DQG KDELW 7D\ORU 7HUUD]DV DQG $ULDV DQG LW LV GLYLGHG LQ WULEHV LQ ZKLFK Cereeae, Rhipsalideae, Trichocereeae, Echinocereeae, and +\ORFHUHHDH RFFXU LQ %UD]LO 7D\ORU DQG =DSSL The importance of anatomical studies in Cactaceae DUH XQGHQLDEOH *LEVRQ DQG +RUDN *DVVRQ

Transcript of Garcia et al 2012

Revista Mexicana de Biodiversidad 83: 1036-1044, 2012

DOI: 10.7550/rmb.28124

Recibido: 20 octubre 2011; aceptado: 02 junio 2012

Stem and root anatomy of two species of Echinopsis (Trichocereeae: Cactaceae)

Anatomía de la raíz y del tallo de dos especies de Echinopsis (Trichocereeae: Cactaceae)

Joelma dos Santos Garcia1, Edna Scremin-Dias1 DQG�3DWULFLD�6RI¿DWWL2

1Universidade Federal de Mato Grosso do Sul, CCBS, Departamento de Biologia, Programa de Pós Graduação em Biologia Vegetal Cidade

Universitária, S/N, Caixa Postal 549, CEP 79.070.900 Campo Grande, MS, Brasil.

2Universidade Federal do Paraná, SCB, Departamento de Botânica, Programa de Pós-Graduação em Botânica, Caixa Postal 19031, CEP 81.531.990

Curitiba, PR, Brasil.

�SVRI¿DWWL�XISU#JPDLO�FRP

Abstract. This study characterizes and compares the stem and root anatomy of Echinopsis calochlora and E. rhodotricha

(Cactaceae) occurring in the Central-Western Region of Brazil, in Mato Grosso do Sul State. Three individuals of each

VSHFLHV�ZHUH�FROOHFWHG��¿[HG��VWRUHG�DQG�SUHSDUHG�IROORZLQJ�XVXDO�DQDWRP\�WHFKQLTXHV��IRU�VXEVHTXHQW�REVHUYDWLRQ�LQ�light and scanning electronic microscopy. Echinopsis calochlora revealed uniseriated epidermis, while E. rhodotricha

KDG�SDWFKHV�RI�ELVVHULDWHG�HSLGHUPLV��DOO�VSHFLHV�VKRZHG�WKLFN�FXWLFOH��SDUDOOHORF\WLF�VWRPDWD�DW�WKH�HSLGHUPLV�OHYHO��DQG�D�ZHOO�GHYHORSHG�K\SRGHUPLV��&RUWLFDO�DQG�PHGXOODU\�EXQGOHV�DUH�SUHVHQW�LQ�WKH�VWXGLHG�VSHFLHV��DV�ZHOO�DV�PXFLODJH�FHOOV� LQ� WKH� FRUWH[� UHJLRQ��7KH� VHFRQGDU\�SKORHP� LV� FRPSRVHG�E\� VLHYH� WXEH� HOHPHQWV�� FRPSDQLRQ�FHOOV�� D[LDO� DQG�UDGLDO�SDUHQFK\PD��6FOHUHLGV�ZHUH� IRXQG�DW� WKH�RXWHU� UHJLRQV�RI�SKORHP� LQ� WKH� URRWV��7KH� VHFRQGDU\�[\OHP� LV�QRQ�¿EURXV�LQ�WKH�VWHPV�RI�E. calochlora��DQG�¿EURXV�LQ�WKH�VWHPV�RI�E. rhodotricha�DQG�LQ�WKH�URRWV�RI�ERWK�VSHFLHV��0DQ\�RI�WKHVH�FKDUDFWHULVWLFV�DUH�FRPPRQO\�IRXQG�LQ�&DFWDFHDH��DQG�UHSUHVHQW�LPSRUWDQW�DGDSWDWLRQV�IRU�VXUYLYDO�LQ�[HULF�environments.

.H\�ZRUGV��FRUWLFDO�EXQGOHV��HSLGHUPLV��QRQ�¿EURXV�ZRRG��ZRRG�DQDWRP\�

Resumen. Este estudio está enfocado a caracterizar y comparar la anatomía de tallos y raíces de Echinopsis calochlora

y E. rhodotricha� �&DFWDFHDH��TXH�KDELWDQ�HQ�OD�UHJLyQ�FHQWUR�RHVWH�GH�%UDVLO��HQ�HO�(VWDGR�GH�0DWR�*URVVR�GR�6XO��6H� UHFROHFWDURQ� �� LQGLYLGXRV� GH� FDGD� HVSHFLH�� ORV� FXDOHV� IXHURQ� ¿MDGRV�� DOPDFHQDGRV� \� SUHSDUDGRV� VLJXLHQGR� ODV�WpFQLFDV�FRPXQHV�GH�DQDWRPtD��SDUD�REVHUYDUORV�HQ�PLFURVFRStD�GH�OX]�\�HOHFWUyQLFD�GH�EDUULGR��Echinopsis calochlora

PRVWUy�HSLGHUPLV�XQLVHULDGD��PLHQWUDV�TXH�OD�GH�E. rhodotricha�IXH�ELVHULDGD��WRGDV�ODV�HVSHFLHV�SUHVHQWDURQ�FXWtFXOD�JUXHVD�� HVWRPDV� SDUDOHORFtWLFRV� D� QLYHO� GH� OD� HSLGHUPLV� \� XQD� KLSRGHUPLV� ELHQ� GHVDUUROODGD�� 6H� SUHVHQWDURQ� KDFHV�YDVFXODUHV�FRUWLFDOHV�\�PHGXODUHV�HQ�ODV�HVSHFLHV�HVWXGLDGDV��DVt�FRPR�FpOXODV�PXFLODJLQRVDV�HQ�OD�UHJLyQ�FRUWLFDO��(O�ÀRHPD�VHFXQGDULR�HVWi�FRPSXHVWR�GH�HOHPHQWRV�GH�WXER�FULERVR��FpOXODV�DFRPSDxDQWHV�\�SDUpQTXLPD�D[LDO�\�UDGLDO��6H�HQFRQWUDURQ�HVFOHUHLGDV�HQ� ODV�UHJLRQHV�H[WHUQDV�GHO�ÀRHPD�HQ� ODV�UDtFHV��(O�[LOHPD�VHFXQGDULR�HV�QR�¿EURVR�HQ�los tallos de E. calochlora� \�¿EURVR� HQ� ORV�GH�E. rhodotricha� \� HQ� ODV� UDtFHV�GH� DPEDV� HVSHFLHV��0XFKDV�GH� HVWDV�FDUDFWHUtVWLFDV�HVWiQ�SUHVHQWHV�FRP~QPHQWH�HQ�HVSHFLHV�GH�&DFWDFHDH��OR�TXH�UHSUHVHQWD�DGDSWDFLRQHV�LPSRUWDQWHV�SDUD�OD�VXSHUYLYHQFLD�HQ�DPELHQWHV�[pULFRV�

3DODEUDV�FODYH��KDFHV�FRUWLFDOHV��HSLGHUPLV��PDGHUD�QR�¿EURVD��DQDWRPtD�GH�OD�PDGHUD�

*LEVRQ�DQG�1REHO��������6RI¿DWWL�DQG�$QJ\DORVV\��������������������7HUUD]DV�DQG�$ULDV��������$UUXGD�HW�DO����������considering the representativeness of the family, and the

SUREOHPV� LQ� FLUFXPVFULELQJ� JURXSV�� HVSHFLDOO\� ZLWKLQ�WULEHV�� DV�ZHOO� DV� HQDEOLQJ� D� EHWWHU� XQGHUVWDQGLQJ� RI� WKH�JURXS� HYROXWLRQ� �7HUUD]DV� DQG� $ULDV�� ������ DQG� WKHLU�adaptations for survival in arid environments (Mauseth,

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Cactaceae in Brazil in the Brazilian southwestern region,

HODERUDWLQJ�D�FKHFNOLVW�RI����QDWLYH�VSHFLHV�IRU�WKH�VWDWHV�RI�0DWR�*URVVR�DQG�0DWR�*URVVR�GR�6XO��)URP�WKRVH�����

Introduction

7KH� VXEIDPLO\� &DFWRLGHDH� �&DFWDFHDH�� FRPSULVHV�WKH� ODUJHVW� QXPEHU� RI� VSHFLHV�� LV� WKH�PRVW� GLYHUVL¿HG� LQ�WHUPV� RI� OLIH�IRUPV� DQG� KDELW� �7D\ORU�� ������ 7HUUD]DV�DQG�$ULDV�� ������� DQG� LW� LV� GLYLGHG� LQ� �� WULEHV�� LQ�ZKLFK�Cereeae, Rhipsalideae, Trichocereeae, Echinocereeae, and

+\ORFHUHHDH�RFFXU�LQ�%UD]LO��7D\ORU�DQG�=DSSL���������The importance of anatomical studies in Cactaceae

DUH�XQGHQLDEOH� �*LEVRQ�DQG�+RUDN��������*DVVRQ��������

����Revista Mexicana de Biodiversidad 83: 1036-1044, 2012

DOI: 10.7550/rmb.28124

VSHFLHV�EHORQJ�WR�&DFWRLGHDH��DQG���JHQHUD�ZHUH�UHFRUGHG�IRU�WULEH�7ULFKRFHUHHDH��Arthrocereus �$��%HUJHU��%DFNHE��et F.M. Knuth, Cleistocactus Lem., Discocactus Pfeiff.,

Echinopsis Zucc., Gymnocalycium Pfeiffer in Pfeiffer et

Otto, and Harrisia %ULWWRQ��DOO�EHLQJ�YHU\�SRRUO\�VWXGLHG��especially in the state of Mato Grosso do Sul.

Echinopsis is one of the largest genus of Trichocereeae,

FRPSRVHG�E\�����VSHFLHV�GLVWULEXWHG�LQ�VHYHUDO�UHJLRQV�RI�6RXWK�$PHULFD�� DQG� GXH� WR� WKH� ODUJH� QXPEHU� RI� VSHFLHV��LW�VKRZV�D�YHU\�FRQWURYHUVLDO�FLUFXPVFULSWLRQ��$QGHUVRQ�������� Echinopsis calochlora K. Schum. is endemic to the

KLJK�DOWLWXGHV�RI�WKH�0DFLoR�GR�8UXFXP��(JJOL���������LQ�western Mato Grosso do Sul, and was recently included

LQ�WKH�%UD]LOLDQ�OLVW�RI�WKUHDWHQHG�VSHFLHV��,%$0$���������EHFDXVH�LWV�KDELWDW�KDV�EHHQ�LQWHQVLYHO\�H[SORLWHG��IRU�LURQ�DQG� PDQJDQHVH� PLQLQJ� �6LOYD� HW� DO��� ������� Echinopsis

rhodotricha� .�� 6FKXP�� KDV� D� EURDGHU� GLVWULEXWLRQ��RFFXUULQJ� %UD]LO�� 3DUDJXD\� DQG� $UJHQWLQD� �$QGHUVRQ���������7KLV�ZRUN�GHVFULEHV�FRPSDUDWLYHO\�WKH�DQDWRP\�RI�stem and root of E. calochlora and E. rhodotricha.

Materials and methods

,QGLYLGXDOV� ZHUH� FROOHFWHG� LQ� URFN\� RXWFURSV� DQG�sandy-clay formations from natural populations as

follows: E. calochlora� DW� &RUXPEi� &RXQW\�� LQ� 3DQWDQDO��YRXFKHU�QXPEHU�&*06��������J. S. Garcia 54), and E.

rhodotricha��DW�3RUWR�0XUWLQKR��LQ�&KDFR��YRXFKHU�QXPEHU�&*06��������J. S. Garcia 61). Vouchers were deposited

DW� WKH�+HUEDULXP� ��&*06�8)06��)HGHUDO�8QLYHUVLW\�RI�Mato Grosso do Sul.

)RU� WKH� TXDOLWDWLYH� DQDWRPLFDO� VWXG\� RI� WKH� VWHP�� ��LQGLYLGXDOV� RI� HDFK� VSHFLHV� ZHUH� VHFWLRQHG� DW� WKH� EDVDO��PHGLXP��DQG�DSLFDO�UHJLRQV��)RU�WKH�URRW��RQO\�WKH�YHU\�EDVH�ZDV�XVHG��6DPSOHV�ZHUH�¿[HG�ZLWK�)$$����IRU����KRXUV��ULQVHG�LQ�ZDWHU�DQG�VWRUHG�LQ�HWKDQRO������-HQVHQ���������3DUDGHUPDO�VHFWLRQV�ZHUH�WDNHQ�DW�WKH�PHGLDQ�UHJLRQ�RI�WKH�stem. Transverse and longitudinal sections of stems and

URRWV�ZHUH�PDGH�E\�KDQG��6DPSOHV�ZHUH� DOVR� HPEHGGHG�LQ� SRO\HWK\OHQH� JO\FRO� �3(*�� ����� �5LFKWHU�� ������ DQG�transverse and longitudinal sections were made in rotative

microtome. Root samples of E. rhodotricha were sectioned

LQ�D�VOLGLQJ�PLFURWRPH��6HFWLRQV�ZHUH�VWDLQHG�ZLWK�DVWUDEOXH�DQG�VDIUDQLQ�DQG�PRXQWHG�LQ�JO\FHULQH������0DFHUDWLRQV�ZHUH�SUHSDUHG�XVLQJ�D�PRGL¿HG�)UDQNOLQ�PHWKRG��)UDQNOLQ���������VWDLQHG�ZLWK�VDIUDQLQ�DQG�PRXQWHG�LQ�JO\FHULQ�

The following hystochemical tests were carried out:

ÀRURJOXF\Q��-RKDQVHQ��������WR�GHWHFW�OLJQL¿HG�VHFRQGDU\�ZDOOV��OXJRO��-RKDQVHQ��������WR�GHWHFW�VWDUFK��IRU�PXFLODJH�ZH�IROORZHG�5LFKWHU���������7KH�DQDO\VLV�ZDV�FDUULHG�RXW�in a Leica DMLB light microscope, connected to an image

FDSWXUH�V\VWHP��DQG�D�GLJLWDO�FDPHUD�'&����)�

Paradermal and transversal sections of the dermal

system of the stem median region were prepared for

scanning electron microscopy (SEM) analysis. Samples

were dehydrated, critical-point dried, mounted on aluminum

VWXEV��DQG�FRDWHG�ZLWK�JROG�SDOODGLXP�LQ�D�VSXWWHU�V\VWHP��SEM analysis and electron micrographs were made with a

-(2/�-60������VFDQQLQJ�HOHFWURQ�PLFURVFRSH��

Results

Stem. Uniseriate epidermis (Fig. 1) occurred in

photosynthesizing regions of the stem, as well as in

DSLFDO� DQG� PHGLDQ� UHJLRQV�� ,W� ZDV� FRPSRVHG� E\� VTXDUH�WR� UHFWDQJXODU� FHOOV� LQ� FURVV� VHFWLRQV��ZLWK� WKLFN� FXWLFOH��Echinopsis rhodotricha� VKRZHG� SDWFKHV� RI� ELVVHULDWH�HSLGHUPLV� LQ� WKH� PHGLDQ� UHJLRQ� �)LJ�� ���� IRUPHG� E\�SHULFOLQDO�GLYLVLRQV�RI�HSLGHUPDO�FHOOV��UDUHO\�REVHUYHG�LQ�E. calochlora.

3DUDOOHORF\WLF� VWRPDWD�ZHUH� IRXQG� LQ� ERWK� VSHFLHV� DW�the same level of epidermal cells in E. calochlora (Fig.

1) while E. rhodotricha� KDG� VXQNHQ� VWRPDWD� �)LJ�� ����6XEVWRPDWDO�FKDPEHUV�FURVVHG�WKH�K\SRGHUPLV��)LJV���������

The phellogen was originated in the epidermis.

Peridermis covered the non photosynthesizing portions

RI� WKH�VWHP��)LJV���������FRPSRVHG�RI�SKHORGHUP�IRUPHG�E\�UHFWDQJXODU�FHOOV� LQ�FURVV�VHFWLRQ��ZLWK�SULPDU\�ZDOOV��&RUN�KDG�OD\HUV�RI�FHOOV�ZLWK�VXEHUL]HG�ZDOOV�WKDW�DOWHUQDWH�ZLWK� OD\HUV� RI� FHOOV�ZLWK� OLJQL¿HG�ZDOOV��ZKLFK� DUH� YHU\�numerous in E. rhodotricha��)LJ����.

:H�REVHUYHG�D�FROOHQFK\PDWLF�K\SRGHUPLV�LQ�WKH�VWHP��)LJV������������FRPSRVHG�RI�FHOOV�ZLWK�LUUHJXODUO\�WKLFNHQHG�SULPDU\�ZDOOV��FRQQHFWLQJ�WKURXJK�UDPL¿HG�FKDQQHOV��

&RUWH[� ZDV� GLYLGHG� LQ� DQ� RXWHU� UHJLRQ� IRUPLQJ� D�SDOLVDGH��)LJ������DQG�DQ� LQQHU� UHJLRQ�FRPSRVHG�RI� ODUJH�SDUHQFK\PD�FHOOV�DQG�YDVFXODU�EXQGOHV��)LJ������FRQVWLWXWHG�E\� SKORHP� DQG� [\OHP�� DUUDQJHG� FROODWHUDOO\�� 'UXVHV�ZHUH� UDUH� LQ� WKH� FRUWH[�� /DUJH� PXFLODJH� FHOOV� RFFXUUHG�WKURXJKRXW� WKH� ZKROH� FRUWH[� �)LJ�� ��� DQG� SLWK��9DVFXODU�EXQGOHV�RFFXUUHG�DOVR�LQ�WKH�SLWK��)LJ�����

$W� WKH� EDVH� RI� VWHP�� SULPDU\� SKORHP�ZDV� FROODSVHG�and in E. rhodotricha caps of sclereids were seen outside

VHFRQGDU\� SKORHP� �)LJ�� ���� 6HFRQGDU\� SKORHP� ZDV�FRPSRVHG�RI�QDUURZ�VLHYH�WXEH�HOHPHQWV�DQG�FRPSDQLRQ�FHOOV��ZLWK�VLPSOH�VLHYH�SODWH��WUDQVYHUVH�WR�VOLJKW�REOLTXH��D[LDO�DQG�UDGLDO�SDUHQFK\PD��

%RWK�VSHFLHV�KDG�QRQ�¿EURXV�ZRRG��PDLQO\�FRPSRVHG�RI� ZLGH� EDQG� WUDFKHLGV� DQG� D� IHZ� YHVVHO� HOHPHQWV��VXUURXQGHG� E\� D[LDO� DQG� UDGLDO� XQOLJQL¿HG� SDUHQFK\PD�FHOOV� �)LJV�� ������� :LGH� EDQG� WUDFKHLGV� SUHVHQWHG� ULQJ��KHOLFDO� RU� PL[HG� ZDOO� WKLFNHQLQJV� �)LJ�� ����� 9HVVHO�elements had simple perforation plates. Both species had

D�VWRULHG�VWUXFWXUH�LQ�WKH�QRQ�¿EURXV�ZRRG��)LJ�������EHLQJ�

���� Garcia et al.- Anatomy of two species of Echinopsis

Figures 1-6.� 6WHP� FURVV� VHFWLRQV�� ����� HSLGHUPLV� FRYHUHG� E\� D� WKLFN� FXWLFOH� �DUURZ��� 6WRPDWD� FKDPEHU� FURVVHG� K\SRGHUPLV�� ���Echinopsis calochlora. Uniseriate epidermis. Stomata at the same level as ordinary epidermal cells. 2, E. rhodotricha. Scanning electron

micrograph (SEM).�%LVVHULDWH�HSLGHUPLV�DQG�VXQNHQ�VWRPDWD�������SHULGHUPLV�ZLWK�SKHORGHUP�FRPSRVHG�RI�OD\HUV�RI�SDUHQFK\PD�FHOOV��FRUN�FRPSRVHG�RI�OD\HUV�RI�FHOOV�ZLWK�VXEHUL]HG�ZDOOV�DOWHUQDWLQJ�ZLWK�OD\HUV�RI�FHOOV�ZLWK�OLJQL¿HG�ZDOOV�(*). 3, E. calochlora�������E. rhodotricha�����RUJDQL]DWLRQ�RI�FRUWH[��K\SRGHUPLV�IROORZHG�E\�OD\HUV�RI�SDOLVDGH�SDUHQFK\PD�DQG�LQQHU�LVRGLDPHWULF�SDUHQFK\PD�FHOOV��&RUWLFDO�EXQGOHV��DUURZV������FRUWH[��1XPHURXV�VFDWWHUHG�PXFLODJH�FHOOV��6FDOH�EDUV��� ������P��� ����P����� ������P��� ������P��� ������P��&P��PXFLODJH�FHOO��(S��HSLGHUPLV��+S��K\SRGHUPLV��3S��SDOLVDGH�SDUHQFK\PD��6W��VWRPDWD��6X��VXEHU�

����Revista Mexicana de Biodiversidad 83: 1036-1044, 2012

DOI: 10.7550/rmb.28124

Figures 7-12.�����������FURVV�VHFWLRQV������WDQJHQWLDO�VHFWLRQ�����Echinopsis calochlora��3LWK�ZLWK�PHGXOODU\�EXQGOH�����E. rhodotricha.

9DVFXODU�V\VWHP��3KORHP��&DSV�RI�VFOHUHLGV�LQ�WKH�SHULSKHU\�RI�VHFRQGDU\�SKORHP�����E. calochlora��1RQ�¿EURXV�ZRRG�PDLQO\�FRPSRVHG�RI�ZLGH�EDQG�WUDFKHLGV������E. rhodotricha��)LEURXV�DQG�QRQ�¿EURXV�ZRRG������E. calochlora.�1RQ�¿EURXV�ZRRG�FRPSRVHG�RI�ZLGH�EDQG�WUDFKHLGV�ZLWK�KHOLFDO�WKLFNHQLQJV������E. rhodotricha.�)LEURXV�ZRRG�FRPSRVHG�RI�YHVVHO�HOHPHQWV��¿EUHV�DQG�VFDQW\�SDUDWUDFKHDO�SDUHQFK\PD��SDUW�RI�D�UD\�LV�VHHQ�RQ�WKH�OHIW���6FDOH�EDUV��������� ������P��� ������P��):��¿EURXV�ZRRG��1):��QRQ�¿EURXV�ZRRG�

���� Garcia et al.- Anatomy of two species of Echinopsis

most pronounced in E. calochlora��)LEURXV�ZRRG�RFFXUUHG�only in E. rhodotricha��RXWVLGH�QRQ�¿EURXV�ZRRG��)LJV������������� GLIIXVH�SRURXV��ZLWK�ZLGH�EDQG� WUDFKHLGV� DEVHQW��composed of vessels predominantly solitary and multiples

RI���WR����DQG�OHVV�IUHTXHQW��RI���WR����YHVVHO�HOHPHQWV�ZDOOV�KDG�KHOLFDO�WR�SLWWHG�VHFRQGDU\�ZDOOV��LQWHUYHVVHO�SLWV�ZHUH�ERUGHUHG��YHVVHO�SDUHQFK\PD�SLWV�VLPSOH�WR�KDOI�ERUGHUHG��VFDODULIRUP��RSSRVLWH�WR�DOWHUQDWH��SHUIRUDWLRQ�SODWHV�ZHUH�VLPSOH��)LJ�������WUDQVYHUVH�WR�REOLTXH��VRPH�YHVVHOV�KDG���SHUIRUDWLRQ�SODWHV��LQWHUPHGLDWH�IRUPV�EHWZHHQ�VFDODULIRUP�ZLWK� RQH� LQFRPSOHWH� EDU� WR� LUUHJXODU� DQG� UHWLFXODWH�ZHUH�REVHUYHG� �)LJ�� ����� 3DUHQFK\PD�ZDV� VFDQW\� SDUDWUDFKHDO�(Fig. 12). Multiseriate wide and high heterogeneous rays

�)LJ�� ����ZHUH� FRPSRVHG� RI� SURFXPEHQW�� XSULJKW� RU�DQG�VTXDUH�FHOOV��ZLWK�OLJQL¿HG�DQG�XQOLJQL¿HG�SRUWLRQV��VWDUFK�JUDLQV� RFFXUUHG� LQ� UD\� FHOOV� �)LJV�� ���� ����� /LEULIRUP�¿EUHV� ZHUH� SUHVHQW�� UDUH� VHSWDWH�� %DQGV� RI� XQOLJQL¿HG�SDUHQFK\PD� LQFOXGLQJ�D[LDO�DQG� UDGLDO�SDUHQFK\PD�ZHUH�SUHVHQW�LQ�¿EURXV�ZRRG��)LJ�������Root��$GXOW�URRWV�ZHUH�FRYHUHG�E\�D�SHULGHUPLV��VLPLODU�WR�WKH�VWHP��)LJ�������FRPSRVHG�RI�SKHORGHUP�IRUPHG�E\���WR���OD\HUV�RI�UHFWDQJXODU�FHOOV�LQ�FURVV�VHFWLRQ��ZLWK�SULPDU\�ZDOOV�� FRUN� ZDV� IRUPHG� E\� VHYHUDO� OD\HUV� RI� FHOOV� ZLWK�VXEHUL]HG�ZDOOV��ZKLFK�DOWHUQDWH�ZLWK�OD\HUV�RI�FHOOV�ZLWK�OLJQL¿HG�ZDOOV�

&RUWH[� ZDV� FRPSRVHG� RI� SDUHQFK\PD� FHOOV� ZLWK�primary walls. Druses were OHVV�IUHTXHQW�LQ�E. calochlora,

DQG�DEVHQW�LQ�E. rhodotricha .

7KH� SULPDU\� SKORHP� ZDV� FROODSVHG� �)LJV�� ���� �����with caps of sclereids outside secondary phloem (Fig.

����� 6HFRQGDU\� SKORHP� �)LJ�� ���� FRPSULVHG� QDUURZ�DQG� VKRUW� VLHYH� WXEH� HOHPHQWV��ZLWK� VLPSOH� VLHYH� SODWH��WUDQVYHUVDO� WR� VOLJKWO\� REOLTXH�� FRPSDQLRQ� FHOOV�� D[LDO�DQG�UDGLDO�SDUHQFK\PD��6HFRQGDU\�[\OHP�ZDV�¿EURXV�LQ�ERWK�VSHFLHV��)LJV����������GLIIXVH�SRURXV��FRPSRVHG�RI�vessels predominantly solitary and multiples of 2 to 3,

DQG�OHVV�IUHTXHQW��RI���WR���YHVVHO�HOHPHQWV�ZLWK�VLPSOH�SHUIRUDWLRQ�SODWH��WUDQVYHUVH�WR�REOLTXH��LQ�E. rhodotricha,

LQWHUPHGLDWH� IRUPV� EHWZHHQ� VFDODULIRUP� ZLWK� RQH�LQFRPSOHWH�EDU�WR�LUUHJXODU�DQG�UHWLFXODWH�ZHUH�REVHUYHG��VLPLODU� WR� WKH� VWHP� ¿EURXV�ZRRG�� LQWHUYHVVHO� SLWV� ZHUH�ERUGHUHG�� YHVVHO�SDUHQFK\PD� SLWV� ZHUH� VLPSOH� WR� KDOI�ERUGHUHG�� VFDODULIRUP�� RSSRVLWH� WR� DOWHUQDWH��:LGH� EDQG�WUDFKHLGV�ZHUH� DEVHQW�� 6FDQW\� SDUDWUDFKHDO� SDUHQFK\PD�SUHVHQW� �)LJ�� ����� 8QOLJQL¿HG�� PXOWLVHULDWH� ZLGH� DQG�KLJK� KHWHURJHQHRXV� UD\V� �)LJV�� ������� ZHUH� FRPSRVHG�RI� SURFXPEHQW�� XSULJKW� RU�DQG� VTXDUH� FHOOV��'UXVHV� DQG�VWDUFK� JUDLQV� �)LJ�� ���� RFFXUUHG� LQ� UD\� FHOOV�� /LEULIRUP�¿EUHV�ZHUH�SUHVHQW��VHSWDWH��UDUH��RQO\�LQ�E. rhodotricha

�)LJ�������8QOLJQL¿HG�SDUHQFK\PD�ZDV�SUHVHQW�LQFOXGLQJ�DOVR� D[LDO� SDUHQFK\PD�� IRUPLQJ�EDQGV��PRUH� HYLGHQW� LQ�E. rhodotricha.

Discussion

7KH�DQDWRPLFDO�FKDUDFWHULVWLFV�REVHUYHG�LQ�WKH�VWXGLHG�VSHFLHV�DUH�FRPPRQO\�REVHUYHG�LQ�RWKHU�&DFWDFHDH�VSHFLHV��DV�H[WHQVLYHO\�GHVFULEHG�LQ�WKH�OLWHUDWXUH��7HUUD]DV�6DOJDGR�DQG�0DXVHWK��������6RI¿DWWL�DQG�$QJ\DORVV\�������������������� 7HUUD]DV� DQG� $ULDV�� ������ $UUXGD� HW� DO��� ������0DXVHWK���������

The 2 Echinopsis� VSHFLHV� KDYH� D� WKLFN� FXWLFOH�� DQ�LPSRUWDQW� DGDSWDWLRQ� IRU� [HULF� KDELWDWV�� SURWHFWLQJ� WKH�SODQW� ERG\� IURP� ZDWHU� ORVV� DQG� SDWKRJHQV�� 8QXVXDO� LQ�&DFWDFHDH� �*LEVRQ� DQG� 1REHO�� ������ /R]D�&RUQHMR� DQG�7HUUD]DV�� ������� WKH\� VKRZHG� D� VHFRQGDULO\� ELVHULDWHG�epidermis, ZKLFK�KDV� DOVR�EHHQ� UHSRUWHG� IRU� D� IHZ�RWKHU�genera in Trichocereeae, such as Harrisia (Mauseth et

DO��� ������� D� PXOWLVHULDWH� HSLGHUPLV� ZDV� PHQWLRQHG� IRU�Espostoa� %ULWWRQ� DQG� 5RVH� �0DXVHWK�� ������� 5HSRUWV�RI� PXOWLVHULDWHG� HSLGHUPLV� H[LVW� IRU� RWKHU� WULEHV� VXFK�DV� 3DFK\FHUHHDH� �*LEVRQ� DQG� +RUDN�� ������ %DUWKORWW�DQG� +XQW�� ������ 7HUUD]DV�6DOJDGR� DQG� 0DXVHWK��������� &DFWHDH� �7HUUD]DV�6DOJDGR� DQG� 0DXVHWK�� �������1RWRFDFWHDH��1\IIHOHU�DQG�(JJOL��������7HUUD]DV�6DOJDGR�DQG� 0DXVHWK�� ������� &HUHHDH� �'DUOLQJ�� ������ 0DXVHWK��������� DQG�%URZQLQJLHDH� �0DXVHWK�� �������$V� VWDWHG� E\�/R]D�&RUQHMR�DQG�7HUUD]DV���������PXOWLSOH�HSLGHUPLV�LQ�cacti occur in unrelated genera and seem to have appeared

several times in Cactaceae.

3DUDOOHORF\WLF�VWRPDWD�DQG�D�ORQJ�VXEVWRPDWDO�FKDPEHU�ZHUH� REVHUYHG� LQ� WKH� LQYHVWLJDWHG� VSHFLHV�� D� FRPPRQ�IHDWXUH�IRU�WKH�PHPEHUV�RI�WKH�IDPLO\��*DVVRQ��������(JJOL���������,Q�FRQWUDVW�ZLWK�PRVW�[HURSK\WHV��VHYHUDO�VSHFLHV�RI�&DFWDFHDH�VKRZ�VXSHU¿FLDO�VWRPDWD��(JJOL��������*LEVRQ�DQG�+RUDN��������)DKQ�DQG�&XWOHU��������/R]D�&RQHMR�DQG�7HUUD]DV�� ������ 6RI¿DWWL� DQG� $QJ\DORVV\�� ������� ZKHUH�guard cells are on the same level as the other epidermal

FHOOV�� 7KH� ORQJ� VXEVWRPDWDO� FKDPEHU�� GXH� WR� WKH� WKLFN�hypodermis, reduces water loss when the stomata are open

�)DKQ�DQG�&XWOHU���������$FFRUGLQJ�WR�'DUOLQJ���������WKH�UHGXFHG� WUDQVSLUDWLRQ� UDWHV� DUH� FRPELQHG� ZLWK� ORZ� &2

2

and O2� H[FKDQJH� UDWHV�� SULRULWL]LQJ� ZDWHU� FRQVHUYDWLRQ��

6XQNHQ� VWRPDWD� ZHUH� REVHUYHG� LQ� E. rhodotricha, a

situation reported for some other genera (Metcalfe and

&KDON�� ������ *DVVRQ�� ������ (JJOL�� ������ /R]D�&RUQHMR�DQG� 7HUUD]DV�� ������� PD[LPL]LQJ� ZDWHU� ORVV� SUHYHQWLRQ��1HYHUWKHOHVV��WKLV�LV�D�YDULDEOH�IHDWXUH�LQ�WKH�JHQXV��RQFH�VXSHU¿FLDO�VWRPDWD�ZHUH�REVHUYHG�LQ�RWKHU�VSHFLHV��VXFK�DV�in Echinopsis aurea�%ULWWRQ�HW�5RVH��(JJOL�������, and in

E. calochlora, while in E. eyriesii (Turpin) Zucc. these are

VXQNHQ��DV�LQ�E. rhodotricha.

7KH� RUJDQL]DWLRQ� RI� FRUWLFDO� WLVVXHV� LQ� ERWK� VWXGLHG�Echinopsis�VSHFLHV�LV�W\SLFDO�RI�FDFWL��*LEVRQ�DQG�1REHO�������� /R]D�&RUQHMR� DQG� 7HUUD]DV�� ������ 0DXVHWK��

����Revista Mexicana de Biodiversidad 83: 1036-1044, 2012

DOI: 10.7550/rmb.28124

Figures 13-20.��������VWHP�VHFRQGDU\�[\OHP��WDQJHQWLDO�VHFWLRQV��Echinopsis rhodotricha������ODUJH�UD\V������VLPSOH�SHUIRUDWLRQ�SODWH�RI�D�YHVVHO�HOHPHQW������LQFRPSOHWH�SHUIRUDWLRQ�SODWH�RI�D�YHVVHO�HOHPHQW���������URRW�DQG�FURVV�VHFWLRQV��������������WDQJHQWLDO�VHFWLRQ���������E. calochlora������JHQHUDO�YLHZ��QRWH�SHULGHUPLV�FRPSRVHG�RI�VHYHUDO�OD\HUV�RI�FHOOV�ZLWK�OLJQL¿HG�ZDOOV�DOWHUQDWH�ZLWK�FHOOV�ZLWK�VXEHUL]HG�ZDOOV��¿EURXV�ZRRG�ZLWK�ODUJH�XQOLJQL¿HG�UD\V������SKORHP�DQG�[\OHP��&ROODSVHG�SKORHP��DUURZV��RXWVLGH�IXQFWLRQDO�VHFRQGDU\�SKORHP���������E. rhodotricha.�����¿EURXV�ZRRG�ZLWK�ODUJH�XQOLJQL¿HG�UD\V������GHWDLO�RI�XQOLJQL¿HG�UD\V�FRQWDLQLQJ�VWDUFK�JUDLQV������E. calochlora�WDQJHQWLDO�VHFWLRQV��VHSWDWH�¿EUHV��DUURZV���6FDOH�EDUV���� ������P������� �����P�������� ������P���� ������P������� ������P��&R�FRUWH[��3��SHULGHUPLV��3K��SKORHP��5��UD\V��;\��[\OHP

���� Garcia et al.- Anatomy of two species of Echinopsis

������ 6RI¿DWWL� DQG� $QJ\DORVV\�� ������� WKHUH� LV� D� WKLFN�hypodermis composed of several layers of colenchymatic

FHOOV�� IROORZHG�E\�D�SKRWRV\QWKHWLF�SDOLVDGH�SDUHQFK\PD�DQG� WKH� LQQHU� FRUWH[�� FRPSRVHG� RI� LVRGLDPHWULF�SDUHQFK\PDWLF� FHOOV�� 7KH� K\SRGHUPLV�� EHLQJ� D� ÀH[LEOH�tissue, has important functions in giving support to the

stem and accommodating the stem changes in volume due

WR�VKULQNDJH�DQG�VZHOOLQJ��DOVR�SURWHFWLQJ�WKH�LQQHU�WLVVXHV�DJDLQVW�SDWKRJHQV��*LEVRQ�DQG�1REHO���������$V�WKH�VWHPV�in most cacti are the main photosynthesizing organ, the

organization of the chlorenchyma or palisade parenchyma

LV�YHU\�VLPLODU�WR�D�OHDI��6DMHYD�DQG�0DXVHWK���������7KH�LQWHUQDO�FRUWH[�FHOOV�VWRUH�ZDWHU��0DXVHWK������E���DQG�WKH�ODUJH�FHOOV�DUH�DEOH� WR�H[SDQG�DQG�FRQWUDFW�GHSHQGLQJ�RQ�K\GUDWLRQ�VWDWH�RU�ZDWHU�DYDLODELOLW\��0DXVHWK��������

%RWK�VSHFLHV�SUHVHQWHG�FRUWLFDO�DQG�PHGXOODU\�EXQGOHV�with secondary growth, spread out in all directions very

VLPLODU� WR� OHDI� YHLQV�� /LNH� OHDI� YHLQV�� WKH\� DUH� FROODWHUDO�DQG� IRUP� D� QHWZRUN� WKDW� H[WHQGV� WR� WKH� EDVH� RI� WKH�SDOLVDGH� SDUHQFK\PD�� &RUWLFDO� EXQGOHV� DUH� FRQVLGHUHG� D�synapomorphy for Cactoideae, present in nearly all species

�0DXVHWK�� ������� 7KH\� SHUIRUP� LPSRUWDQW� IXQFWLRQV�LQ� VXJDU� DQG� ZDWHU� WUDQVSRUW� WR� DQG� IURP� FRUWH[� FHOOV�DQG� WR� WKH� YDVFXODU� V\VWHP� �0DXVHWK� DQG� 6DMHYD�� ������7HUUD]DV�6DOJDGR� DQG� 0DXVHWK�� ������ 0DXVHWK�� �������0DXVHWK� ������� FRQVLGHUV� WKDW� WKH� SUHVHQFH� RI� FRUWLFDO�EXQGOHV�H[SODLQV�WKH�ODUJH�YROXPH�RI�FRUWH[��DV�VXFFXOHQW�HXSKRUELDV�� IRU� LQVWDQFH�� GR�QRW�SRVVHVV� FRUWLFDO� EXQGOHV�and their stems never reach such a large size as cacti.

0HGXOODU\� EXQGOHV� KDYH� WKH� VDPH� IXQFWLRQ� DV� FRUWLFDO�EXQGOHV��WUDQVSRUWLQJ�ZDWHU�DQG�QXWULHQWV��0DXVHWK������D�������� 7HUUD]DV�6DOJDGR� DQG� 0DXVHWK�� ������� RFFXUULQJ�RQO\�LQ�&DFWRLGHDH��0DXVHWK������D���������$FFRUGLQJ�WR�*LEVRQ�DQG�1REHO��������SLWK�GLDPHWHU� LQFUHDVHG�GXULQJ�&DFWDFHDH� HYROXWLRQ� DQG� WKLV� HYHQW� SUREDEO\� OHG� WR� WKH�HPHUJHQFH�RI�PHGXOODU\�EXQGOHV�

,Q�WKH�EDVDO�DQG�ROGHU�UHJLRQV�RI�WKH�VWHP��DV�ZHOO�DV�in the root of these 2 Echinopsis species, the epidermis

LV�VXEVWLWXWHG�E\�D�SHULGHUPLV��DV�QRWHG�IRU�PDQ\�VSHFLHV�in the family. The origin of phellogen from periclinal

divisions of epidermal cells is very common in cacti

�*LEVRQ�DQG�1REHO��������7HUUD]DV�6DOJDGR�DQG�0DXVHWK�������� 6RI¿DWWL� DQG� $QJ\DORVV\�� ������ 0DXVHWK�� �������$OWKRXJK�WKH�RULJLQ�RI�SKHOORJHQ�LQ�D�SODQW�RUJDQ�FDQ�EH�YDULDEOH�� IURP� GLIIHUHQW� FHOO� W\SHV�� VXFK� DV� HSLGHUPDO��colenchymatic, parenchymatic or even phloematic cells,

WKH� PRVW� FRPPRQ� RULJLQ� LV� IURP� VXEHSLGHUPDO� OD\HUV�RI� FHOOV�� DFFRUGLQJ� WR� (YHUW� �������� ,W� ZDV� VHHQ� LQ� E.

calochlora and E. rhodotricha� WKDW� SKHOOHP� LQ� ERWK� URRW�DQG�VWHP�LV�FRPSRVHG�E\�OD\HUV�RI�VXEHUL]HG�FHOOV�ZKLFK�DOWHUQDWH� ZLWK� OD\HUV� RI� OLJQL¿HG� FHOOV�� DOVR� D� FRPPRQ�IHDWXUH�IRU�VSHFLHV�RI�WKH�IDPLO\��0DXVHWK��������

7KH� SKORHP� ZDV� FRPSRVHG� RI� VLHYH� WXEH� HOHPHQWV�DQG� FRPSDQLRQ� FHOOV�� D[LDO� DQG� UDGLDO� SDUHQFK\PD� DQG�VRPHWLPHV��QRQ�IXQFWLRQDO�SKORHP�PLJKW�EHFRPH�OLJQL¿HG��VLPLODU�WR�ZKDW�ZDV�GHVFULEHG�IRU�VHYHUDO�&DFWDFHDH�VWHPV��0DXVHWK�� ������ 7HUUD]DV�6DOJDGR� DQG� 0DXVHWK�� ������6RI¿DWWL� DQG� $QJ\DORVV\�� ������ $UUXGD� HW� DO��� ������0DXVHWK�� ������� DQG� URRWV� �0DXVHWK� DQG� 5RVV�� ������0DXVHWK�� ������� Echinopsis rhodotricha KDV� ¿EUH� FDSV�RXWVLGH�SKORHP��LQ�E. calochlora� WKH\�DUH�DEVHQW�� W\SLFDO�RI�VSHFLHV�ZKLFK�KDYH�QRQ�¿EURXV�ZRRG��0DXVHWK�HW�DO���������

Echinopsis calochlora and E. rhodotricha present

¿EURXV� ZRRG� LQ� WKH� URRWV� ZKLOH� LQ� WKH� VWHPV� WKH\� ERWK�KDYH�QRQ�¿EURXV�ZRRG��FRUURERUDWLQJ�WKH�IDFW�HVWDEOLVKHG�E\�0DXVHWK� DQG� 6WRQH�3DOPTXLVW� �������� 7KHVH� DXWKRUV�stated that the structure of wood in the stem and root of the

VDPH�SODQW�FDQ�EH�YHU\�GLIIHUHQW��Echinopsis rhodotricha

KDV�DOVR�¿EURXV�ZRRG�LQ�WKH�VWHP��FKDUDFWHUL]LQJ�ZKDW�LV�termed dimorphic wood: when young the plant produces

RQH� W\SH� RI� ZRRG� DQG� ODWHU� RQ�� WKH� FDPELXP� SURGXFHV�DQRWKHU� W\SH� �0DXVHWK�DQG�3OHPRQV��������0DXVHWK�DQG�3OHPRQV�5RGUtJXH]�� ������� ,Q� WKH� SUHVHQW� ZRUN�� LW� ZDV�REVHUYHG�D�W\SLFDO�VLWXDWLRQ�LQ�E. rhodotricha��ZKLFK�¿UVW�IRUPV� QRQ� ¿EURXV� ZRRG�� ZKHQ� WKH� ERG\� LV� \RXQJ� DQG�small and does not need special investment in support,

DQG� ZKHQ� PDWXUH�� WKH� FDPELXP� VWDUWV� IRUPLQJ� ¿EURXV�wood since support is then needed due to the larger size

�7HUUD]DV�6DOJDGR� DQG�0DXVHWK�� ������� 7KH� ¿EURXV� DQG�QRQ�¿EURXV�ZRRG�W\SHV�REVHUYHG�LQ�WKH�SUHVHQW�VWXG\�KDYH�WKH�VDPH�VWUXFWXUH�DV�GHVFULEHG�LQ�OLWHUDWXUH��ZLWK�¿EURXV�ZRRG�FRPSRVHG�RI�YHVVHO�HOHPHQWV�LQ�D�PDWUL[�RI�OLEULIRUP�¿EUHV��DQG�QRQ�¿EURXV�ZRRG�FRPSRVHG�RI�YHVVHO�HOHPHQWV�LQ�D�PDWUL[�RI�ZLGH�EDQG�WUDFKHLGV��*LEVRQ��������*LEVRQ�DQG� 1REHO�� ������ 7HUUD]DV�6DOJDGR� DQG�0DXVHWK�� ������0DXVHWK��������6RI¿DWWL�DQG�$QJ\DORVV\���������7KH�RQO\�GLIIHUHQFH�EHWZHHQ�WKH�¿EURXV�ZRRG�RI�WKH�VWHP�DQG�URRW�of E. rhodotricha is the presence of the storied structure in

WKH�VWHP��DEVHQW�LQ�WKH�URRW��$�QRWLFHDEOH�IHDWXUH�REVHUYHG�LQ�WKH�VHFRQGDU\�[\OHP�RI�WKH�VWHP�LQ�ERWK�VSHFLHV�LV�WKH�RFFXUUHQFH�RI�D�VWRULHG�VWUXFWXUH�LQ�WKH�VHFRQGDU\�[\OHP��7KLV�IHDWXUH�KDV�EHHQ�GHVFULEHG�LQ�WKH�OLWHUDWXUH�IRU�VHYHUDO�species of Cactoideae, including E. calochlora��E\�*LEVRQ���������7KLV�DXWKRU�REVHUYHG�WKDW�WKH�UD\V�DUH�UDUHO\�VWRULHG��and mentions that the storied structure is more evident in

VSHFLDOL]HG�JURZWK� IRUPV�� VXFK�DV�JORERVH�DQG�HSLSK\WLF�VSHFLHV��*LEVRQ��������DOVR�VWDWHG�WKDW�LQ�QRQ�¿EURXV�ZRRG�LW�LV�D�GLI¿FXOW�IHDWXUH�WR�GLVWLQJXLVK�GXH�WR�GLIIHUHQFHV�LQ�WKH�OHQJWK�RI�YHVVHO�HOHPHQWV�DQG�ZLGH�EDQG�WUDFKHLGV��EXW�this was not the case in the present study. Storied wood is

a feature that occurs in several phylogenetically unrelated

JURXSV�DQG�LW� LV�IRXQG�LQ�JURXSV�ZKHUH�IXVLIRUP�FDPELDO�LQLWLDOV�DUH�VKRUWHU��&DUOTXLVW���������,Q�WKH�SUHVHQW�VWXG\��

����Revista Mexicana de Biodiversidad 83: 1036-1044, 2012

DOI: 10.7550/rmb.28124

WKH�QRQ�¿EURXV�ZRRG�VKRZHG�D�VWRULHG�VWUXFWXUH��HVSHFLDOO\�UHJDUGLQJ� ZLGH� EDQG� WUDFKHLGV�� PRUH� SURQRXQFHG� LQ� E.

calochlora than E. rhodotricha.

0DQ\� RI� WKH� FKDUDFWHULVWLFV� SUHVHQWHG� LQ� WKLV� ZRUN�FRUURERUDWH�WKH�GHVFULSWLRQV�WKDW�KDG�DOUHDG\�EHHQ�UHSRUWHG�for other species of Cactoideae. Some anatomical features

FDQ�EH�XVHG�DV�GLDJQRVWLF��DOORZLQJ�IRU�WKH�VHJUHJDWLRQ�RI�the 2 species of Echinopsis studied, such as: presence of

ELVHULDWHG�HSLGHUPLV��VXQNHQ�VWRPDWD��DQG�¿EURXV�DQG�QRQ�¿EURXV�ZRRG�LQ�WKH�VWHP�RI�E. rhodotricha��$V�Echinopsis

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Acknowledgements

7KDQNV�WR�WKH�³)XQGDomR�GH�$SRLR�DR�'HVHQYROYLPHQWR�do Ensino, Ciência e Tecnologia do Estado de Mato Grosso

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University of Paraná, especially to the Dr. Cleusa Bona

GXH�WR�WKH�HTXLSPHQW�XVH��WR�'U��*HUDOGR�$OYHV�'DPDVFHQR�Jr. and the Msc. Vali Joana Pott for identifying the species

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Literature cited

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cactus (Cereus giganteus): anatomy and spectral properties.

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