Garcia et al 2012
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)
AnatomÃa de la raÃz 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.
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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.
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Resumen. Este estudio está enfocado a caracterizar y comparar la anatomÃa de tallos y raÃces 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�
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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�
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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
FRPSULVHV� D� ODUJH� QXPEHU� RI� VSHFLHV� �FD�� ���� VSHFLHV��$QGHUVRQ���������VRPH�LQWUDJHQHULF�YDULDELOLW\�LV�H[SHFWHG�IRU� WKLV� JHQXV�� DQG�PRUH� VSHFLHV� VKRXOG� EH� LQYHVWLJDWHG��LQ� RUGHU� WR� HVWDEOLVK� WKH� GLDJQRVWLF� YDOXH� RI� WKRVH� WUDLWV�mentioned.
Acknowledgements
7KDQNV�WR�WKH�³)XQGDomR�GH�$SRLR�DR�'HVHQYROYLPHQWR�do Ensino, Ciência e Tecnologia do Estado de Mato Grosso
GR� 6XO´� �)81'(&7��� IRU� WKH� PDVWHU� VFKRODUVKLS� WR� WKH�¿UVW�DXWKRU��WR�WKH�5HVHDUFK�&RRUGLQDWRU�±�3UR�5HLWRULD�GH�3HVTXLVD�H�3yV�*UDGXDomR�GD�8QLYHUVLGDGH�)HGHUDO�0DWR�*URVVR�GR�6XO��IURP�WKH�3URJUDPD�GH�3yV�*UDGXDomR�HP�%LRORJLD�9HJHWDO�� DQG� WKH�1DWLRQDO� &RXQFLO� RI� 5HVHDUFK��&13T���SURFHVV�Q�����������������:H�ZRXOG�OLNH�WR�WKDQN�the Scanning Electronic Microscope Center of the Federal
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
DQG�WR�WKH�³3DQWDQDO�6HHGV�1HW´�E\�WKH�ORJLVWLF�VXSSRUW�RQ�WKH�¿HOG�VDPSOLQJ�
Literature cited
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%DUWKORWW��:��DQG�'��5��+XQW��������&DFWDFHDH��In The families
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&DUOTXLVW�� 6�� ������ &RPSDUDWLYH� ZRRG� DQDWRP\�� 6\VWHPDWLF��ecological, and evolutionary aspects of dicotyledon wood,
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cactus (Cereus giganteus): anatomy and spectral properties.
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