morphology, ploidy and molecular phylogenetics reveal a new … new gps.pdf · 2013-01-17 · 1242...

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1240 TAXON 'HFHPEHU ± 3HWWLJUHZ DO Adansonia kilima sp. nov., a diploid baobab from Africa 1240 9HUVLRQ RI 5HFRUG LGHQWLFDO WR SULQW YHUVLRQ INTRODUCTION 7KH $IULFDQ EDREDE WUHH Adansonia digitata / VRPH- WLPHV NQRZQ DV WKH 7UHH RI /LIH KDV P\ULDG XVHV LQ PDQ\ GLI- IHUHQW FXOWXUHV VXSSO\LQJ D ZLGH YDULHW\ RI IRRGVWXIIV DV ZHOO DV ZDWHU ILEHUV IRU ZHDYLQJ DQG URSHPDNLQJ YHVVHOV DUWLVWLF PDWHULDOV HWF :LFNHQV *HEDXHU DO 6LGLEH :LOOLDPV :LFNHQV /RZH 'H &DOXZp DO %DREDEV DOVR VXSSO\ UDZ PDWHULDOV IRU KXQWLQJ DQG ILVK- LQJ HTXLSPHQW IRU PDNLQJ PXVLFDO LQVWUXPHQWV DQG DV D VRXUFH RI PDQ\ KHUEDO UHPHGLHV +ROORZ EDREDEV DUH XVHG IRU VKHOWHU DQG WR VWRUH ZDWHU DOVR DV SULVRQV EXULDO VLWHV VWDEOHV VWRUDJH URRPV ZDWFKWRZHUV HWF 9LUWXDOO\ DOO SDUWV RI WKH SODQW FDQ EH XVHG DQG WKH WUHH LV LQGHOLEO\ LPSODQWHG LQ WKH IRON WUDGLWLRQV DQG P\WKRORJ\ RI PDQ\ FXOWXUHV 7KHLU LFRQLF VKDSH H[WUDRUGL- QDU\ ORQJHYLW\ LQGLYLGXDO WUHHV FDQ OLYH IRU ZHOO RYHU WKRXVDQG \HDUV $GDQVRQ 6ZDUW 3DWUXW DO HFR- QRPLF LPSRUWDQFH HWKQRERWDQLFDO VLJQLILFDQFH DQG LQWULJXLQJ QDWXUDO KLVWRU\ KDYH FDSWXUHG SRSXODU LPDJLQDWLRQ IRU PDQ\ FHQWXULHV 7KH\ ZHUH NQRZQ WR WKH $QFLHQW (J\SWLDQV DQG KDYH EHHQ PHQWLRQHG LQ WUDYHO DFFRXQWV ZLWK UHODWLYH IUHTXHQF\ VLQFH WKH V :LFNHQV %DXP %OHQFK 0LFKHO $GDQVRQ GHVFULEHG WKH EDREDE LQ $GDQVRQ DQG /LQQHDXV PHPRULDOL]HG KLP DV WKH WUHH¶V QDPHVDNH /LQQHDXV 'DUZLQ GRFXPHQWHG EDREDE WUHHV RQ 6W -DJR LQ WKH &DSH 9HUGH ,VODQGV LQ FRPPHQWLQJ RQ WKHLU VL]H DQG ORQJHYLW\ $UPVWURQJ /LYLQJVWRQH PHQWLRQV EDREDE WUHHV VHYHUDO WLPHV GXULQJ KLV DFFRXQW RI KLV WUDYHOV LQ VRXWKHUQ $IULFD GXULQJ WKH V DQG V /LYLQJVWRQH 7KH JHQXV Adansonia FRQVLVWV RI HLJKW NQRZQ VSHFLHV A. digitata ZKLFK LV QDWLYH WR PDLQODQG $IULFD EXW ZKLFK KDV EHHQ ZLGHO\ GLVWULEXWHG WKURXJKRXW WKH WURSLFV E\ KXPDQV Morphology, ploidy and molecular phylogenetics reveal a new diploid species from Africa in the baobab genus Adansonia (Malvaceae: Bombacoideae) Jack D. Pettigrew FRS, Karen L. Bell, Adhil Bhagwandin, Eunice Grinan, Ngalla Jillani, Jean Meyer, Emily Wabuyele & Claudia E. Vickers 1 Queensland Brain Institute, The University of Queensland, St Lucia 4072, Australia 2 Royal Botanic Gardens Melbourne, Birdwood Avenue, South Yarra, Victoria 3141, Australia School of Geography and Environmental Sciences, Monash University, Clayton, Victoria 3800, Australia 4 School of Anatomical Sciences, Faculty of Health Science, University of the Witwatersrand, Johannesburg, Gauteng, 2193, South Africa Centre for Microscopy & Microanalysis, The University of Queensland, St Lucia 4072 Australia 6 Pretoria National Herbarium, Cussonia Ave, Bremmeria 0001, South Africa East African Herbarium, Museum Hill Road, P.O Box 45166, 00100 Nairobi, Kenya Australian Institute for Bioengineering and Nanotechnology, The University of Queensland, St Lucia 4072 Australia $XWKRU IRU FRUUHVSRQGHQFH Jack D. Pettigrew, [email protected] Abstract 7KH JHQXV Adansonia KDV D GLVMXQFW JHRJUDSKLFDO GLVWULEXWLRQ VL[ VSHFLHV DUH HQGHPLF LQ 0DGDJDVFDU RQH LQ $IULFD DQG RQH LQ $XVWUDOLD 7KH ZHOONQRZQ $IULFDQ EDREDE Adansonia digitata LV DQ LFRQLF WUHH ZLWK FRQVLGHUDEOH HWKQRERWDQLFDO VLJQLILFDQFH ,Q FRQWUDVW WR WKH RWKHU VHYHQ VSHFLHV ZKLFK DUH GLSORLG A. digitata LV WHWUDSORLG $ FRPPRQ DQFHVWRU RI A. digitata DQG WKH RWKHU GLSORLG EDREDE VSHFLHV ZRXOG EH GLSORLG KRZHYHU WKHUH DUH QR GLSORLG VSHFLHV UHFRUGHG RQ WKH $IULFDQ PDLQODQG ([DPLQLQJ YDULDWLRQ LQ IORUDO DQG SROOHQ FKDUDFWHUV DQG FKURPRVRPH QXPEHU LQ VSHFLPHQV IURP $IULFD LGHQWLILHG D QHZ GLSORLG EDREDE VSHFLHV Adansonia kilima VS QRY ZKLFK FRH[LVWV ZLWK A. digitata LQ $IULFD Adansonia kilima LV UHVWULFWHG WR PRGHUDWH HOHYDWLRQV ± P LQ FRQWUDVW WR A. digitata ZKLFK LV ZLGHVSUHDG WKURXJKRXW $IULFD EXW SUHIHUV HOHYDWLRQV EHORZ P Adansonia kilima LV VXSHUILFLDOO\ VLPLODU WR A. digitata EXW FDQ EH GLIIHUHQWLDWHG RQ WKH EDVLV RI IORUDO PRUSKRORJ\ SROOHQ DQG FKURPRVRPH QXPEHU :H XVHG WZR FKORURSODVW '1$ PDUNHUV DQG WKH QXFOHDU ,76 WR H[DPLQH SK\ORJHQHWLF UHODWLRQVKLSV ZLWKLQ Adansonia 7KUHH OLQHDJHV ZHUH REVHUYHG RQH FRQWDLQLQJ WKH 0DODJDV\ VSHFLHV RQH FRQWDLQLQJ WKH $XVWUDOLDQ VSHFLHV DQG RQH FRQWDLQLQJ WKH $IULFDQ VSHFLHV 7KH UHODWLRQVKLSV EHWZHHQ WKHVH FODGHV ZHUH GLIILFXOW WR UHVROYH EXW D OLQN EHWZHHQ WKH $IULFDQ DQG $XVWUDOLDQ FODGHV HPHUJHG ZKHQ WKH DQDO\VLV XVHG IHZHU UHSOLFDWH VDPSOHV RI LQGLYLGXDO 0DODJDV\ WD[D LQFOXGHG LQGHO FKDUDFWHUV DQG LQFOXGHG IHZHU RXWJURXS WD[D 7KH ,76 SK\ORJHQ\ GHPRQVWUDWHG WKDW A. digitata DQG A. kilima DUH JHQHWL- FDOO\ VLPLODU VXJJHVWLQJ WKDW WHWUDSORLG\ HYROYHG UHODWLYHO\ UHFHQWO\ Keywords Adansonia kilima $IULFD EDREDE GLSORLG PROHFXODU SK\ORJHQ\ Supplementary Material )LJXUHV 6±6 DUH DYDLODEOH LQ WKH (OHFWURQLF 6XSSOHPHQW WR WKH RQOLQH YHUVLRQ RI WKLV DUWLFOH KWWS ZZZLQJHQWDFRQQHFWFRPFRQWHQWLDSWWD[

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Page 1: Morphology, ploidy and molecular phylogenetics reveal a new … new GPS.pdf · 2013-01-17 · 1242 3HWWLJUHZ DO Adansonia kilima sp. nov., a diploid baobab from Africa TAXON 'HFHPEHU

1240

TAXON����������'HFHPEHU�����������±����3HWWLJUHZ��DO����Adansonia kilima sp. nov., a diploid baobab from Africa

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INTRODUCTION

7KH�$IULFDQ�EDREDE�WUHH��Adansonia digitata /���VRPH-WLPHV�NQRZQ�DV�WKH�7UHH�RI�/LIH��KDV�P\ULDG�XVHV�LQ�PDQ\�GLI-IHUHQW�FXOWXUHV��VXSSO\LQJ�D�ZLGH�YDULHW\�RI�IRRGVWXIIV��DV�ZHOO�DV�ZDWHU��ILEHUV�IRU�ZHDYLQJ�DQG�URSH�PDNLQJ��YHVVHOV��DUWLVWLF�PDWHULDOV��HWF���:LFNHQV��������*HEDXHU��DO���������6LGLEH��:LOOLDPV��������:LFNHQV��/RZH��������'H�&DOXZp��DO����������%DREDEV�DOVR�VXSSO\�UDZ�PDWHULDOV�IRU�KXQWLQJ�DQG�ILVK-LQJ�HTXLSPHQW��IRU�PDNLQJ�PXVLFDO�LQVWUXPHQWV��DQG�DV�D�VRXUFH�RI�PDQ\�KHUEDO�UHPHGLHV��+ROORZ�EDREDEV�DUH�XVHG�IRU�VKHOWHU�DQG�WR�VWRUH�ZDWHU��DOVR�DV�SULVRQV��EXULDO�VLWHV��VWDEOHV��VWRUDJH�URRPV��ZDWFKWRZHUV��HWF��9LUWXDOO\�DOO�SDUWV�RI�WKH�SODQW�FDQ�EH�XVHG��DQG�WKH�WUHH�LV�LQGHOLEO\�LPSODQWHG�LQ�WKH�IRON�WUDGLWLRQV�DQG�P\WKRORJ\�RI�PDQ\�FXOWXUHV��7KHLU�LFRQLF�VKDSH��H[WUDRUGL-QDU\�ORQJHYLW\��LQGLYLGXDO�WUHHV�FDQ�OLYH�IRU�ZHOO�RYHU�WKRXVDQG�

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7KH� JHQXV�Adansonia� FRQVLVWV� RI� HLJKW� NQRZQ� VSHFLHV��A. digitata��ZKLFK�LV�QDWLYH�WR�PDLQODQG�$IULFD�EXW�ZKLFK�KDV�EHHQ�ZLGHO\� GLVWULEXWHG� WKURXJKRXW� WKH� WURSLFV� E\� KXPDQV��

Morphology, ploidy and molecular phylogenetics reveal a new diploid species from Africa in the baobab genus Adansonia (Malvaceae: Bombacoideae)Jack D. Pettigrew FRS,!"# Karen L. Bell,$"% Adhil Bhagwandin,& Eunice Grinan,' Ngalla Jillani,& Jean Meyer,( Emily Wabuyele) & Claudia E. Vickers#

1 Queensland Brain Institute, The University of Queensland, St Lucia 4072, Australia2 Royal Botanic Gardens Melbourne, Birdwood Avenue, South Yarra, Victoria 3141, Australia�� School of Geography and Environmental Sciences, Monash University, Clayton, Victoria 3800, Australia4 School of Anatomical Sciences, Faculty of Health Science, University of the Witwatersrand, Johannesburg,

Gauteng, 2193, South Africa�� Centre for Microscopy & Microanalysis, The University of Queensland, St Lucia 4072 Australia6 Pretoria National Herbarium, Cussonia Ave, Bremmeria 0001, South Africa�� East African Herbarium, Museum Hill Road, P.O Box 45166, 00100 Nairobi, Kenya�� Australian Institute for Bioengineering and Nanotechnology, The University of Queensland, St Lucia 4072 Australia$XWKRU�IRU�FRUUHVSRQGHQFH� Jack D. Pettigrew, [email protected]

Abstract 7KH�JHQXV�Adansonia KDV�D�GLVMXQFW�JHRJUDSKLFDO�GLVWULEXWLRQ��VL[�VSHFLHV�DUH�HQGHPLF�LQ�0DGDJDVFDU��RQH�LQ�$IULFD��DQG�RQH�LQ�$XVWUDOLD��7KH�ZHOO�NQRZQ�$IULFDQ�EDREDE��Adansonia digitata��LV�DQ�LFRQLF�WUHH�ZLWK�FRQVLGHUDEOH�HWKQRERWDQLFDO�VLJQLILFDQFH��,Q�FRQWUDVW�WR�WKH�RWKHU�VHYHQ�VSHFLHV��ZKLFK�DUH�GLSORLG��A. digitata�LV�WHWUDSORLG��$�FRPPRQ�DQFHVWRU�RI�A. digitata DQG�WKH�RWKHU�GLSORLG�EDREDE�VSHFLHV�ZRXOG�EH�GLSORLG��KRZHYHU��WKHUH�DUH�QR�GLSORLG�VSHFLHV�UHFRUGHG�RQ�WKH�$IULFDQ�PDLQODQG��([DPLQLQJ�YDULDWLRQ�LQ�IORUDO�DQG�SROOHQ�FKDUDFWHUV�DQG�FKURPRVRPH�QXPEHU�LQ�VSHFLPHQV�IURP�$IULFD�LGHQWLILHG�D�QHZ�GLSORLG�EDREDE�VSHFLHV��Adansonia kilima VS��QRY���ZKLFK�FR�H[LVWV�ZLWK�A. digitata�LQ�$IULFD��Adansonia kilima�LV�UHVWULFWHG�WR�PRGHUDWH�HOHYDWLRQV�����±�����P���LQ�FRQWUDVW�WR�A. digitata� ZKLFK�LV�ZLGHVSUHDG�WKURXJKRXW�$IULFD�EXW�SUHIHUV�HOHYDWLRQV�EHORZ�����P��Adansonia kilima LV�VXSHUILFLDOO\�VLPLODU�WR�A. digitata��EXW�FDQ�EH�GLIIHUHQWLDWHG�RQ�WKH�EDVLV�RI�IORUDO�PRUSKRORJ\��SROOHQ��DQG�FKURPRVRPH�QXPEHU��:H�XVHG�WZR�FKORURSODVW�'1$�PDUNHUV�DQG�WKH�QXFOHDU�,76�WR�H[DPLQH�SK\ORJHQHWLF�UHODWLRQVKLSV�ZLWKLQ�Adansonia��7KUHH�OLQHDJHV�ZHUH�REVHUYHG��RQH�FRQWDLQLQJ�WKH�0DODJDV\�VSHFLHV��RQH�FRQWDLQLQJ�WKH�$XVWUDOLDQ�VSHFLHV��DQG�RQH�FRQWDLQLQJ�WKH�$IULFDQ�VSHFLHV��7KH�UHODWLRQVKLSV�EHWZHHQ�WKHVH�FODGHV�ZHUH�GLIILFXOW�WR�UHVROYH��EXW�D�OLQN�EHWZHHQ�WKH�$IULFDQ�DQG�$XVWUDOLDQ�FODGHV�HPHUJHG�ZKHQ�WKH�DQDO\VLV�XVHG�IHZHU�UHSOLFDWH�VDPSOHV�RI�LQGLYLGXDO�0DODJDV\�WD[D��LQFOXGHG�LQGHO�FKDUDFWHUV�DQG�LQFOXGHG�IHZHU�RXWJURXS�WD[D��7KH�,76�SK\ORJHQ\�GHPRQVWUDWHG�WKDW�A. digitata�DQG�A. kilima�DUH�JHQHWL-FDOO\�VLPLODU��VXJJHVWLQJ�WKDW�WHWUDSORLG\�HYROYHG�UHODWLYHO\�UHFHQWO\�

Keywords Adansonia kilima� $IULFD��EDREDE��GLSORLG��PROHFXODU�SK\ORJHQ\

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Page 2: Morphology, ploidy and molecular phylogenetics reveal a new … new GPS.pdf · 2013-01-17 · 1242 3HWWLJUHZ DO Adansonia kilima sp. nov., a diploid baobab from Africa TAXON 'HFHPEHU

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MATERIALS AND METHODS

Sampling. — /HDYHV��IORZHUV��DQG�VHHGV�ZHUH�FROOHFWHG�IURP�EDREDE�WUHHV�LQ�D�QXPEHU�RI�ORFDWLRQV�LQ�$IULFD��7KH�VXUYH\�UHJLRQ�UDQJHG�IURP�WKH�HDVWHUQ�VORSHV�RI�0W��.LOLPDQMDUR��7VDYR�:HVW�1DWLRQDO�3DUN��LQ�.HQ\D�WR�WKH�VRXWK�.HQ\DQ�FRDVW��WKH�FHQWUDO�SODWHDX�RI�7DQ]DQLD�DQG�=DPELD��ZHVW�WR�%RWVZDQD�DQG�1DPLELD�DQG�VRXWK�WR�WKH�OLPLWV�RI�EDREDE�GLVWULEXWLRQ�LQ�WKH�/LPSRSR�UHJLRQ�RI�6RXWK�$IULFD��$�IHZ�VDPSOHV�IURP�6HQHJDO�LQ�:HVW�$IULFD�ZHUH�DOVR�SURYLGHG�E\�$ODLQ�&RDFKH��2&,6���/HDI�PDWHULDO�IURP�WKH�$XVWUDOLDQ�VSHFLHV�A. gregorii ZDV�FROOHFWHG�IURP�WUHHV�JURZLQJ�ZLOG�LQ�WKH�UHPRWH�.LPEHUOH\�UHJLRQ�RI�:HVW-HUQ�$XVWUDOLD��/HDI�PDWHULDO�IURP�0DODJDV\�VSHFLHV�ZDV�NLQGO\�SURYLGHG�E\�3DVFDO�'DQWKX��&,5$'��0DGDJDVFDU��DQG�&KULVWLDQ�.XOO�DQG�+DULSUL\D�5DQJDQ��0RQDVK�8QLYHUVLW\��$XVWUDOLD��

Hawk mothpollinated

Limitedinformation

Mammalpollinated

Mammalpollinated

Bombacoid outgroups

A. perrieri

A. madagascariensis

A. za

A. rubrostipa

A. suarezensis

A. grandidieri

A. gregorii

hypothetical diploid

A. digitata

Malagasy

Australian

African

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Page 3: Morphology, ploidy and molecular phylogenetics reveal a new … new GPS.pdf · 2013-01-17 · 1242 3HWWLJUHZ DO Adansonia kilima sp. nov., a diploid baobab from Africa TAXON 'HFHPEHU

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DNA sequencing. —�/HDYHV�ZHUH�GHK\GUDWHG�E\�DGGLQJ�WKHP�WR�DQ�H[FHVV����������YROXPH�RI�VLOLFD�JHO�JUDQXOHV�DQG�VWRU-LQJ�LQ�D�VHDOHG�EDJ��'1$�ZDV�LVRODWHG�XVLQJ�WKH�'1HDV\�3ODQW�.LW��4LDJHQ��'RQFDVWHU��9LFWRULD��$XVWUDOLD���ZLWK�WKH�IROORZLQJ�PRGLILFDWLRQV��DSSUR[LPDWHO\����PJ�RI�SRO\YLQ\OS\UUROLGRQH�ZDV�DGGHG�WR�HDFK�VDPSOH�SULRU�WR�JULQGLQJ�LQ�D�PRUWDU�DQG�SHVWOH��WKH�YROXPH�RI�EXIIHU�$3��ZDV�LQFUHDVHG�WR�����ȝO��WKH�YROXPH�RI�EXIIHU�$3��ZDV�LQFUHDVHG�WR�����ȝO��DIWHU�SUHFLSLWD-WLRQ�RQ�LFH�WKH�VDPSOHV�ZHUH�FHQWULIXJHG�DW��������g�DW���&�IRU����PLQ��'1$�ZDV�HOXWHG�IURP�WKH�VSLQ�FROXPQV�WKUHH�WLPHV��HDFK�ZLWK�����ȝO�RI�EXIIHU�$(�

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Morphological analyses. —�6WRPDWDO�GLPHQVLRQV�ZHUH�GHWHUPLQHG�E\�XVLQJ�D�VPDOO��SRUWDEOH�ILHOG�PLFURVFRSH��3\VHU�(YROXWLRQ��3\VHU�6*,�/WG��(GHQEULGJH��8�.����&OHDU�QDLO�YDUQLVK�OLIWV�RI�WKH�DGD[LDO�OHDI�VXUIDFH�ZHUH�H[DPLQHG�XQGHU�î����PDJ-QLILFDWLRQ��,PDJHV�ZHUH�FDSWXUHG�WKURXJK�WKH�H\HSLHFH�XVLQJ�DQ�L3KRQH��$SSOH��DQG�VWRPDWDO�DSHUWXUH�PHDVXUHG�ZLWK�FDOLSHUV�DW�WKH�PD[LPXP�PDJQLILFDWLRQ�RQ�WKH�L3KRQH�VFUHHQ��7ULDOV�RQ�GLSORLG��A. gregorii� A. madagascariensis��DQG�WHWUDSORLG�(A. digitata��EDREDEV�IURP�WKH�%ULVEDQH�%RWDQLFDO�*DUGHQV��0W��&RRW�WKD��%ULVEDQH��GHPRQVWUDWHG�WKH�XWLOLW\�RI�WKLV�PHWKRG�IRU�GLIIHUHQWLDWLQJ�EHWZHHQ�GLSORLG�DQG�WHWUDSORLG�VSHFLHV��0HDQ�VWRPDWDO�OHQJWK�ZDV�PHDVXUHG�DQG�VWRPDWDO�GHQVLW\�ZDV�FDO-FXODWHG�IRU�OHDYHV�VDPSOHG�LQ�$IULFD��7UHHV�ZHUH�DVVLJQHG�WR�HLWKHU�GLSORLG�A. kilima RU�WHWUDSORLG�A. digitata�EDVHG�RQ�VL]H�RI�VWRPDWDO�DSHUWXUH��3ROOHQ�PHDVXUHPHQWV�XVLQJ�WKH�SRUWDEOH�PLFURVFRSH�ZHUH�DYDLODEOH�LQ�PDQ\�FDVHV�DQG�ZHUH�XVHG�WR�VXS-SRUW�WKH�GLDJQRVLV�EHFDXVH�RI�WKH�PXFK�ODUJHU�SROOHQ�VL]H�RI�WKH�WHWUDSORLG��VHH�EHORZ���0HDVXUHPHQWV�RI�DQGURHFLXP�OHQJWK�DQG�VWDON�GLDPHWHU��PD[LPXP�FDO\[�GLDPHWHU��DQG�VWDPLQDO�FRUROOD�ZLGWK�ZHUH�WDNHQ�XVLQJ�D�UXOHU��7KH�QXPEHU�RI�IUHH�VWDPLQDO�ILODPHQWV�ZDV�FRXQWHG�

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Chromosome number. —�6HHGV�ZHUH�VRDNHG�LQ�FRQFHQ-WUDWHG�K\GURFKORULF�DFLG�IRU����KU�WR�EUHDN�GRUPDQF\�DQG�WKHQ�JHUPLQDWHG�LQ�FRPPHUFLDO�VHHG�UDLVLQJ�PL[��6HDUOHV��.LOFR\��$XVWUDOLD��XQWLO�WKH�FRW\OHGRQV�ZHUH�H[SDQGHG��6HHGOLQJV�ZHUH�WKHQ�UHPRYHG�IURP�WKH�VRLO��ULQVHG��DQG�SODFHG�LQ�WHVW�WXEHV�ILOOHG�ZLWK�DTXHRXV������FROFKLFLQH��7XEHV�ZHUH�ZUDSSHG�LQ�DOXPLQLXP�IRLO�DQG�LQFXEDWHG�DW���&�IRU����KU��5RRW�WLSV�DQG�

Page 4: Morphology, ploidy and molecular phylogenetics reveal a new … new GPS.pdf · 2013-01-17 · 1242 3HWWLJUHZ DO Adansonia kilima sp. nov., a diploid baobab from Africa TAXON 'HFHPEHU

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RESULTS

Morphological differentiation between the diploid and the tetraploid. —�:H�SHUIRUPHG�D�VXUYH\�RI�EDREDEV�IURP�VHYHUDO�GLIIHUHQW�ORFDWLRQV�LQ�$IULFD�DQG�ZHUH�DEOH�WR�LGHQWLI\�WZR�VWRPDWDO�FODVVHV��)LJ��6����7KH�FODVV�ZLWK�D�ODUJHU�VWRPD-WDO�DSHUWXUH�DQG�ORZHU�VWRPDWDO�GHQVLW\��������P�DQG�����SHU������Pð��UHVSHFWLYHO\��KDG�D�FKURPRVRPH�FRXQW�RI��n = 160 �7DEOH�����7KLV�LV�LQ�DJUHHPHQW�ZLWK�SUHYLRXV�ILQGLQJV�IRU�WHWUD-SORLG�A. digitata��%DXP��2JLQXPD���������1XPHURXV�WUHHV�ZHUH�IRXQG�ZLWK�D�VPDOOHU�VWRPDWDO�DSHUWXUH��PHDQ�OHQJWK� ����1��P���KLJKHU�VWRPDWDO�GHQVLW\����SHU������Pð��DQG�PXFK�

Table . 0RUSKRORJLFDO�PHDVXUHPHQWV�DQG�FKURPRVRPH�QXPEHUV�IRU�A. kilima�DQG�A. digitata��1XPEHUV�DUH�DYHUDJHV�RU�UDQJHV�IURP n = ��WUHHV�SHU�VSHFLHV��A. kilima VDPSOHV�IURP�WKH�&HQWUDO�+LJKODQGV�RI�7DQ]DQLD��ILYH�A. digitata�VDPSOHV�IURP�WKH�FRDVWDO�UHJLRQ�DURXQG�0RPEDVD��.HQ\D�DQG�RQH�IURP�%ULVEDQH�%RWDQLF�*DUGHQV��IRU�DOO�PHDVXUHPHQWV�H[FHSW�IUHH�VWDPLQDO�ILODPHQWV��n� �����:KHUH�DYHUDJHV�DUH�XVHG��HUURUV�DUH�JLYHQ�DV�VWDQGDUG�GHYLDWLRQ��6�'���

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A trnL-trnF B ITS

D ITSC trnL-trnF

Cavanillesia platanifolia HQ696737

A. gregorii HQ696740

A. madagascariensis Ama1 JN400307

A. za HQ696741

A. rubrostipa Aru1 JN400306

A. digitata HQ696378

A. kilima 402 JX178941

A. kilima 488 JX178942

A. grandidieri HQ696739

A. suarezensis DIEB005.i JN400304

2.0 substitutions

66

85

0.88

0.72

Cavanillesia platanifolia HQ696737

A. madagascariensis Ama1 JN400307

A. rubrostipa Aru1 JN400306

A. za HQ696741

A. gregorii HQ696740

A. kilima 402 JX178941

A. kilima 488 JX178942

A. sp. HQ696378

A. grandidieri HQ696739

A. suarezensis DIEB005.i JN400304

0.02 subs/site

Cavanillesia platanifolia HQ658371

A. gregorii AF028524

A. sp. AF028525

A. kilima 488 JX178940

A. kilima 402 JX178939

A. rubrostipa AF028531

A. digitata AF028526

A. digitata Adi2 JX254900

A. suarezensis AF028529

A. za AF028536

A. grandidieri AF028528

A. grandidieri AF028527

A. perrieri AF028538

A. madagascariensis AF028533

A. madagascariensis AF028534

20.0 substitutions

100

94

89

77

62

61

54 87

0.79

0.76

0.86

0.96

1

0.89

0.92

1

Cavanillesia platanifolia HQ658371

A. rubrostipa AF028531 A. gregorii AF028524

A. za AF028536

A. suarezensis AF028529

A. kilima 402 JX178939 A. kilima 488 JX178940 A. sp. AF028525

A. perrieri AF028538

A. grandidieri AF028527 A. grandidieri AF028528

A. digitata AF028526 A. digitata Adi2 JX254900

A. madagascariensis AF028533 A. madagascariensis AF028534

0.03 subs/site

1

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DQG�$)�������DUH�ODEHOHG�DV�Adansonia VS��LQ�)LJ���%�DQG�'��VLQFH�WKH\�ZHUH�FROOHFWHG�SULRU�WR�UHFRJQLWLRQ�RI�WKH�GLSORLG�VSHFLHV��$)�������ZDV�VDPSOHG�IURP�D�WUHH�LQ�%XUNLQD�)DVR��LWV�JHRJUDSKLF�ORFDWLRQ��KRW��FRDVWDO��LQGLFDWHV�WKDW�LW�LV�PRVW�OLNHO\�A. digitata��)XUWKHUPRUH��WKH�VHHG�EDWFK�ZKLFK�SURYLGHG�WKLV�'1$�VDPSOH�ZDV�DOVR�XVHG�IRU�FKURPRVRPH�FRXQWV�LQ�D�VXEVHTXHQW�VWXG\�E\�%DXP��2JLQXPD��������'��%DXP��SHUV��FRPP����WKH�FKURPRVRPH�QXPEHU�LQGLFDWHV�LW�LV�LQGHHG�WHWUD-SORLG��7KLV�VXSSRUWV�LWV�SRVLWLRQ�RQ�WKH�,76�WUHH��ZKLFK�SODFHV�LW�ZLWK�WKH�A. digitata�DFFHVVLRQV��$)�������ZDV�VDPSOHG�IURP�D�FXOWLYDWHG�WUHH�LQ�,RZD�6WDWH�8QLYHUVLW\��VHHG�VRXUFH�XQNQRZQ���LWV�SODFHPHQW�LQ�WKH�,76�SK\ORJHQ\�VXJJHVWV�WKDW�LW�LV�LQ�IDFW�A. kilima��7KH�DFFHVVLRQ�RI�$)�������LV�XQNQRZQ��DV�WKHUH�LV�QR�SXEOLFDWLRQ�DVVRFLDWHG�ZLWK�WKLV�*HQ%DQN�VHTXHQFH��,WV�SODFHPHQW�LQ�WKH�SK\ORJHQ\�VXJJHVWV�WKDW�LW� LV�A. digitata.�,I�ZH�FRQVLGHU�Adansonia VS� $)�������WR�EH�A. kilima��WKH�SK\-ORJHQHWLF�DQDO\VLV�VXJJHVWV�WKDW�WKH�WHWUDSORLG�LV�PRQRSK\OHWLF�ZLWKLQ�D�SDUDSK\OHWLF�GLSORLG��FRQVLVWHQW�ZLWK�WKH�WZR�JURXSV�EHLQJ�VHSDUDWH�VSHFLHV��2QO\�D�VPDOO�QXPEHU�RI�VXEVWLWXWLRQV�VHSDUDWH�WKHP��DSSUR[LPDWHO\�DQ�RUGHU�RI�PDJQLWXGH�OHVV�WKDQ�WKH�GLYHUJHQFH�EHWZHHQ�WKHVH�WZR�VSHFLHV�DQG�A. gregorii.

DISCUSSION

Ploidy level and morphology differentiate two African species. —�3RO\SORLGL]DWLRQ�LV�NQRZQ�WR�KDYH�VHYHUDO�GLUHFW�HIIHFWV�RQ�SODQW�PRUSKRORJ\��,QFUHDVHV�LQ�WKH�VL]H�RI�FHOOV�DQG�RUJDQV��H�J���IORZHUV��VWRPDWD��SROOHQ��VHHGV��UHODWLYH�WR�GLSORLGV�LV�FRPPRQ��6WHEELQV��������5DPVH\��6FKHPVNH���������,Q�SDUWLFXODU��LQFUHDVLQJ�SORLG\�OHYHO�RIWHQ�UHVXOWV�LQ�GHFUHDVHG�QXPEHUV�RI�VWRPDWD�DQG�LQFUHDVHG�JXDUG�FHOO�VL]H��(YDQV��������%LQJKDP��������0LVKUD��������$U\DYDQG��DO���������&KHQ��DO���������DQG�WKHVH�WUDLWV�FDQ�VRPHWLPHV�EH�XVHG�WR�LQIHU�SORLG\��:H�XVHG�WKLV�DSSURDFK�WR�VXFFHVVIXOO\�GLIIHUHQWLDWH�EH-WZHHQ�GLSORLG�A. kilima�DQG�WHWUDSORLG�A. digitata�LQ�WKH�ILHOG��ZLWK�WKH�GLDJQRVLV�FRQILUPHG�E\�FKURPRVRPH�FRXQWV��7DEOH�����)XUWKHU�DQDO\VLV�UHYHDOHG�VHYHUDO�PRUSKRORJLFDO�GLIIHUHQFHV��7DEOH����)LJV���±���)LJV��6���6���6����)LJXUH�6��VKRZV�WKH�GLV-FRYHUHG�ORFDWLRQV�RI�WKH�$IULFDQ�GLSORLG�ZLWKLQ�WKH�VDPSOHG�UHJLRQV�

Phylogenetic conclusions for Adansonia. —�3K\ORJHQLHV�LQIHUUHG�IURP�DOO�WKUHH�JHQH�UHJLRQV�LQ�RXU�VWXG\�ZHUH�FRQFRUG-DQW�LQ�VXJJHVWLQJ��D�FORVH�UHODWLRQVKLS�EHWZHHQ�A. kilima DQG�A. digitata��WKH�PRQRSK\O\�RI�A. gregorii��DQG�FORVH�UHODWLRQ-VKLSV�DPRQJ�WKH�0DODJDV\�VSHFLHV��,Q�DGGLWLRQ��WKH�H[LVWHQFH�RI�WKUHH�PDMRU�VXEFODGHV�ZLWKLQ�Adansonia��FRUUHVSRQGLQJ�WR�WKH�WKUHH�JHRJUDSKLFDO�DUHDV��$IULFD��$XVWUDOLD��0DGDJDVFDU���LV�VWURQJO\�VXSSRUWHG�E\�WKH�,76�SK\ORJHQ\��3UHYLRXV�VWXGLHV�EDVHG�RQ�D�FRPELQDWLRQ�RI�FKORURSODVW�DQG�QXFOHDU�'1$�DQG�PRUSKRORJLFDO�FKDUDFWHUV��%DXP��DO����������DQG�SK\ORJHQHWLF�DQDO\VLV�RI�%RPEDFRLGHDH�EDVHG�RQ�,76�VHTXHQFHV��'XDUWH��DO���������KDV�VXJJHVWHG��DOEHLW�ZHDNO\��WKDW�WKH�$XVWUDOLDQ�DQG�$IULFDQ�FODGHV�ZHUH�VLVWHUV��2XU�GDWD�DOVR�WHQG�WR�VXSSRUW�WKLV�FRQFOXVLRQ��EXW�DJDLQ�VXSSRUW�ZDV�ORZ��LQGLFDWLQJ�WKDW�DG-GLWLRQDO�GDWD�ZLOO�EH�QHHGHG�WR�UHVROYH�WKH�GHHS�UHODWLRQVKLSV�DPRQJ�WKH�PDMRU�OLQHDJHV�RI�Adansonia�

Page 8: Morphology, ploidy and molecular phylogenetics reveal a new … new GPS.pdf · 2013-01-17 · 1242 3HWWLJUHZ DO Adansonia kilima sp. nov., a diploid baobab from Africa TAXON 'HFHPEHU

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Adansonia digitata DQG�A. kilima�DUH�GLIIHUHQWLDWHG�E\�SORLG\�OHYHO��DQG�E\�PRUSKRORJLFDO�FKDUDFWHUV��SDUWLFXODUO\�LQ�WKH�IORZHUV�DQG�SROOHQ��2I�WKH�WKUHH�PDUNHUV�XVHG��RQO\�WKH�,76�UHJLRQ��)LJ���%��'��)LJ��6�&��)��FRQWDLQHG�VXIILFLHQW�LQIRUPD-WLRQ�WR�GLVWLQJXLVK�EHWZHHQ�WKH�WZR�VSHFLHV��2WKHU�VWXGLHV�KDYH�IRXQG�WKH�,76�UHJLRQ�WR�EH�PRUH�LQIRUPDWLYH�WKDQ�FKORURSODVW�PDUNHUV�LQ�PDNLQJ�FRUUHFW�'1$�LGHQWLILFDWLRQV�WR�WKH�VSHFLHV�OHYHO��*RQ]DOH]��DO���������&KHQ��DO���������5HQ��DO����������7KH�SRWHQWLDO�WR�GLDJQRVH�WKHVH�VSHFLHV�ZLWK�,76�VHTXHQFHV�FRXOG�EH�SDUWLFXODUO\�XVHIXO�VLQFH�PRVW�RI�WKH�GLDJQRVWLF�PRU-SKRORJLFDO�FKDUDFWHUV�RFFXU�LQ�UHSURGXFWLYH�VWUXFWXUHV��ZKLFK�PD\�QRW�EH�DYDLODEOH�

$�UHFHQW�GLYHUJHQFH�VXFK�DV�WKDW�EHWZHHQ�A. digitata�DQG�A. kilima�FDQQRW�EH�FDOFXODWHG�ZLWK�KLJK�DFFXUDF\�ZLWK�WKH�FXUUHQWO\�DYDLODEOH�GDWD��6HYHUDO�FORFN�FDOLEUDWLRQV�DUH�DYDLO-DEOH�IRU�,76��UHYLHZHG�LQ�.D\��DO����������DQG�WKHVH�YDU\�E\�PRUH�WKDQ�DQ�RUGHU�RI�PDJQLWXGH��EHWZHHQ������î���–"�DQG����î���–"�VXEVWLWXWLRQV�VLWH�\HDU���7KLV�ZRXOG�FRUUHVSRQG�WR�DQ�DJH�RI���±�����0D�IRU�WKH�DJH�RI�WKH�GLYHUJHQFH�RI�A. kilima DQG�A. digitata��UDQJLQJ�IURP�WKH�0LRFHQH�WR�WKH�ODWH�3OHLVWRFHQH���,I�ZH�FRQVLGHUHG�WKH�DJH�RI�WKH�HDUOLHVW�GLYHUJHQFH�LQ�Adansonia WR�EH���±��0D�DV�LQIHUUHG�E\�%DXP��DO����������DQG�HVWLPDWHG�WKH�GDWH�RI�WKH�GLYHUJHQFH�EHWZHHQ�A. kilima DQG�A. digitata EDVHG�RQ�WKH�QXPEHU�RI�,76�VXEVWLWXWLRQV��LW�ZRXOG�EH�EHWZHHQ�����DQG������0D��3OHLVWRFHQH���(YLGHQFH�WKDW�PXWDWLRQ�UDWHV�DUH�IDVWHU�DW�KLJKHU�WHPSHUDWXUHV��*LOORRO\��DO���������IRU�H[-DPSOH�LQ�WKH�+RORFHQH��UHODWLYH�WR�WKH�3OHLVWRFHQH���PD\�OHDG�WR�RYHUHVWLPDWHV�RI�GLYHUJHQFH�WLPH�IRU�UHFHQW�GLYHUJHQFHV��VR�WKH�GLYHUJHQFH�EHWZHHQ�A. digitata�DQG�A. kilima�FRXOG�EH�PRUH�UHFHQW�WKDQ�ZH�KDYH�HVWLPDWHG�KHUH��7KH�IDFW�WKDW�WKH�,76�VHTXHQFH�EXW�QRW�WKH�FKORURSODVW�VHTXHQFHV�FRXOG�EH�XVHG�WR�GLVWLQJXLVK�WKH�WZR�VSHFLHV�VXSSRUWV�D�UHODWLYHO\�UHFHQW�GLYHU-JHQFH��)LJXUH���VKRZV�D�FODGRJUDP�RI�WKH�UHODWLRQVKLSV�ZLWKLQ�Adansonia��WDNLQJ�LQWR�DFFRXQW�WKH�QHZ�VSHFLHV�A. kilima�DQG�WKH�PROHFXODU�ZRUN�SUHVHQWHG�KHUH�

Ploidy of Adansonia digitata. — Adansonia digitata� LV�SUHVXPHG�WR�EH�DQ�DXWRWHWUDSORLG��%DXP��2JLQXPD���������:H�GLG�QRW�ILQG�D�VHFRQG�$IULFDQ�GLSORLG�GXULQJ�RXU�VXUYH\��WKRXJK�ZH�GLG�QRW�LQFOXGH�D�WKRURXJK�VXUYH\�RI�:HVW�$IULFD��:H�ZHUH�SURYLGHG�ZLWK�VRPH�VDPSOHV�IURP�:HVW�$IULFD��VHH�0DWHULDOV�DQG�0HWKRGV��WKDW�ZHUH�DOO�A. digitata��,W�LV�XQOLNHO\�WKDW�D�VHFRQG�GLSORLG�ZLOO�EH�IRXQG�LQ�:HVW�$IULFD�LI�LW�KDV�WKH�VDPH�SUHIHUHQFH�IRU�D�FRROHU�FOLPDWH�DV�A. kilima��,W� LV�DOVR�SRVVLEOH�WKDW�D�VHFRQG�GLSORLG�PD\�RQFH�KDYH�H[LVWHG�EXW�LV�QR�ORQJHU�H[WDQW��,Q�HLWKHU�FDVH��WKH�SRWHQWLDO�H[LVWV�WKDW�A. digitata LV�DQ�DOORWHWUDSORLG��7KH�PRVW�SDUVLPRQLRXV�FKDUDFWHU�UHFRQ-VWUXFWLRQ�IRU�SORLG\�OHYHO�LQIHUV�D�GLSORLG�DQFHVWRU�IRU�WKH�JHQXV�Adansonia��ZLWK�D�UHFHQW�WUDQVLWLRQ�WR�WHWUDSORLG\�RFFXUULQJ�GXULQJ�RU�VKRUWO\�DIWHU�WKH�VSHFLDWLRQ�HYHQW�EHWZHHQ�A. digitata DQG�A. kilima.�7KLV�VXSSRUWV�WKH�LGHD�WKDW�WKH�WZR�VKDUHG�D�FRP-PRQ�DQFHVWRU�YHU\�UHFHQWO\��,W�VHHPV�OLNHO\�WKDW�A. digitata�LV�DQ�DXWRWHWUDSORLG�RI�WKLV�FRPPRQ�DQFHVWRU�

+HWHURVLV�LQ�SRO\SORLGV�FDQ�FRQIHU�RQ�WKHP�JUHDWHU�YLJ-RXU�UHODWLYH�WR�WKHLU�GLSORLG�DQFHVWRUV��DQG�JHQH�GXSOLFDWLRQ�FDQ�DOORZ�WKHP�WR�EH�PRUH�UHVLVWDQW�WR�WKH�HIIHFWV�RI�GHOHWHUL-RXV�PXWDWLRQV��&RPDL���������+RZHYHU��RYHU�ODUJH�JHRORJLFDO�WLPH�VFDOHV��WKHUH�DSSHDUV�WR�EH�QR�VWURQJ�EHQHILW�WR�SRO\SORLGV�

RYHU�WKHLU�GLSORLG�DQFHVWRUV��:RRG��DO����������'HVSLWH�WKLV�LW�VKRXOG�EH�QRWHG�WKDW�WKH�WHWUDSORLG�A. digitata�KDV�VHYHUDO�IHDWXUHV�ZKLFK�PLJKW�IDYRXU�DUWLILFLDO�VHOHFWLRQ�E\�KXPDQV�UHODWLYH�WR�A. kilima��LQ�SDUWLFXODU��WKH�ODUJHU�IUXLWV��7KLV��FRX-SOHG�ZLWK�WKH�VWRPDWDO�SURSHUWLHV�WKDW�PD\�DOORZ�A. digitata�WR�FRORQLVH�GU\HU��KRWWHU�DQG�FRDVWDO�HQYLURQPHQWV��PLJKW�H[SODLQ�LWV�EURDGHU�GLVWULEXWLRQ�LQ�$IULFD�UHODWLYH�WR�A. kilima�

Altitudinal distribution of African species. —�$OWKRXJK�VWRPDWD�LQ�GLSORLGV�DUH�VPDOOHU�WKDQ�WKRVH�RI�WHWUDSORLGV��WKH�PXFK�KLJKHU�VWRPDWDO�GHQVLW\�LQ�GLSORLGV�OHDGV�WR�D�QHW�LQFUHDVH�LQ�ZDWHU�ORVV�IURP�FORVHG�VWRPDWD�DW�QLJKW��&DLUG��DO����������7KH�GHFUHDVHG�VWRPDWDO�GHQVLW\�PHDQV�WKDW�SRO\SORLGV�FDQ�EH�PRUH�WROHUDQW�RI�ZDWHU�GHILFLW�WKDQ�GLSORLGV��/L��DO����������7KLV�PD\�H[SODLQ�WKH�SUHIHUHQFH�RI�WKH�GLSORLG�A. kilima�IRU�HOHYDWLRQV�DERYH�����P��ZKHUH�KLJKHU�PRLVWXUH�LV�DYDLODEOH��,Q�WKH�/LPSRSR�UHJLRQ��WKHUH�ZDV�PRUH�RYHUODS�EHWZHHQ�GLSORLGV�DQG�WHWUDSORLGV�DW�ORZHU�DOWLWXGHV�WKDQ�ZH�REVHUYHG�LQ�.HQ\D�DQG�7DQ]DQLD��7KLV�LV�FRQVLVWHQW�ZLWK�WKH�LGHD�WKDW�GLVWULEX-WLRQ�LV�UHODWHG�WR�VWRPDWDO�FKDUDFWHULVWLFV��VLQFH�WKH�FOLPDWH�LV�FRROHU�DQG�PRLVWHU�DW�WKLV�VRXWKHUQ�OLPLW�RI�EDREDE�GLVWULEX-WLRQ��$Q�DOWLWXGH�FHLOLQJ�RI�a�����P�FRXOG�EH�UHODWHG�WR�WKH�H[WUHPH�VHQVLWLYLW\�RI�WKH�KLJK�ZDWHU�FRQWHQW�EDREDE�WR�IURVW��:LFNHQV��������:LFNHQV��/RZH���������IURVW�LV�RIILFLDOO\�UHFRUGHG�GRZQ�WR������P�IRU�PDQ\�FURSV�LQ�WKH�.HQ\DQ�DQG�7DQ]DQLDQ�KLJKODQGV��EXW�SUREDEO\�UHDFKHV�HYHQ�ORZHU�DW�LQ-IUHTXHQW�LQWHUYDOV�

Concluding comments. —�,W�LV�VRPHZKDW�VXUSULVLQJ�WKDW�D�VHSDUDWH�EDREDE�VSHFLHV�KDV�FR�H[LVWHG�DORQJVLGH�A. digitata ZLWKRXW�GHWHFWLRQ�ZKHQ�ERWK�DUH�FRPPRQ�DQG�ZLGHO\�GLVWULE-XWHG��7KHUH�KDYH�EHHQ�RFFDVLRQDO�VXJJHVWLRQV�WKDW�WKHUH�PLJKW�EH�PRUH�WKDQ�RQH�VSHFLHV�RI�EDREDE�LQ�$IULFD��,Q�������%XUWRQ�LGHQWLILHG�LQODQG�DQG�FRDVWDO�YDULHWLHV�DV�ZHOO�DV�QRUWKHUQ�DQG�VRXWKHUQ�YDULHWLHV��EDVHG�RQ�GLIIHULQJ�PRUSKRORJ\��%XUWRQ���������2QH�RI�WKHVH�ZDV�GHVFULEHG�DV�UDUH�DQG�IRXQG�RQO\�LQ�WKH�8VDJDUD�PRXQWDLQV��LW�KDG�VPDOOHU�OHDYHV�WKDQ�WKH�ORZODQG�EDREDE��,Q�������&KHYDOLHU�GLVWLQJXLVKHG�EHWZHHQ�WZR�VSH-FLHV��RQH�LQ�WKH�ZHVW�DQG�RQH�LQ�WKH�HDVW��KLV�GLVWLQFWLRQ�ZDV�EDVHG�RQ�GLIIHUHQWO\�VL]HG�IUXLWV��&KHYDOLHU���������1RQH�RI�WKHVH�YDULHWLHV�ZHUH�ZLGHO\�DFFHSWHG��:LWK�WKH�H[FHSWLRQ�RI�%XUWRQ¶V�8VDJDUD�PRXQWDLQ�YDULHW\��QRQH�RI�WKH�GHVFULSWLRQV�FRUUHVSRQGV�ZLWK�WKDW�RI�A. kilima��DQG�LW�VHHPV�OLNHO\�WKDW WKH�RWKHU�YDULHWLHV�DUH�VLPSO\�D�PDQLIHVWDWLRQ�RI�WKH�JUHDW�PRU-SKRORJLFDO�DQG�JHQHWLF�GLYHUVLW\�ZKLFK�LV�REVHUYHG�ZLWKLQ�WKH�$IULFDQ�EDREDE�SRSXODWLRQ��:LFNHQV��������6LGLEH��:LOOLDPV��������$VVRJEDGMR��DO���������.\QGW��DO���������3RFN�7V\��DO���������&XQL�6DQFKH]��DO����������6RPH�RI�WKH�REVHUYHG�YDULDELOLW\��DQG�SRVVLEO\�%XUWRQ¶V�8VDJDUD�PRXQWDLQ�YDULHW\��LV�OLNHO\�H[SODLQDEOH�E\�WKH�SUHVHQFH�RI�A. kilima� LQ�$IULFD��7KH�IDFW�WKDW�YDULDELOLW\�LV�ZHOO�NQRZQ��LQ�FRPELQDWLRQ�ZLWK�WKH�VXSHUILFLDO�VLPLODULW\�RI�WKH�WZR�VSHFLHV��PD\�H[SODLQ�ZK\�A. kilima�UHPDLQHG�XQGHWHFWHG�IRU�VR�ORQJ�DPRQJVW�WKH�A. digi-tata�SRSXODWLRQ�

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TAXONOMIC CONCLUSIONS

:H�KDYH�LGHQWLILHG�D�QHZ�GLSORLG�$IULFDQ�VSHFLHV�LQ�Adan-sonia��:H�QDPH�WKLV�VSHFLHV�A. kilima 3HWWLJUHZ��%HOO��%KDJ-ZDQGLQ��*ULQDQ�� -LOODQL��0H\HU��:DEX\HOH��9LFNHUV��7KH�HSLWKHW�kilima�LV�GHULYHG�IURP�6ZDKLOL��µRI�WKH�KLOOV¶��7KH�IORUDO�FKDUDFWHULVWLFV�RI�A. kilima�SODFH�LW�ZLWK�A. digitata�LQ�A. VHFW��Adansonia.

Adansonia kilima�3HWWLJUHZ��%HOO��%KDJZDQGLQ��*ULQDQ��-LO-ODQL��0H\HU��:DEX\HOH��9LFNHUV��sp. nov.�±�+RORW\SH��6RXWK�$IULFD��/LPSRSR��RQ�VRXWK�VLGH�RI�WKH�URDG�EHWZHHQ�7VKNXZL�DQG�7VKLWXQL�����NP�(DVW�RI�WKH�1��+LJKZD\������������6����������(��Jack Pettigrew & Chloë Callis-temon 1�����1RY�������35(������������3DUDW\SH��1DPLELD��&DSULYL�6WULS�����������6����������(��

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Diagnosis. – Adansoniae digitatae� WHWUDSORLGHDH�DI¿QLV�VHG�GLIIHUW�ÀRULEXV�GLPLGLR�PLQRULEXV��SHWDOLV�DQGURHFLR�VX-SHUDQWLEXV��IROLLV�EUHYLRULEXV��VWRPDWLEXV�PLQRULEXV�GHQVL-RULEXVTXH��SROOLQH�VSLQRVLRUH��VSLQLV����±�����LQ�A. digitata ��±������PLQRUH������P��LQ�A. digitata ����P�GLDPHWUR���¿ODPHQWLV�OLEHULV�VWDPLQXP�SDXFLRULEXV�����±�����LQ�A. digi-tata����±�������&KURPRVRPDWXP�QXPHUXV�GLSORLGHXV�����

/LNH�WKH�WHWUDSORLG�A. digitata��EXW�GLIIHUV�LQ�KDYLQJ�IORZ-HUV�KDOI�WKH�VL]H��SHWDOV�VKRUWHU�WKDQ�WKH�DQGURHFLXP�ZLWK�QR�NLQHVLV��VPDOOHU�OHDYHV��VPDOOHU�DQG�GHQVHU�VWRPDWD��VPDOOHU�DQG�PRUH�GHQVHO\�VSLQRVH�SROOHQ��44��P�LQ�GLDPHWHU�FRPSDUHG�ZLWK�����P�LQ�A. digitata���IHZHU�IUHH�VWDPLQDO�ILODPHQWV�����±�����FRPSDUHG�ZLWK����±�����LQ�A. digitata���GLSORLG�FKURPRVRPH�QXPEHU��n� ����

Description� �PRGLILHG�DQG�H[WHQGHG�IURP�WKH�GHVFULS-WLRQ�RI�A. digitata�E\�%DXP�������ZKHUH�DSSURSULDWH���±�7UHH�UHDFKLQJ����P�LQ�KHLJKW�DQG�XS�WR���P�GLDP���XVXDOO\�ZLWK�VLQJOH� F\OLQGULFDO� WUXQNV� DQG� VSUHDGLQJ�� URXQGHG� FURZQV��%UDQFKHV� LUUHJXODUO\� GLVWULEXWHG�� %DUN� GDUN� JUH\�� VPRRWK�WR�LUUHJXODUO\�WXEHUFXODWH��/HDYHV��±��±���IROLRODWH��OHDIOHWV�

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Distribution and ecology. – ,Q�RXU�OLPLWHG�VXUYH\��A. kilima ZDV�IRXQG�IURP�WKH�HDVWHUQ�VORSHV�RI�0W��.LOLPDQMDUR��VRXWK�WR�VRXWKHUQ�7DQ]DQLD�DQG�ZHVW�WR�QRUWKHUQ�1DPLELD��DV�ZHOO�DV�LQ�WKH�9HQGD�RI�6RXWK�$IULFD��UHVWULFWHG�WR�HOHYDWLRQV�EHWZHHQ�����DQG������P��Adansonia digitata�EXW�QRW�A. kilima�ZHUH�IRXQG�LQ�FRDVWDO�1DPLELD�DQG�6HQHJDO�

ACKNOWLEDGEMENTS

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Adanson, M. ������'HVFULSWLRQ�G¶XQ�DUEUH�G¶XQ�QRXYHDX�JHQUH�DSSHOp�%DREDE��REVHUYp�DX�6pQpJDO��Mem. Acad. Roy. Sci. (Paris)��������±����

Armstrong, P. ������%DREDEV��5HPQDQW�RI�*RQGZDQDODQG"�Trees South Africa�������±���

Armstrong, P. ������7KH�GLVMXQFW�GLVWULEXWLRQ�RI�WKH�JHQXV�Adansonia��Natl. Geogr. J. India��������±����

Armstrong, P. ������6W�-DJR��&DSH�9HUGH��6RXUFH�RI�LQVSLUDWLRQ��3S����±���LQ��$UPVWURQJ��3���HG����Darwin’s other islands��/RQGRQ��1HZ�<RUN��&RQWLQXXP�

Aryavand, A., Ehdaie, B., Tran, B. & Waines, J.G. ������6WRPDWDO�IUHTXHQF\�DQG�VL]H�GLIIHUHQWLDWH�SORLG\�OHYHOV�LQ�Aegilops neglecta��Genet. Resources Crop Evol.��������±����

Assogbadjo, A., Kyndt, T., Sinsin, B., Gheysen, G. & Van Damme, P. ������3DWWHUQV�RI�JHQHWLF�DQG�PRUSKRPHWULF�GLYHUVLW\�LQ�%DREDE�(Adansonia digitata��SRSXODWLRQV�DFURVV�GLIIHUHQW�FOLPDWLF�]RQHV�RI�%HQLQ��:HVW�$IULFD���Ann. Bot. (London)��������±����

Assogbadjo, A.E., Sinsin, B., Codjia, J.T.C. & Van Damme, P. ������(FRORJLFDO�GLYHUVLW\�DQG�SXOS��VHHG�DQG�NHUQDO�SURGXFWLRQ�RI�WKH�EDREDE��Adansonia digitata��LQ�%HQLQ��Belg. J. Bot.��������±���

Baum, D.A. ������$�V\VWHPDWLF�UHYLVLRQ�RI�Adansonia��%RPEDFDFHDH���Ann. Missouri Bot. Gard.��������±����

Baum, D.A. & Oginuma, K. ������$�UHYLHZ�RI�FKURPRVRPH�QXPEHUV�LQ�%RPEDFDFHDH�ZLWK�QHZ�FRXQWV�IRU�Adansonia��Taxon�������±���

Baum, D.A., Small, R.L. & Wendel, J.F. ������%LRJHRJUDSK\�DQG�IORUDO�HYROXWLRQ�RI�EDREDEV��Adansonia��%RPEDFDFHDH��DV�LQIHUUHG�IURP�PXOWLSOH�GDWD�VHWV��Syst. Biol.��������±����

Baum, D.A., Witt Smith, S., Yen, A., Alverson, W.S., Nyffeler, R., Whitlock, B.A. & Oldham, R.L. ������3K\ORJHQHWLF�UHODWLRQ-VKLSV�RI�Malvatheca �%RPEDFRLGHDH�DQG�0DOYRLGHDH��0DOYDFHDH�VHQVX�ODWR��DV�LQIHUUHG�IURP�SODVWLG�'1$�VHTXHQFHV��Amer. J. Bot. ��������±�����

Bingham, E.T. ������6WRPDWDO�FKORURSODVWV�LQ�DOIDOID�DW�IRXU�SORLG\�OHYHOV��Crop Sci.�������±����

Blench, R. ������7KH�LQWHUWZLQHG�KLVWRU\�RI�WKH�VLON�FRWWRQ�DQG�EDREDE��3S���±���LQ��9DQ�GHU�9HHQ��0���HG����Fields of change: Progress in African archeobotany.�*URQLQJHQ��%DUNKXLV��*URQLQJHQ�8QL-YHUVLW\�

Bowman, D.M.J.S. ������2EVHUYDWLRQV�RQ�WKH�GHPRJUDSK\�RI�WKH�$XV-WUDOLDQ�%RDE��Adansonia gibbosa��LQ�WKH�1RUWK�ZHVW�RI�WKH�1RUWK-HUQ�7HUULWRU\��$XVWUDOLD��Austral. J. Bot.��������±����

Brummitt, R.K. ������5HSRUW�RI�WKH�&RPPLWWHH�IRU�6SHUPDWRSK\WD������Taxon��������±����

Burton, R.F. ������7KH� ODNH�UHJLRQV�RI�&HQWUDO�(TXDWRULDO�$IULFD��J. Roy. Geogr. Soc.������±����

Caird, M.A., Richards, J.H. & Donovan, L.A. ������1LJKWWLPH�VWRPD-WDO�FRQGXFWDQFH�DQG�WUDQVSLUDWLRQ�LQ�&#�DQG�&$�SODQWV��Pl. Physiol. ������±���

Chen, G., Sun, W.-B. & Sun, H. ������0RUSKRORJLFDO�FKDUDFWHULVWLFV�RI�OHDI�HSLGHUPLV�DQG�VL]H�YDULDWLRQ�RI�OHDI��IORZHU�DQG�IUXLW�LQ�GLIIHUHQW�SORLG\�OHYHOV�LQ�Buddleja macrostachya��%XGGOHMDFHDH���J. Syst. Evol.��������±����

Chen, S.L., Yao, H., Han, J.P., Liu, C., Song, J.Y., Shi, L.C., Zhu, Y.J., Ma, X.Y., Gao, T., Pang, X.H., Luo, K., Li, Y., Li, X.W., Jia, X.C., Lin, Y.L. & Leon, C. ������9DOLGDWLRQ�RI�WKH�,76��UHJLRQ�DV�D�QRYHO�'1$�EDUFRGH�IRU�LGHQWLI\LQJ�PHGLFLQDO�SODQW�VSHFLHV��PLoS One����H������GRL����������MRXUQDO�SRQH���������

Chevalier, A. ������/HV�%DREDEV��Adansonia��GH�O¶$IULTXH�FRQWLQHQWDOH��Bull. Soc. Bot. France��������±����

Comai, L. ������7KH�DGYDQWDJHV�DQG�GLVDGYDQWDJHV�RI�EHLQJ�SRO\SORLG��Nat. Rev. Genet.�������±����

Cuni Sanchez, A., Haq, N. & Assogbadjo, A. ������9DULDWLRQ�LQ�EDREDE�(Adansonia digitata�/���OHDI�PRUSKRORJ\�DQG�LWV�UHODWLRQ�WR�GURXJKW�WROHUDQFH��Genet. Resources Crop Evol.�������±���

De Caluwé, E., Halamová, K. & Van Damme, P. ������%DREDE�

LITERATURE CITED (Adansonia digitata�/����$�UHYLHZ�RI�WUDGLWLRQDO�XVHV��SK\WRFKHP-LVWU\�DQG�SKDUPDFRORJ\��3S����±���LQ��-XOLDQL��+�5���6LPRQ��-���+R��&��7���HGV����African natural plant products: New discover-ies and challenges in chemistry and quality��:DVKLQJWRQ��'�&���$PHULFDQ�&KHPLFDO�6RFLHW\�

Drummond, A.J., Ashton, B., Buxton, S., Cheung, M., Cooper, A., Duran, C., Field, M., Heled, J., Kearse, M., Markowitz, S., Moir, R., Stones-Havas, S., Sturrock, S., Thierer, T. & Wilson, A. ������*HQHLRXV��YHUVLRQ������KWWS���ZZZ�JHQHLRXV�FRP��

Duarte, M.C., Esteves, G.L., Salatino, M.L.F., Walsh, K.C. & Baum, D.A. ������3K\ORJHQHWLF�DQDO\VHV�RI�Eriotheca�DQG�UHODWHG�JHQHUD��%RPEDFRLGHDH��0DOYDFHDH���Syst. Bot.��������±����

Evans, A.M. ������7KH�SURGXFWLRQ�DQG�LGHQWLILFDWLRQ�RI�SRO\SORLGV�LQ�UHG�FORYHU��ZKLWH�FORYHU�DQG�OXFHUQH��New Phytol.��������±����

Gebauer, J., ElSiddig, K. & Ebert, G. ������%DREDE��Adansonia digi-tata�/����$�UHYLHZ�RQ�D�PXOWLSXUSRVH�WUHH�ZLWK�SURPLVLQJ�IXWXUH�LQ�WKH�6XGDQ��Gartenbauwissenschaft�����6XSSO�������±����

Gillooly, J.F., McCoy, M.W. & Allen, A.P. ������(IIHFWV�RI�PHWDEROLF�UDWH�RQ�SURWHLQ�HYROXWLRQ��Biol. Lett.�������±����

Gonzalez, M.A., Baraloto, C., Engel, J., Mori, S.A., Petronelli, P., Riera, B., Roger, A., Thebaud, C. & Chave, J. ������,GHQWLIL-FDWLRQ�RI�DPD]RQLDQ�WUHHV�ZLWK�'1$�EDUFRGHV��PLoS One �������H������GRL����������MRXUQDO�SRQH���������

Hochreutiner, B.P.G. ������8Q�QRXYHDX�EDREDE�±�5HYLVLRQ�GX�JHQUH�Adansonia��Annuaire Conserv. Jard. Bot. Genève����������±����

Huelsenbeck, J.P. & Ronquist, F. ������05%$<(6��%D\HVLDQ�LQIHU-HQFH�RI�SK\ORJHQHWLF�WUHHV��Bioinformatics��������±����

Kay, K., Whittall, J. & Hodges, S. ������$�VXUYH\�RI�QXFOHDU�ULERVRPDO�LQWHUQDO�WUDQVFULEHG�VSDFHU�VXEVWLWXWLRQ�UDWHV�DFURVV�DQJLRVSHUPV��$Q�DSSUR[LPDWH�PROHFXODU�FORFN�ZLWK�OLIH�KLVWRU\�HIIHFWV��B. M. C. Evol. Biol.��������GRL�������������������������

Kress, W.J., Erickson, D.L., Jones, F.A., Swenson, N.G., Perez, R., Sanjur, O. & Bermingham, E. ������3ODQW�'1$�EDUFRGHV�DQG�D�FRPPXQLW\�SK\ORJHQ\�RI�D�WURSLFDO�IRUHVW�G\QDPLFV�SORW�LQ�3DQDPD��Proc. Natl. Acad. Sci. U.S.A. ����������±������

Kress, W.J., Wurdack, K.J., Zimmer, E.A., Weigt, L.A. & Janzen, D.H. ������8VH�RI�'1$�EDUFRGHV�WR�LGHQWLI\�IORZHULQJ�SODQWV��Proc. Natl. Acad. Sci. U.S.A.����������±�����

Kyndt, T., Assogbadjo, A.E., Hardy, O.J., Glele Kakai, R., Sinsin, B., Van Damme, P. & Gheysen, G. ������6SDWLDO�JHQHWLF�VWUXFWXU-LQJ�RI�EDREDE��Adansonia digitata��0DOYDFHDH��LQ�WKH�WUDGLWLRQDO�DJURIRUHVWU\�V\VWHPV�RI�:HVW�$IULFD��Amer. J. Bot.��������±����

Li, W.-L., Berlyn, G.P. & Ashton, P.M.S. ������3RO\SORLGV�DQG�WKHLU�VWUXFWXUDO�DQG�SK\VLRORJLFDO�FKDUDFWHULVWLFV�UHODWLYH�WR�ZDWHU�GHILFLW�LQ�Betula papyrifera �%HWXODFHDH���Amer. J. Bot.�������±���

Linneaus, C. ������Species plantarum���QG�HG��6WRFNKROP��6DOYLXV�Livingstone, D. ������A popular account of missionary travels and

researches in South Africa��/RQGRQ��0XUUD\�Mishra, M. ������6WRPDWDO�FKDUDFWHULVWLFV�DW�GLIIHUHQW�SORLG\�OHYHOV�LQ�

Coffea�/��Ann. Bot. (London)��������±����Patrut, A., Reden, K.F. von, Lowy, D.A., Alberts, A.H., Pohlman,

J.W., Wittmann, R., Gerlach, D., Xu, L. & Mitchell, C.S. ������5DGLRFDUERQ�GDWLQJ�RI�D�YHU\�ODUJH�$IULFDQ�EDREDE��Tree Physiol. ��������±�����

Pock Tsy, J.-M.L., Lumaret, R., Mayne, D., Vall, A.O.M., Abutaba, Y.I.M., Sagna, M., Raoseta, S.O.R. & Danthu, P. ������&KOR-URSODVW�'1$�SK\ORJHRJUDSK\�VXJJHVWV�D�:HVW�$IULFDQ�FHQWUH�RI�RULJLQ�IRU�WKH�EDREDE��Adansonia digitata�/���%RPEDFRLGHDH��0DO-YDFHDH���Molec. Ecol.���������±�����

Ramsey, J. & Schemske, D.W. ������1HRSRO\SORLG\� LQ� IORZHULQJ�SODQWV��Annual Rev. Ecol. Syst.��������±����

Ren, B.Q., Xiang, X.G. & Chen, Z.D. ������6SHFLHV�LGHQWLILFDWLRQ�RI�Alnus��%HWXODFHDH��XVLQJ�QU'1$�DQG�FS'1$�JHQHWLF�PDUNHUV��Molec. Ecol. Resources��������±����

Ronquist, F. & Huelsenbeck, J.P. ������0U%D\HV����%D\HVLDQ�SK\ORJH-QHWLF�LQIHUHQFH�XQGHU�PL[HG�PRGHOV��Bioinformatics���������±�����

Sidibe, M. & Williams, J.T. ������Boabab: Adansonia digitata L.:

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Venter, S.M. & Witkowski, E.T.F. ������%DREDE��Adansonia digi-tata�/���GHQVLW\��VL]H�FODVV�GLVWULEXWLRQ�DQG�SRSXODWLRQ�WUHQGV�EH-WZHHQ�IRXU�ODQG�XVH�W\SHV�LQ�QRUWKHUQ�9HQGD��6RXWK�$IULFD��Forest Ecol. Managem.���������±����

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Wood, T.E., Takebayashi, N., Barker, M.S., Mayrose, I., Greenspoon, P.B. & Rieseberg, L.H. ������7KH�IUHTXHQF\�RI�SRO\SORLG�VSHFLDWLRQ�LQ�YDVFXODU�SODQWV��Proc. Natl. Acad. Sci. U.S.A.�����������±������

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Danthu Adi2 �0(/���±��±�� -;��������A. digitata,a,c U.S.A., ,RZD��FXOWLYDWHG�VSHFLPHQ���Small s.n. �,6&���$)��������A. digitata,b,c Burkina Faso, 3REp�0HQJDR�7RXIOp��Baum 349 �02���±��±��$)��������A. digitata,a,d XQSXEOLVKHG� ±��±� $)��������A. digitata,e Kenya, FROOHFWRU�XQNQRZQ 770032002 �37%*���±��+4��������±��A. digitata,f XQSXEOLVKHG� ±� $<��������±��A. grandidieri Baill.,c,e Madagascar, 7ROLDUD�����������6��������(��Baum 345 �02���±��+4��������$)��������A. grandidieri,c Madagascar, 7ROLDUD��Baum 347 �02���±��±��$)��������A. gregorii Muell., Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 81 �QR�YRXFKHU��� -1�������� -1��������±��A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 76 �QR�YRXFKHU��� -1��������±��±��A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 37 �QR�YRXFKHU��� -1�������� -1��������±��A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 233 �QR�YRXFKHU���±�� -1��������±��A. gregorii, South Africa, *DXWHQJ��FXOWLYDWHG�VSHFLPHQ������������6����������(��J. Pettigrew 400 �QR�YRXFKHU���±�� -1��������±��A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 70 �QR�YRXFKHU���±��±�� -1��������A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6����������(��J. Pettigrew 220 �QR�YRXFKHU���±��±�� -1��������A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 85 �QR�YRXFKHU���±��±�� -1��������A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 73 �QR�YRXFKHU���±��±�� -1��������A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 201 �QR�YRXFKHU���±��±�� -1��������A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. Pettigrew 222 �QR�YRXFKHU���±��±�� -1��������A. gregorii, Australia, :HVWHUQ�$XVWUDOLD�����������6�����������(��J. 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Perez 445949 �6&=���*4��������±��±��C. platanifolia,e U.S.A., )ORULGD��FROOHFWRU�XQNQRZQ FG83343A �)7*���±��+4��������+4������D�,GHQWLILHG�DV�A. digitata�LQ�WKH�RULJLQDO�SXEOLFDWLRQ��9RXFKHU�QRW�H[DPLQHG�LQ�WKH�FXUUHQW�VWXG\�E�,GHQWLILHG�DV�A. digitata�LQ�WKH�RULJLQDO�SXEOLFDWLRQ��,GHQWLW\�FRQILUPHG�E\�'��%DXP��SHUV��FRPP��F�%DXP��DO�������G�6KL��<XDQ��XQSXE�H�'XDUWH��DO�������I�<XDQ��DO���XQSXE�J�.UHVV��DO�������