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INTRODUCTION

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Molecular and morphological analysis of subfamily Alooideae (Asphodelaceae) and the inclusion of Chortolirion in Aloe

Barnabas H. Daru,! John C. Manning,"#$ James S. Boatwright,% Olivier Maurin,! Norman Maclean,& Hanno Schaefer,' Maria Kuzmina( & Michelle van der Bank!

1 African Centre for DNA Barcoding, University of Johannesburg, P.O. Box 524 Auckland Park 2006, Johannesburg, South Africa2 Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa�� Research Centre for Plant Growth and Development, School of Biological and Conservation Sciences, University of

KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa�� Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, Cape Town,

South Africa�� School of Biological Sciences, University of Southampton, Highfield, Southampton, Hants, SO16 7PX, U.K.6 Technische Universitaet Muenchen, Biodiversitaet der Pflanzen, Maximus-von-Imhof Forum 2, 85354 Freising, Germany�� International Barcode of Life Project, Biodiversity Institute of Ontario, University of Guelph, Guelph, Ontario N1G 2W1, Canada$XWKRU�IRU�FRUUHVSRQGHQFH� Barnabas H. Daru, darunabas@gmail.com

Abstract $VSKRGHODFHDH�VXEIDP��$ORRLGHDH��$VSDUDJDOHV��FXUUHQWO\�FRPSULVHV�ILYH�JHQHUD��IRXU�RI�ZKLFK�DUH�HQGHPLF�WR�VRXWKHUQ�$IULFD��'HVSLWH�WKHLU�LPSRUWDQFH�LQ�FRPPHUFLDO�KRUWLFXOWXUH�WKH�HYROXWLRQDU\�UHODWLRQVKLSV�DPRQJ�WKH�JHQHUD�DUH�VWLOO�LQFRPSOHWHO\�XQGHUVWRRG��7KLV�VWXG\�H[DPLQHV�SK\ORJHQHWLF�UHODWLRQVKLSV�LQ�WKH�VXEIDPLO\�XVLQJ�DQ�H[SDQGHG�PROHFXODU�VHTXHQFH�GDWDVHW�IURP�WKUHH�SODVWLG�UHJLRQV��matK��rbcLa� trnH-psbA��DQG�WKH�ILUVW�VXEXQLW�RI�WKH�QXFOHDU�ULERVRPDO�LQWHUQDO�WUDQVFULEHG�VSDFHU��,76����6HTXHQFH�GDWD�ZHUH�DQDO\VHG�XVLQJ�PD[LPXP�SDUVLPRQ\�DQG�%D\HVLDQ�VWDWLVWLFV��DQG�VHOHFWHG�PRUSKRORJLFDO�WUDLWV�ZHUH�PDSSHG�RQWR�WKH�PROHFXODU�SK\ORJHQ\��Haworthia LV�FRQILUPHG�DV�EHLQJ�SRO\SK\OHWLF��FRPSULVLQJ�WKUHH�PDLQ�FODGHV�WKDW�ODUJHO\�FRUUHODWH�ZLWK�FXUUHQW�VXEJHQHULF�FLUFXPVFULSWLRQV��Astroloba DQG�Gasteria DUH�HYLGHQWO\�HDFK�PRQRSK\OHWLF�DQG�VLVWHU�UHVSHFWLYHO\�WR�Astroloba DQG�H. VXEJ� Robustipedunculares��Chortolirion LV�VKRZQ�WR�EH�GHHSO\�QHVWHG�ZLWKLQ�Aloe DQG�LV�IRUPDOO\�LQFOXGHG�LQ�WKDW�JHQXV��Aloe LWVHOI�LV�FOHDUO\�SRO\SK\OHWLF��ZLWK�WKH�GZDUI�VSHFLHV�A. aristata DOOLHG�WR�Haworthia VXEJ��Robustipedunculares��7KH�WD[RQRPLF�LPSOLFDWLRQV�RI�WKHVH�ILQGLQJV�DUH�H[DPLQHG�EXW�EUDQFK�VXSSRUW�DW�FULWLFDO�ORZHU�QRGHV�LV�LQVXIILFLHQW�DW�WKLV�VWDJH�WR�MXVWLI\�LPSOHPHQWLQJ�PDMRU�WD[RQRPLF�FKDQJHV�

Keywords $VSKRGHODFHDH� ,76���matK� PROHFXODU SK\ORJHQ\��PRUSKRORJ\��QRPHQFODWXUH� rbcLa� trnH-psbA��WD[RQRP\

Supplementary Material 7KH�(OHFWURQLF�6XSSOHPHQW��)LJV��6�±6���$SSHQGLFHV�6�±6���DQG�WKH�DOLJQPHQW�DUH�DYDLODEOH�LQ�WKH�6XSSOHPHQWDU\�'DWD�VHFWLRQ�RI�WKH�RQOLQH�YHUVLRQ�RI�WKLV�DUWLFOH��KWWS���ZZZ�LQJHQWDFRQQHFW�FRP�FRQWHQW�LDSW�WD[��

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MATERIALS AND METHODS

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XVHG�WR�SURGXFH�D�����PDMRULW\�UXOH�FRQVHQVXV�WUHH�VKRZLQJ�WKH�IUHTXHQFLHV��SRVWHULRU�SUREDELOLWLHV�RU�33��RI�DOO�REVHUYHG�EL�SDUWLWLRQV��7KH�IROORZLQJ�VFDOH�ZDV�XVHG�WR�HYDOXDWH�WKH�33V��EHORZ�������ZHDNO\�VXSSRUWHG������±�����VWURQJO\�VXSSRUWHG��7R�PDS�WKH�ERRWVWUDS�YDOXHV��%6��DQG�33�YDOXHV��0U%D\HV�WUHH��RQWR�WKH�WUHH��WKH�QH[XV�WUHH�ILOH�IURP�WKH�%,�DQDO\VLV�ZDV�UHVFDOHG�XVLQJ�³DSH´�Y��������3DUDGLV��DO���������DQG�³DGH-SK\OR´�Y������-RPEDUW��'UD\��������SDFNDJHV�LPSOHPHQWHG�LQ�5��5�'HYHORSPHQW�&RUH�7HDP��������

Coding of morphological characters. —� $�PDWUL[� RI����PRUSKRORJLFDO�FKDUDFWHUV�ZDV�SUHSDUHG�IRU�WKH���� WD[D RI�$ORRLGHDH�DQG�RXWJURXSV�LQFOXGHG�LQ�WKH�DQDO\VHV��0RVW�WD[RQRPLF�VWXGLHV�LQ�$ORRLGHDH�LGHQWLILHG�WKHVH�GLDJQRVWLF�FKDUDFWHUV��RIWHQ�DW�VSHFLHV�OHYHO��WR�LQIHU�UHODWLRQVKLSV�ZLWKLQ�WKH�VXEIDPLO\��5H\QROGV��������������-HSSH��������%D\HU��������������������������9DQ�-DDUVYHOG��������������������������9DQ�-DDUVYHOG��DO���������*OHQ��6PLWK��������0|VVPHU��DO���������6PLWK������D��E��6PLWK��DO.��������0H\HU��6PLWK��������������*OHQ��+DUG\��������9DQ�:\N��6PLWK��������9DQ�-DDUVYHOG��9DQ�:\N��������������������6PLWK��6WH\Q��������*HUPLVKXL]HQ��DO���������*LOGHQKX\V��������.ORSSHU��6PLWK���������7KHVH�FKDUDFWHUV�ZHUH�VFRUHG�DV�SUHVHQW�RU�DEVHQW�LQ�D�����PDWUL[�DV�LQGLFDWHG�LQ�$SSHQGL[�6���7KH�SDWWHUQV�RI�HYROXWLRQ�RI�WKHVH�FKDUDFWHUV�ZHUH�H[DPLQHG�E\�UHFRQVWUXFW-LQJ�WKHP�RQWR�WKH�PDMRULW\�UXOH�FRQVHQVXV�WUHH�SURGXFHG�E\�WKH�%,�DQDO\VLV�XVLQJ�0HVTXLWH�Y�������0DGGLVRQ��0DGGLVRQ���������0RUSKRORJLFDO�FKDUDFWHUV�DQG�FKDUDFWHU�VWDWHV�DUH�GH-ILQHG�LQ�$SSHQGL[�6��DQG�WKH�GDWD�PDWUL[�XVHG�IRU�FKDUDFWHU�UHFRQVWUXFWLRQV�LV�SUHVHQWHG�LQ�$SSHQGL[�6��

RESULTS

Comparison of sequence partitions. —�7KH�VWDWLVWLFV�IURP�WKH�03�DQDO\VHV�IRU�WKH�VLQJOH�SODVWLG�DQDO\VLV��FRPELQHG�SODV-WLG��PDW.࣠�UEF/D࣠�trnH-psbA���,76��DQG�WKH�WRWDO�FRPELQHG�GDWDVHW�DUH�VKRZQ�LQ�7DEOH����,76��KDG�D�VLJQLILFDQWO\�KLJKHU�

QXPEHU�RI�YDULDEOH�VLWHV�������FRPSDUHG�WR�WKH�SODVWLG�UHJLRQV�FRPELQHG��KHQFHIRUWK�FDOOHG�SODVWLG�GDWDVHW���7KH�QXPEHU�RI�SRWHQWLDOO\�SDUVLPRQ\�LQIRUPDWLYH�FKDUDFWHUV�IRU�WKH�SODVWLG�GDWDVHW�ZLWKLQ�$ORRLGHDH�������LV�PXFK�ORZHU�WKDQ�IRU�,76���������7KH�DYHUDJH�QXPEHU�RI�FKDQJHV�SHU�YDULDEOH�VLWH�IRU�,76�������FKDQJHV�SHU�YDULDEOH�VLWH��UHWHQWLRQ�LQGH[�5,� �������DQG�FRQVLVWHQF\�LQGH[�&,� �������LV�KLJKHU�WKDQ�LQ�WKH�SODVWLG�GDWDVHW������FKDQJHV�SHU�YDULDEOH�VLWH��5,� �������&,� ��������$QDO\VLV�RI�HDFK�RI�WKH�WKUHH�SODVWLG�UHJLRQV��matK� rbcLa� trnH-psbA) UHVXOWHG�LQ�WUHHV�WKDW�ZHUH�VLPLODU�LQ�WRSRORJ\��7UHHV�UHVXOWLQJ�IURP�WKH�IROORZLQJ�DQDO\VHV�DUH�SUHVHQWHG��VLPSOLILHG�WUHH�WR-SRORJLHV�RI�WKH�FRPELQHG�SODVWLG�UHJLRQV��PDW.࣠�UEF/D࣠+࣠WUQ+�psbA��)LJ���$���,76���)LJ���%��DQG�FRPELQHG�SODVWLG�DQG�QXFOHDU�(PDW.࣠�UEF/D࣠+࣠WUQ+�SVE$�,76���)LJ���&���FRPELQHG�SODVWLG�UHJLRQV��PDW.࣠�UEF/D࣠+࣠WUQ+�SVE$���)LJ��6����,76���)LJ��6���DQG�FRPELQHG�SODVWLG�DQG�QXFOHDU�UHJLRQV��PDW.࣠�UEF/D࣠+࣠WUQ+�psbA�,76���)LJ�����

Combined plastid data. —�,QGLYLGXDO�SODVWLG�QXFOHRWLGH�VHTXHQFH�DQDO\VHV��UHVXOWV�QRW�VKRZQ��ZHUH�WRSRORJLFDOO\�FRQ-VLVWHQW��QHJOLJLEOH�WR�]HUR�LQFRQJUXHQFH���DQG�IRU�WKH�SXUSRVH�RI�WKH�UHVXOWV�DQG�GLVFXVVLRQ�ZHUH�FRPELQHG�DQG�WUHDWHG�DV�D�VLQJOH�GDWDVHW��7KH�VWDWLVWLFV�IRU�03�DQDO\VLV�IRU�WKH�FRPELQHG�SODVWLG�GDWD�LV�SUHVHQWHG�LQ�7DEOH����)URP�WKH�KHXULVWLF�VHDUFK��ZH�IRXQG�����PRVW�SDUVLPRQLRXV�SK\ORJHQHWLF�WUHHV�RI�ZKLFK�RQH�LV�SUHVHQWHG�LQ�)LJXUH�6���$ORRLGHDH�DUH�VXSSRUWHG�DV�D�PRQRSK\OHWLF�JURXS��EXW�UHVROXWLRQ�LQ�WKH�UHVW�RI�WKH�WUHH�LV�ORZ��6RPH�JURXSV�LQ�Aloe��Haworthia�VXEJ��Haworthia��H��VXEJ��Robustipedunculares�DQG�Gasteria�ZHUH�PRGHUDWHO\�VXSSRUWHG�

ITS data. —�6XPPDU\�VWDWLVWLFV�IRU�WKH�,76��GDWD�PDWUL[�LV�SUHVHQWHG�LQ�7DEOH����$QDO\VLV�RI�WKH�QXFOHDU�GDWDVHW�DOVR�UHWULHYHG�D�PRQRSK\OHWLF�$ORRLGHDH��DOWKRXJK�WKHUH�LV�VRPH�LQFRQJUXHQFH�ZLWK�WKH�SODVWLG�GDWD��LW�DOORZV�XV�WR�LGHQWLI\�ILYH�PDMRU�$ORRLGHDH�JURXSLQJV��)LJ��6���ZLWK�PRGHUDWH�WR�VWURQJ�VXSSRUW�EXW�SRRU�UHVROXWLRQ�ZLWKLQ�HDFK�JURXS��Haworthia VXEJ��Haworthia��Aloe VHFW��Macrifoliae��A. VHFW. Kumara��D�SRO\WRP\�LQFOXGLQJ WKH�µWUHH�Aloe¶�VSHFLHV�SOXV�Aloe V�VWU���LQFOXGLQJ�Chortolirion�� DQG�D�FODGH�FRPSULVLQJ�Astroloba�

Table .�6WDWLVWLFV�IURP�0RGHO7HVW�DQG�03�DQDO\VHV�REWDLQHG�IURP�VHSDUDWH�DQG�FRPELQHG�GDWDVHWV�

matK rbcLa trnH-psbA&RPELQHG�SODVWLG� ,76�

&RPELQHG� SODVWLG�,76�

1R��RI�WD[D�LQFOXGHG ��� ��� ��� ��� ��� ���1R��RI�LQFOXGHG�FKDUDFWHUV ��� ��� ��� ���� ��� ����1R��RI�FRQVWDQW�FKDUDFWHUV ��� ��� ��� ���� ��� ����1R��RI�YDULDEOH�VLWHV ��������� �������� ��������� ��������� ��������� ���������1R��RI�SDUVLPRQ\�LQIRUPDWLYH�VLWHV ��������� �������� ������� ��������� �������� ���������1R��RI�WUHHV��)LWFK� ��� ��� ��� 661 ��� ����1R��RI�VWHSV��7UHH�OHQJWK� ��� ��� ��� ��� ��� ����&RQVLVWHQF\�LQGH[ ���� ���� ���� ���� ���� ����5HWHQWLRQ�LQGH[ ���� ���� ���� ���� ���� ����$YHUDJH�QXPEHU�RI�FKDQJHV�SHU�YDULDEOH�VLWH� �QXPEHU�RI�VWHSV�QXPEHU�RI�YDULDEOH�VLWHV� ��� ��� ��� ��� ��� ���0RGHO�VHOHFWHG�E\�$NDLNH�LQIRUPDWLRQ�FULWHULRQ 790�* +.<�* 790�,�* 790�,�* 751�* *75�,�*

��

TAXON����������)HEUXDU\���������±��'DUX��DO����Phylogeny of Alooideae

�� 9HUVLRQ�RI�5HFRUG��LGHQWLFDO�WR�SULQW�YHUVLRQ��

Gasteria� Haworthia�VXEJ��Hexangulares�DQG�VXEJ��Robusti-pedunculares� DQG Aloe aristata�

Combined plastid and nuclear dataset. —�$OWKRXJK�WKH�SDUWLWLRQHG�%UHPHU�VXSSRUW�WHVW�LQGLFDWHG�VRPH�LQFRQJUXHQ-FLHV�EHWZHHQ�WKH�QXFOHDU�DQG�SODVWLG�GDWDVHWV��YLVXDO�LQVSHF-WLRQ�RI�WKH�VHSDUDWH�DQDO\VHV��)LJ�����VKRZV�WKDW�QRQH�RI�WKH�VWURQJO\�VXSSRUWHG�FODGHV�ZHUH�PXWXDOO\�LQFRPSDWLEOH��DQG�WHVWV�IRU�LQFRQJUXHQFH�KDYH�EHHQ�VKRZQ�WR�EH�RI�YDULDEOH�UHOLDELOLW\��5HHYHV��DO���������<RGHU��DO����������%DVHG�RQ�WKH�HYLGHQW�FRQJUXHQFH�EHWZHHQ�WKH�WZR�GDWDVHWV��ZH�WKHUHIRUH�FRPELQHG�DOO�GDWD��6HHODQDQ��DO���������:LHQV���������)LJ������6WDWLVWLFV�IRU�WKH�SK\ORJHQHWLF�IUDPHZRUN�RI�WKH�FRQFDWHQDWHG�GDWDVHW�LV�VXP-PDULVHG�LQ�7DEOH����$�PRQRSK\OHWLF�$ORRLGHDH�ZDV�UHFRYHUHG�ZLWK�KLJK�VXSSRUW��)LJ�����%6� �����33� ��������7KH�FRPELQHG�GDWDVHW�UHVROYHG�HLJKW�PDMRU�JURXSLQJV�UHSUHVHQWHG�LQ�GLIIHUHQW�FRORXUV�DQG�FDSLWDO�OHWWHUV�LQ�)LJXUH�����$��WKH�WUHH�Aloe VSHFLHV���%��Aloe plicatilis� �&��Aloe VHFW��Macrifoliae���'��Haworthia VXEJ��Haworthia� �(��WKH�µWUXH¶�Aloe VSHFLHV���)��WKH�µ+DZRU-WKLRLG¶ FODGH��H. VXEJ��5REXVWLSHGXQFXODUHV࣠+࣠$VWUR�ORED࣠+࣠$ORH�aristata�� �*��Haworthia VXEJ��Hexangulares DQG��+��Gasteria�

The ‘tree aloes’. —�7KH�DQDO\VLV�UHWULHYHV�PRVW�RI�WKH�WUHH�Aloe�VSHFLHV��A. barberae��A. dichotoma��A. eminens��DV�D�FODGH�ZLWK�VWURQJ�VXSSRUW�LQ�WKH�%,�DQDO\VLV��%6� �����33� ��������7KLV�OLQHDJH�FRQVWLWXWHV�WKH�HDUOLHVW�GLYHUJLQJ�HOHPHQWV�LQ�VXEIDPLO\�$ORRLGHDH��)LJ���%�. 7KH�UHPDLQLQJ�WUHH�VSHFLHV��A. plicatilis� RFFXSLHV�DQ�LVRODWHG�SRVLWLRQ�LQ�DQ�XQUHVROYHG�SRO\WRP\�ZLWK�A. VHFW��Macrifoliae DQG�Haworthia VXEJ��Haworthia�

Haworthia subg. Haworthia. —�7KLV�FODGH�LV�ZHOO�VXS-SRUWHG� LQ� ERWK�03� DQG�%,� DQDO\VHV� �%6� � ���� 33� � ������)LJ���%�.�7KH�JUDVV�OLNH�Haworthia blackburniae LV�UHVROYHG�DV�

VLVWHU�WR�D�ZHOO�VXSSRUWHG��%6� �������33� �������EXW�LQWHUQDOO\�XQUHVROYHG�FODGH�LQFOXGLQJ�WKH�RWKHU�PHPEHUV�RI�WKH�VXEJHQXV�

The ‘rambling aloes’. —�7KLV�JURXS��UHSUHVHQWLQJ�Aloe VHFW��Macrifoliae (A. ciliaris� A. commixta��A. gracilis� A. stria-tula��A. tenuior��ZDV�UHFRYHUHG�LQ�ERWK�DQDO\VHV�DV�D�VWURQJO\�VXSSRUWHG�FODGH��%6� �����33� �������)LJ���%��

The ‘true aloes’. —�7KLV�JURXS��ZKLFK�FRPSULVHV�WKH�PD-MRULW\�RI�Aloe VSHFLHV�SOXV�Chortolirion �)LJ���$��ZDV�UHWULHYHG�ZLWK�VWURQJ�VXSSRUW�LQ�WKH�%,��33� ��������6HYHUDO�LQWHUQDO�FODGHV��QRQH�RI�ZKLFK�FRUUHVSRQG�H[DFWO\�WR�FXUUHQW�VHFWLRQV�UHFRJQLVHG�LQ�WKH�WD[RQRP\�RI�WKH�JHQXV��ZHUH�ZHOO�VXSSRUWHG�LQ�WKH�%,�DQDO\VLV��$PRQJ�WKHVH��WKH�PDMRULW\�RI�WKH�µJUDVV�DORHV¶��A. VHFW��Leptoaloe� JURXSHG�WRJHWKHU��%6� �����33� �������LQ�D�FODGH�WKDW�LQFOXGHG�Chortolirion angolense �%DNHU��$��%HUJHU� A. kouebok-keveldensis� A. buhrii� A. lutescens� A. reynoldsii� A. spicata��A. striata VXEVS. komaggasensis DQG�A. striata VXEVS��striata DOEHLW�ZLWK�ZHDN�%,�VXSSRUW��33� ������� ,Q�WKH�03�WUHH�WKLV�FODGH�ZDV�UHFRYHUHG�DV�SDUW�RI�D�ODUJHU�SRO\WRP\�ZLWK�QR�ERRW-VWUDS�VXSSRUW��$�VHFRQG�FODGH�FRPSULVHG�VHYHUDO�RI�WKH�µVLQJOH�VWHPPHG�DORHV¶��Aloe VHFW��Pachydendron��SOXV�RWKHU�VSHFLHV��ZLWK�ZHDN�03�VXSSRUW��%6� �����EXW�VWURQJ�VXSSRUW�LQ�WKH�%,�DQDO\VLV��33� ��������$�WKLUG�FODGH�FRPSULVLQJ�D�KHWHURJHQRXV�DVVHPEODJH�RI�Aloe VSHFLHV��A. excelsa� A. petricola� A. munchii� A. chabaudii��A. vryheidensis��ZDV�ZHOO�VXSSRUWHG�LQ�WKH�%,��%6� �����33� ��������$�IRXUWK�FODGH�UHFRYHUHG�DQRWKHU�KHWHURJHQRXV�DVVHPEODJH�RI�Aloe VSHFLHV�FRQWDLQLQJ�WKH�W\SH�RI�WKH�JHQXV��A. perfoliata� ZLWK�ZHDN�WR�PRGHUDWH�VXSSRUW��%6� �����33� ��������7ZR�DGGLWLRQDO�µJUDVV�DORHV¶��A. albida DQG�A. chortoliri-oides� ZHUH�UHFRYHUHG�DV�VLVWHUV�ZLWK�VWURQJ�VXSSRUW��%6� �����33� ��������7KH�UHPDLQLQJ�µWUXH�DORHV¶�DUH�ODUJHO\�XQUHVROYHG�

Fig. .�&ODGRJUDPV�VXPPDULVLQJ�UHODWLRQVKLSV�RI�PDMRU�WD[RQRPLF�JURXSLQJV�ZLWKLQ�$ORRLGHDH��1XPEHUV�DERYH�WKH�EUDQFKHV�DUH�03�ERRWVWUDS�VXSSRUW�YDOXHV�DQG�RQHV�EHORZ�DUH�%,�SRVWHULRU�SUREDELOLWLHV�

Gasteria (n = 20)

Haworthia subg. Hexangulares (n = 15)52

6 2

65

92

Aloe haworthioides

H. subg. Robustipedunculares (n = 5)

Astroloba (n = 4)

Aloe sect. Aristatae (n = 3)

Haworthia koelmaniorum (n = 2)

H. subg. Haworthia (n = 49)

True Aloe spp .

Chortolirion angolense

T ree Aloe spp. (n = 5)

Out g rou p s ( n = 13 ) Out g rou p ( Bulbine s p.)

Gasteria (n = 20)

H. subg. Hexangulares (n = 16)

80

0.74

92

-

H. subg. Robusti-

pedunculares (n = 5)

Astroloba (n = 4)

Haworthia koelmaniorum (n = 2)

Chortolirion angolense

Aloe sect. Macrifoliae (n = 5)

T ree Aloe spp. (n = 5)

Out g rou p s (n = 13 )

d e n i b m o C ) C ( 1 S T I ) B ( d i t s a l P ) A (

Aloe sect. Aristatae (n = 3)

H. subg. Haworthia (n = 49)

Aloe sect. Kumara (n = 2)

True Aloe spp. (n = 40)

1.00

1.00

58 0.68

77 0.95

-

56 0.97

1.00

-

0.53

93 1.00

77

99

59

Aloe sect. Macrifoliae (n = 5)

57 88

95

T rue Aloe spp .

Chortolirion angolense

Gasteria (n = 20)

H. subg. Robusti-

pedunculares (n = 5)

H. subg. Hexangulares

Aloe sect. Aristatae (n = 3)

57

61

Astroloba (n = 4)

85 85

61

99

T ree Aloe spp. (n = 5)69

Aloe sect. Kumara (n = 2)

Aloe sect. Kumara (n = 2)

H. subg. Haworthia (n = 44)

��

'DUX��DO����Phylogeny of AlooideaeTAXON����������)HEUXDU\���������±��

��9HUVLRQ�RI�5HFRUG��LGHQWLFDO�WR�SULQW�YHUVLRQ��

Aloe succotrina

Aloe hereroensis

Aloe haworthioides

Aloe anivoranoensis

Aloe dewinteri

Aloe propagulifera

Aloe greatheadii subsp. davyana

Aloe striata subsp. karasbergensis

Aloe glauca

Aloe brevifolia

Aloe albida

Aloe chortolirioides

Aloe melanacantha

Aloe pearsonii

Aloe arenicola

Aloe perfoliata

Aloe vryheidensis

Aloe chabaudii

Aloe munchii

Aloe petricola

Aloe excelsa

Aloe angelica

Aloe alooides

Aloe comosa

Aloe rupestris

Aloe thraskii

Aloe lineata

Aloe lineata var. muirii

Aloe lineata var. muirii

Aloe pictifolia

Aloe microstigma

Aloe arborescens

Aloe ferox

Aloe spicata

Chortolirion angolense

Aloe lutescens

Aloe reynoldsii

Aloe striata

Aloe striata subsp. kommagasensis

Aloe buhrii

Aloe kouebokkeveldensis

Aloe saundersiae

Aloe verecunda

Aloe vossii

Aloe challisii

Aloe fouriei

Aloe ecklonis

Aloe nubigena

H. koelmaniorum var. mcmutryi

H. koelmaniorum var. mcmutryi

Aloe aristata

Aloe aristata

Aloe aristata

Astroloba rubriflora

Astroloba foliolosa

Astroloba herrei

Astroloba corrugata

Haworthia marginata

Haworthia kingiana

Haworthia pumila

Haworthia marginata

Haworthia kingiana

Haworthia attenuata

Haworthia venosa

Haworthia venosa var. granulata

Haworthia limifolia

Haworthia limifolia

Haworthia limifolia var. ubomboensis

Haworthia fasciata

Haworthia longiana

Haworthia sordida

Haworthia bruynsii

Haworthia glauca

Haworthia nigra

Haworthia reinwardtii

Haworthia coarctata var. adelaidensis

Haworthia reinwardtii var. brevicula

Haworthia coarctata

G. pillansii var. pillansii

G. pillansii var. ernesti-ruschii

G. croucheri

G. carinata var. carinata

G. carinata var. retusa

G. acinacifolia

G. nitida var. armstrongii

G. batesiana var. dolomitica

G. ellaphieae

G. excelsa

G. vlokii

G. obliqua

G. rawlinsonii

G. glauca

G. glomerata

G. pulchra

G. tukhelensis

G. disticha

G. doreeniae

G. polita

[Fig. 3B, next page]

0.97

1.00

0.73

1.00

0.60

1.00

0.92

0.73

0.99

0.93

0.90

0.89

1.00

1.00 1.00

0.99

1.00

0.92

0.51

1.00

0.60

0.591.00

0.77

1.00

1.00

0.93

1.00

0.68

1.00

0.95

1.00

0.74

0.99

0.97

1.00

1.00

0.67

1.00

0.64

0.75

1.00

1.00

0.88

1.00

0.78

1.00

0.55

0.98

0.55

0.99

0.74

1.00

1.00

0.99

0.93

1.00

0.57

0.95

0.83

0.98

0.77

56(3)

í(3)

80(5)

80

í(0)

90(3)

53(1)í(0)

53(1)84(1)

51(1)66(1)

61(2)

71(4) 70(1)

55(0)

86(2)

í(0)í(0)

91(2)

í(0)

í(0) 95(2)í

85(2)

89(3)

61(1)

92(6)

58(1)

99(6)

77(1)

100(11)

í(1)

79(2)75(2)

100(1)

100(10)

í(0)

80(1)

í(1)

í(1)82(2)

81(5)67(1)

57(2)í(1)

78(2)í

54(3)

í

í(3)

97(4)

88

70(2)

92(3)

61(1)

63(1)

58(1)

í

Gasteria

Haworthia subg.Hexangulares I

Haworthia subg.Robustipedunculares

Astroloba

Aloe sect. Aristatae

Haworthia sect. Hexangulares II

Aloe sect.Leptoaloe

True Aloe spp.

E

F

G

H

(1)

(1)

(1)

Fig. . 7KH�����PDMRULW\�UXOH�FRQVHQVXV�WUHH�REWDLQHG�IURP�WKH�%,�DQDO\VLV�RI�WKH�FRPELQHG�SODV-WLG�DQG�,76��GDWDVHW�IRU�$OR�RLGHDH�SOXV�RXWJURXSV��:H�PDSSHG�WKH�ERRWVWUDS�YDOXHV�IURP�WKH�03�WUHH�RQWR�WKH�%,�PDMRULW\�UXOH�FRQVHQVXV�WUHH�EHFDXVH�WKH�03�DQG�%,�DQDO\VHV�SURGXFHG�VLPLODU�WRSRORJLHV��1XPEHUV�DERYH�QRGHV�DUH�QRQSDUDPHWULF�03�ERRWVWUDS�VXSSRUW�YDOXHV��WKRVH�DERYH�QRGHV�EXW�ZLWKLQ�SDUHQWKHVHV�DUH�GHFD\�LQGLFHV�FDOFXODWHG�XVLQJ�7UHH5RW��6RUHQVHQ��)UDQ]RVD���������WKRVH�EHORZ�QRGHV�DUH�%,�SRVWH-ULRU�SUREDELOLWLHV�DERYH�����

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‘Haworthioid’ clade (Haworthia subg. Robustipeduncu-lares, Astroloba, Aloe aristata, H. koelmaniorum). —�$�PRG-HUDWHO\�VXSSRUWHG�µKDZRUWKLRLG¶�FODGH��%6� �����33� �������ZDV�UHWULHYHG�DV�VLVWHU�WR�WKH�Gasteria-Hexangulares FODGH�ZLWK�VWURQJ�VXSSRUW��%6� �����33� ��������Haworthia koelmaniorum YDU��mcmurtryi (H. VXEJ��Hexangulares��ZDV�UHWULHYHG�ZLWK�ZHDN�03�DQG�%,�VXSSRUW��%6� �����33� �������DV�WKH�HDUOLHVW�GLYHUJLQJ�OLQHDJH�LQ�WKH�FODGH��7KHUH�LV�PRGHUDWH�WR�VWURQJ�VXSSRUW��%6� �����33� �������IRU�D�VLVWHU�UHODWLRQVKLS�EHWZHHQ�

A. aristata DQG�H. VXEJ��5REXVWLSHGXQFXODUHV࣠�Astroloba��7KH�UHPDLQLQJ�VSHFLHV�RI�H. VXEJ��Robustipedunculares ZHUH�UH-FRYHUHG�ZLWK�VWURQJ�VXSSRUW��%6� ������33� �������EXW�DUH�RQO\�ZHDNO\�VXSSRUWHG�LQ�WKH�%,��33� �������DV�VLVWHU�WR�D�PRGHUDWH�WR�ZHOO�VXSSRUWHG��%6� �����33� �������Astroloba.

Haworthia subg. Hexangulares. — Haworthia� VXEJ��Hexangulares� �EXW� H[FOXGLQJ�H. koelmaniorum�� )LJ�� �$��ZDV�UHFRYHUHG�LQ�ERWK�03�DQG�%,�DQDO\VHV�ZLWK�PRGHUDWH�WR�VWURQJ�VXSSRUW��%6� �����33� �������LQ�D�FODGH�ZLWK�Gasteria��

Anthericum liliago

Tecophilaea cyanocrocus

Zephyra elegans

Xanthorrhoea resinosa

Xanthorrhoea sp.

Xanthorrhoea sp.

Asphodeline lutea

Eremurus spectabilis

Kniphofia galpinii

Kniphofia uvaria

Bulbine fistulosa

Bulbine semibarbata

Bulbine frutescens

Aloe barberae

Aloe eminens

Aloe dichotoma subsp. ramosissima

Aloe dichotoma

Aloe pillansii

Aloe plicatilis

Aloe plicatilis

Aloe commixta

Aloe gracilis

Aloe striatula

Aloe ciliaris

Aloe tenuior

Haworthia blackburniae

Haworthia blackburniae var. derustensis

Haworthia blackburniae var. graminifolia

Haworthia mirabilis

Haworthia bayeri

Haworthia zantneriana

Haworthia angustifolia

Haworthia emelyae var. comptoniana

Haworthia truncata

Haworthia variegata

Haworthia pygmaea

Haworthia chloracantha

Haworthia outeniquensis

Haworthia decipiens

Haworthia arachnoidea

Haworthia arachnoidea

Haworthia mucronata var. bijliana

Haworthia mucronata var. morrisiae

Haworthia mucronata var. rycroftiana

Haworthia monticola var. asema

Haworthia pulchella var. globifera

Haworthia lockwoodii

Haworthia mucronata var. habdomadis

Haworthia cooperi

Haworthia cymbiformis var. setulifera

Haworthia cymbiformis var. ramosa

Haworthia decipiens var. virella

Haworthia decipiens var. pringlei

Haworthia cooperi var. doldii

Haworthia cymbiformis

Haworthia cymbiformis var. setulifera

Haworthia vlokii

Haworthia wittebergensis

Haworthia emelyae

Haworthia floribunda

Haworthia marxii

Haworthia semiviva

Haworthia marumiana var. archeri

Haworthia mutica

Haworthia mutica var. nigra

Haworthia retusa

Haworthia mirabilis var. triebneriana

Haworthia mirabilis var. calcarea

Haworthia mirabilis var. paradoxa

Haworthia magnifica var. dekenahii

Haworthia herbacea var. flaccida

Haworthia maculata

Haworthia reticulata

Haworthia herbacea var. paynei

[Fig. 3A, previous page]

1.00

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0.92

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0.51

0.90

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0.53

1.00

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0.94

0.96

0.99

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0.88

0.82

0.76

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Haworthia subg.Haworthia

Aloe sect. Macrifoliae

Aloe sect. Kumara

Tree Aloe spp.

Outgroups

AB

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5HODWLRQVKLSV�DPRQJ�H. attenuata��WKH�UHVW�RI�H. VXEJ��Hexan-gulares�DQG�Gasteria�UHPDLQ�XQUHVROYHG�

Gasteria. — Gasteria�VSHFLHV�IRUP�D�ZHOO�VXSSRUWHG�FODGH�LQ�ERWK�DQDO\VHV��%6� �����33� �������)LJ���$���7KH�FODGH�LV�UHFRYHUHG�LQ�DOO�WUHHV�DV�RQH�HOHPHQW�RI�D�PRGHUDWHO\�WR�ZHOO�VXSSRUWHG�WULFKRWRP\��%6� �����33� �������WKDW�LQFOXGHV�WKH�VSHFLHV�RI�Haworthia VXEJ��Hexangulares.

Evolution of morphological traits. —�$OO����PRUSKR-ORJLFDO�WUDLWV�WKDW�ZHUH�DQDO\VHG�DUH�VKRZQ�WR�EH�KRPRSODVLRXV��QDPHO\�KDELW��OHDI�LQVHUWLRQ��OHDI�PDUJLQV��OHDI�PDFXODWLRQ�DQG�WXEHUFXODWLRQ��OHDI�DSH[��LQIORUHVFHQFH�EUDQFKLQJ�DQG�RULHQWD-WLRQ��LQIORUHVFHQFH�OHQJWK�UHODWLYH�WR�SHGLFHO��LQIORUHVFHQFH�VKDSH�DQG�FRORXU��SHULDQWK�RULHQWDWLRQ�DW�DQWKHVLV��IORZHU�DUUDQJHPHQW�RQ�SHGXQFOH��SHULDQWK�V\PPHWU\�DQG�FRORXU��SHULDQWK�VKDSH�DQG�FXUYDWXUH��SHGLFHO�OHQJWK��WHSDO�FRQQDWLRQ��DQG�VWDPHQ�OHQJWK��$SSHQGL[�6����7KH�GLVWULEXWLRQ�RI�QLQH�FKDUDFWHUV�RI�SDUWLFX-ODU�LPSRUWDQFH�LQ�JHQHULF�DQG�LQIUDJHQHULF�FODVVLILFDWLRQ�ZLWKLQ�$ORRLGHDH�LV�VXPPDULVHG�LQ�7DEOH����VHH�DOVR�)LJV��6�±6���

Habit. —�$FDXOHVFHQFH�LV�SOHVLRPRUSKLF�ZLWKLQ�$ORRLGHDH��ZLWK�FHUWDLQ�OLQHDJHV�FKDUDFWHULVHG�E\�VHFRQGDU\�FDXOHVFHQFH�DQG� DUERUHVFHQFH� �)LJ�� 6�$��� &DXOHVFHQFH� KDV� GHYHORSHG�VHYHUDO� WLPHV� LQ� WKH�VXEIDPLO\�� LQFOXGLQJ�Aloe� LWVHOI� �QRWD-EO\�A. VHFW��Macrifoliae��DQG�LQ�VRPH�RI�WKH�VHJUHJDWH�JHQHUD��QRWDEO\�Astroloba DQG�VRPH�VSHFLHV�RI�Gasteria�DQG�Haworthia VXEJ��Hexangulares��$UERUHVFHQFH�LV�XQFRPPRQ�LQ�WKH�VXE-IDPLO\�DQG�LV�FKDUDFWHULVWLF�RI�VSHFLHV�LQ�Aloe VHFW��Aloidendron DQG�A. VHFW��Dracoaloe� LQFOXGLQJ�A. eminens �)LJ��6�$��

Leaf insertion. — 3RO\VWLFKRXV�OHDI�LQVHUWLRQ�LV�FRPPRQ�ZLWKLQ�WKH�VXEIDPLO\�EXW�GLVWLFKRXV�OHDI�LQVHUWLRQ�LV�UDUH��RF-FXUULQJ�VHYHUDO�WLPHV�LQ�Aloe��RQFH�LQ�Haworthia�DQG�SRVVLEO\�RQFH�LQ�Gasteria��)LJ��6�%��

Leaf tuberculation. — 7KH�SUHVHQFH�RI�ZKLWH�WXEHUFOHV�RQ�WKH�OHDYHV�LV�ODUJHO\�UHVWULFWHG�WR�Gasteria DQG�Haworthia�VXEJ��Hexangulares�DQG�VXEJ��Robustipedunculares�EXW�WXEHUFXOHV�DUH�DOVR�GHYHORSHG�LQ�VRPH�Aloe VSHFLHV��H�J���Aloe aristata� A. haworthioides��A. verecunda��DQG�LQ�Chortolirion��)LJ��6�&��

Table .�&KDUDFWHU�VWDWHV�IRU�QLQH�VHOHFWHG�WUDLWV�ZKLFK�KDYH�EHHQ�HPSOR\HG�LQ�SUHYLRXV�$ORRLGHDH�V\VWHPDWLFV�

&KDUDFWHU &KDUDFWHU�VWDWH 6SHFLHV

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6WHPOHVV 0RVW�Gasteria� KDOI�RI�Haworthia�VXEJ��Hexangulares� H. VXEJ��Robustipedunculares

&DXOHVFHQW Gasteria obliqua��KDOI�RI�Haworthia�VXEJ��Hexangulares� DOO�Astroloba �LQFOXGLQJ�Poellnitzia)

$UERUHVFHQW Aloe plicatilis (A. VHFW��Kumara���A. VHFW��Pachydendron DQG� A. VHFW��Dracoaloe

/HDI�LQVHUWLRQ6SLUDO�SRO\VWLFKRXV� 0RVW�$ORRLGHDH�

'LVWLFKRXV Gasteria obliqua� Aloe nubigena� A. plicatilis� A. verecunda DQG�Haworthia truncata

/HDI�PDFXODWLRQ�,PPDFXODWH� 0RVW�$ORRLGHDH

0DFXODWH $OO Gasteria� VHYHUDO�Aloe VSHFLHV��H�J���A. aristata DQG A. VHFW�� Leptoaloe

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6SUHDGLQJ Chortolirion� Astroloba��Haworthia DQG�D�IHZ�Aloe

3HQGXORXV 0RVW�Aloe DQG�Gasteria VSHFLHV

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$FWLQRPRUSKLF� Haworthia�VXEJ��Robustipedunculares DQG�VHYHUDO�Aloe� H�J���A. albida

:HDNO\�ELODELDWH� Gasteria VSHFLHV

6WURQJO\�ELODELDWH Haworthia VXEJ��Haworthia DQG�VXEJ��Hexangulares SOXV�Chortolirion angolense

3HULDQWK�FXUYDWXUH6WUDLJKW� Astroloba��Chortolirion��Haworthia VXEJ��Hexangulares DQG�VXEJ��

Robustipedunculares DQG�PDMRULW\�RI�Aloe

&XUYHG� Gasteria��Haworthia VXEJ��Haworthia

3HULDQWK&\OLQGULFDO�DQG�WDSHULQJ�LQWR�SHGLFHO� 0RVW�Aloe� Astroloba� Chortolirion� Haworthia VXEJ��Haworthia DQG�

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Perianth colour. —�)URP�RXU�DQDO\VLV��QRW�VKRZQ���YLYLG�\HOORZ��RUDQJH�RU�UHGGLVK�IORZHUV�DUH�FOHDUO\�SOHVLRPRUSKLF�ZLWKLQ� $ORRLGHDH�� +RZHYHU��Chortolirion��Haworthia DQG�Astro loba �H[FOXGLQJ�Poellnitzia��DQG�VHYHUDO�VSHFLHV�RI�Aloe��HVSHFLDOO\�LQ�A. VHFW��Leptoaloe��DUH�FKDUDFWHULVHG�E\�ZKLWLVK�RU�JUHHQLVK�SHULDQWK�

The Haworthia-type flower. —�8VXDOO\�ZKLWH�DQG�VRPH-WLPHV�ZLWK�VWURQJO\�]\JRPRUSKLF�SHULDQWKV��DQG�LQVHUWHG�DQ-WKHUV�DUH�FKDUDFWHULVWLF�RI�WKH�JHQXV�Haworthia DQG�Chorto-lirion��&KDUDFWHU�UHFRQVWUXFWLRQ�VXJJHVWV�WKDW�WKLV�IORZHU�W\SH�LV�KRPRSODVLRXV��DQG�WKDW�LW�KDV�HYROYHG�LQGHSHQGHQWO\�DW�OHDVW�WKUHH�DQG�SRVVLEO\�IRXU�WLPHV��RQFH�HDFK�LQ�Chortolirion DQG�H. VXEJ� Haworthia�DQG�RQFH�RU�WZLFH�LQ�H. VXEJ��Hexangu-lares�VXEJ��Robustipedunculares ZLWK�SRVVLEOH�UHYHUVDOV� LQ�A. aristata��A. haworthioides DQG�Gasteria���)LJ��6�$±&��

The Gasteria-type flower. —�)ODVN�VKDSHG�IORZHUV�ZLWK�LQIODWHG�EDVHV�RFFXU�QRW�RQO\�LQ�Gasteria� EXW�DOVR�LQ�PDQ\�Aloe VSHFLHV�DQG�KDYH�HYLGHQWO\�HYROYHG�VHYHUDO�WLPHV��)LJ��6�$���DOWKRXJK�ZLWKLQ�Aloe�� WKH�VWDPHQV�DUH�RIWHQ�EXW�QRW�DOZD\V�H[VHUWHG��)LJ��6�%���6LPLODU�IORZHUV�DUH�FKDUDFWHULVWLF�DOVR�RI�Astro loba �Ł�Poellnitzia) rubriflora�EXW�KHUH�DUH�XQLTXHO\�KHOG�HUHFW�RQ�DQ�LQFOLQHG�UDFHPH��$�ZHOO�GHYHORSHG�SHULDQWK�WXEH�IRUPHG�E\�WKH�IXVLRQ�RI�ERWK�WHSDO�ZKRUOV�KDV�HYROYHG�LQGH-SHQGHQWO\�LQ�Gasteria��Astroloba�DQG�Aloe kouebokkeveldensis �)LJ��6�&��

Tepal connation. —�7KH�SHULDQWK�LQ�$ORRLGHDH�FRPSULVHV�VL[�WHSDOV�LQ�WZR�ZKRUOV��YDULRXVO\�FRQQDWH�LQWR�D�VKRUW�RU�SURP-LQHQW� WXEH��%DVDOO\�FRQQDWH�WHSDOV�UHSUHVHQW� WKH�SOHVLRPRU-SKLF�FRQGLWLRQ��,Q�VHYHUDO�RI�WKH�RUQLWKRSKLORXV�Aloe�VSHFLHV��KRZHYHU��WKH�RXWHU�WHSDO�ZKRUO�LV�FRQQDWH�LQ�WKH�EDVDO�KDOI��IRUPLQJ�D�GLVWLQFW�SHULDQWK�WXEH��)LJ� 6�&���)ORZHUV�ZLWK�ERWK�ZKRUOV�FRQQDWH�IRU�KDOI�RU�PRUH�RI�WKHLU�OHQJWK�DUH�GLDJQRVWLF�IRU�Gasteria��Astroloba (Poellnitzia) rubriflora��DQG�A. koue-bokkeveldensis�DQG�DSSHDU�WR�KDYH�HYROYHG�LQGHSHQGHQWO\�LQ�WKHVH�WKUHH�OLQHDJHV�IURP�DQFHVWUDO�W\SHV�ZLWK�WHSDOV�FRQQDWH�DW�WKH�EDVH�RQO\��)LJ��6�&)�

DISCUSSION

Relationships within Alooideae. —�7KH�PRQRSK\O\�RI�$ORRLGHDH�ZDV�ILUVW�GHPRQVWUDWHG�E\�&KDVH��DO���������XVLQJ�WKH�WZR�SODVWLG�UHJLRQV�rbcL�DQG�trnL-F� DQG�PRUH�UHFHQWO\�E\�7UHXWOHLQ��DO� �����D��E���XVLQJ�matK�DQG�JHQRPLF�ILQJHU-SULQWLQJ��7KLV�ILQGLQJ�LV�VWURQJO\�VXSSRUWHG�E\�RXU�VWXG\��XVLQJ�VLJQLILFDQWO\�JUHDWHU�WD[RQ�VDPSOLQJ�DQG�D�FRPELQHG�DQDO\VLV�RI�WKUHH�SODVWLG�UHJLRQV��matK� rbcLa� trnH-psbA��SOXV�WKH�QXFOHDU�UHJLRQ�,76���$OWKRXJK�WKH�PRQRSK\O\�RI�WKH�VXEIDP-LO\�LV�QR�ORQJHU�LQ�TXHVWLRQ��WKLV�FDQQRW�EH�DFFHSWHG�IRU�DOO�RI�WKH�JHQHUD��,W�KDV�EHFRPH�LQFUHDVLQJO\�FOHDU�WKDW�QHLWKHU�Aloe QRU�Haworthia DUH�PRQRSK\OHWLF�DV�FXUUHQWO\�FLUFXPVFULEHG��7UHXWOHLQ��DO�������D��E��5DPGKDQL��DO����������2XU�DQDO\VLV�RI�WKH�FRPELQHG�SODVWLG�DQG�QXFOHDU�'1$�GDWDVHWV��)LJ�����FRQ-ILUPV�WKDW�ERWK�Aloe DQG�Haworthia DUH�SRO\SK\OHWLF�DV�FXU-UHQWO\�FLUFXPVFULEHG��:H�LGHQWLI\�HLJKW�SULPDU\�PRQRSK\OHWLF�OLQHDJHV�LQ�$ORRLGHDH�WKDW�ODUJHO\�FRUUHVSRQG�WR�WKH�IROORZLQJ�FXUUHQWO\�UHFRJQLVHG�JHQHULF�DQG�LQIUDJHQHULF�JURXSV������Aloe

VHFW��Dracoaloe�A. VHFW��Aloidendron������A. VHFW��Kumara������A��VHFW��Macrifoliae������Haworthia�VXEJ��Haworthia������A. VHFW� Aristatae�H.�VXEJ��Robustipedunculares�Astroloba��(6) H��VXEJ��Hexangulares������Gasteria������UHPDLQLQJ�VSH-FLHV�RI�Aloe��7KLV�WRSRORJ\�LV�FRQJUXHQW�ZLWK�WKDW�RI�7UHXWOHLQ��DO�������E���ZKLFK�KDG�PXFK�UHGXFHG�VDPSOLQJ��$PRQJ�WKH�VPDOOHU�VHJUHJDWH�JHQHUD��ERWK�Astroloba DQG�Gasteria�DUH�PRQRSK\OHWLF�EXW�Chortolirion� LV�GHHSO\�HPEHGGHG�DPRQJ�WUXH�Aloe�VSHFLHV�

6HYHQ�RI�WKH�HLJKW�SULPDU\�OLQHDJHV�DUH�HQGHPLF�WR�VRXWK-HUQ�$IULFD��ZLWK�RQO\�Aloe V�VWU��H[WHQGLQJ�EH\RQG�WKH�UHJLRQ��7KLV�VXJJHVWV�WKDW�WKH�HDUO\�GLYHUVLILFDWLRQ�RI�WKH�VXEIDPLO\�WRRN�SODFH�LQ�WKH�VXEFRQWLQHQW��ZKLFK�LV�DOVR�WKH�PDLQ�FHQWUH�RI�GLVWULEXWLRQ�IRU�WKH�VXEIDPLO\��6PLWK��9DQ�:\N��������

Relationships of the ‘tree aloes’. —�2XU�DQDO\VLV�GRHV�QRW�UHWUHLYH�WKH�µWUHH�DORHV¶�VHQVX�9DQ�:\N��6PLWK��������DV�D�PRQRSK\OHWLF�JURXS�EXW�DV�WZR�VHSDUDWH�OLQHDJHV��Aloe VHFW��$ORLGHQGURQ࣠�A. VHFW��Dracoaloe��LQFOXGLQJ�A. eminens IURP�6RPDOLD��HPHUJHV�DV�SRVVLEO\�RQH�RI�WKH�HDUO\�GLYHUJLQJ�OLQH-DJHV�ZLWKLQ�WKH�VXEIDPLO\��)LJ���%���7KH�LPPHGLDWH�UHODWLRQ-VKLSV�RI�WKH�UHPDLQLQJ�WUHH�Aloe��A. plicatilis (A. VHFW��Kumara) DUH�XQUHVROYHG�

Relationships of Aloe sect. Macrifoliae. — Aloe VHFW��Macri foliae� WKH�µUDPEOLQJ�DORHV¶��VHQVX�9DQ�:\N��6PLWK��������� FRPSULVLQJ� ILYH� FORVHO\� UHODWHG� VSHFLHV� �A. ciliaris� A. commixta��A. gracilis� A. striatula��A. tenuior���LV�UHFRYHUHG�DV�D�VWURQJO\�VXSSRUWHG�FODGH�EXW�LWV�UHODWLRQVKLS�WR�WKH�UHPDLQ-LQJ�VSHFLHV�LQ�WKH�VXEIDPLO\�UHPDLQV�XQUHVROYHG��7KH�VHFWLRQ�LV�GHILQHG�YHJHWDWLYHO\�E\�LWV�FDQH�OLNH�VWHPV��DQG�VOHQGHU��VKHDWK-LQJ��XQVSRWWHG�DQG�PHVRSK\WLF�OHDYHV�ZLWK�PLQXWH�PDUJLQDO�WHHWK��DQG�IORUDOO\�E\�WKH�PRUH�RU�OHVV�HQWLUHO\�FRQQDWH�RXWHU�WHSDOV��*OHQ��+DUG\��������

Relationships within the ‘true aloes’. —�7KH�UHPDLQLQJ�VSHFLHV�RI�Aloe �H[FOXGLQJ�A. aristata��DUH�UHWULHYHG�DV�D�FODGH�EXW�UHODWLRQVKLSV�DPRQJ�WKHP�DUH�SRRUO\�UHVROYHG��$OWKRXJK�VRPH�RI�WKH�FXUUHQWO\�UHFRJQLVHG�VHFWLRQV�PD\�EH�PRQRSK\-OHWLF��RWKHUV�DUH�QRW��DQG�D�PXFK�PRUH�H[WHQVLYH�VDPSOLQJ�RI�VSHFLHV�DQG�DGGLWLRQDO�JHQH�UHJLRQV�LV�UHTXLUHG�WR�HYDOXDWH�WD[R-QRPLF�DQG�HYROXWLRQDU\�UHODWLRQVKLSV�DPRQJ�WKHP�

$OWKRXJK�Chortolirion� LV�GHHSO\�HPEHGGHG�ZLWKLQ�WKLV�JURXS��DV�SDUW�RI�D�FODGH�LQFOXGLQJ�PRVW�RI�WKH�JUDVV�Aloe VSH-FLHV��LWV�SUHFLVH�UHODWLRQVKLSV�DUH�VWLOO�XQFOHDU��&ORVH�PRUSKRORJ-LFDO�VLPLODULW\�WR�VSHFLHV�VXFK�DV�A. bowiea�DQG�A. inconspicua LQ�YHJHWDWLYH�SDUWV��QDPHO\�WKH�JUDVV�OLNH�OHDYHV�ZLWK�EXOE�OLNH�VZHOOLQJ��DQG�LQ�WKH�VPDOO��ELODELDWH�IORZHUV��VXJJHVW�D�FORVH�UHODWLRQVKLS�WR�SDUW�RI�A. VHFW��Leptoaloe�

Relationships of Gasteria. — Gasteria�IRUPV�D�VWURQJO\�VXSSRUWHG�FODGH�VLVWHU�WR�Haworthia�VXEJ��Hexangulares. ,W�LV�GHILQHG�E\�VHYHUDO�PRUSKRORJLFDO�V\QDSRPRUSKLHV��QRWDEO\�WKH�XQDUPHG��YHUUXFRVH�OHDYHV�DQG�LQFOLQHG�VHFXQG�LQIORUHVFHQFHV�RI�SHQGXORXV��JDVWHULIRUP�IORZHUV�ZLWK�D�ZHOO�GHYHORSHG�IORUDO�WXEH��9DQ�-DDUVYHOG���������DQG�ZDV�XQVXUSULVLQJO\�RQH�RI�WKH�HDUOLHVW�VHJUHJDWHV�RI�Aloe WR�EH�UHFRJQLVHG��2XU�SK\OR-JHQHWLF�DQDO\VLV�SODFHV�WKH�JHQXV�VLVWHU�WR�Haworthia VXEJ��Hexangulares��ZKLFK�LQFOXGHV�VSHFLHV�ZLWK�UHPDUNDEO\�VLPLODU�OHDYHV��7KH�XQLTXH�Gasteria�W\SH�IORZHUV�DUH�PRVW�SDULVPRQL-RXVO\�LQWHUSUHWHG�DV�D�UHYHUVLRQ�WR�ELUG�SROOLQDWLRQ�IURP�WKH�

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HQWRPRSKLORXV�Haworthia�W\SH�IORZHU�ZLWK�LQFOXGHG�VWDPHQV��,W�LV�KDUGO\�VXUSULVLQJ�WKHQ�WKDW�Gasteria IORZHUV�DUH�QRW�SUH-FLVHO\�PDWFKHG�E\�DQ\�ELUG�SROOLQDWHG�IORZHUV�LQ�Aloe�

Relationships of Astroloba. —�6SHFLHV�RI�Astroloba DUH�UHWULHYHG�DV�D�FODGH�VLVWHU�WR�Haworthia VXEJ��Robustipeduncu-lares��7KH�JHQXV�LV�PRUSKRORJLFDOO\�GHILQHG�E\�LWV�FDXOHVFHQW�KDELW�ZLWK�VWLII�� LPEULFDWH�OHDYHV��DQG�VPDOO��DFWLQRPRUSKLF�IORZHUV�ZLWK�LQFOXGHG�VWDPHQV�

Relationships within Haworthia. —�2XU�DQDO\VLV�DFFRUGV�ZLWK�SUHYLRXV�VWXGLHV�LQGLFDWLQJ�WKDW�Haworthia�LV�QRW�PRQR-SK\OHWLF�EXW�UDWKHU�UHSUHVHQWV�WKUHH�OLQHDJHV�FRUUHVSRQGLQJ�WR�WKH�WKUHH�VXEJHQHUD�SURSRVHG�E\�%D\HU���������������6SHFLHV�RI�H. VXEJ��Haworthia�FRPSULVH�D�VWURQJO\�VXSSRUWHG�FODGH��GHILQHG�PRUSKRORJLFDOO\�E\�WKH�EDVDOO\�WULDQJXODU�SHULDQWK��REFODYDWH� IORZHUV�DQG�XSFXUYHG�VW\OH� �%D\HU���������������$GGLWLRQDO�VXSSRUW�IRU�WKLV�DOOLDQFH�FRPHV�IURP�6PLWK��DO����������ZKR�UHSRUWHG�WKH�RFFXUUHQFH�RI�KH[RVH�SRRU�QHFWDU��OHVV�WKDQ�����VXFURVH�HTXLYDOHQWV��LQ�H. VXEJ��Haworthia LQ� FRQWUDVW� WR� KH[RVH�ULFK� QHFWDU� �PRUH� WKDQ� ���� VXFURVH�HTXLYDOHQWV��LQ�H. VXEJ��Hexangulares DQG�H. VXEJ��Robusti-pedunculares. 7KH�ODWWHU�LV�D�VPDOO�JURXS�RI�IRXU�VSHFLHV�WKDW�LV�ZHOO�VXSSRUWHG�DV�PRQRSK\OHWLF�DQG�VLVWHU�WR�Astroloba��,W�LV�GHILQHG�PRUSKRORJLFDOO\�E\�LWV�PRUH�RU�OHVV�VWUDLJKW�SHULDQWK�DEUXSWO\�MRLQHG�WR�WKH�SHGLFHO��%D\HU���������������7KH�IORZHU�W\SH�IRXQG�LQ�H. VXEJ��Robustipedunculares LV�QRW�GLVVLPLODU�WR�WKDW�LQ�Astroloba��GLIIHULQJ�HVVHQWLDOO\�E\�LWV�VOLJKW�]\JRPRUSK\��0HPEHUV�RI�H. VXEJ��Robustipedunculares�DUH�RIWHQ�UREXVW�ZLWK�DWWHQXDWH�OHDYHV��RIWHQ�VFDEULG�DQG�SDWWHUQHG�ZLWK�ZKLWH�WXEHUFOHV��%D\HU���������$�VLVWHU�UHODWLRQVKLS�EHWZHHQ�Astroloba DQG�H. VXEJ��Robustipedunculares�LV�VXSSRUWHG�E\�VLPLODULWLHV�LQ�QHFWDU�VXFURVH�FRQFHQWUDWLRQV��9DQ�:\N��DO�� ������

7KH�GZDUI�Aloe aristata LV VLVWHU�WR�H. VXEJ��Robustipe-dunculares�Astroloba��7KLV�PRUSKRORJLFDOO\�XQXVXDO�VSH-FLHV�LV�XQLTXH�LQ�Aloe LQ�KDYLQJ�³Haworthia-OLNH´ OHDYHV�ZLWK�GU\��DZQ�WLSSHG�DSLFHV�DQG�ZKLWH�WXEHUFOHV�DQG�GLVWLQFWLYH��GRZQFXUYHG�IORZHUV�ZLWK�EDVDO�VZHOOLQJ��*OHQ��+DUG\���������7KH�FORVH�YHJHWDWLYH�VLPLODULW\�EHWZHHQ�WKLV�VSHFLHV�DQG�VRPH�PHPEHUV�RI�H. VXEJ��Robustipedunculares LV�FRQVLVWHQW�ZLWK�D�FORVH�UHODWLRQVKLS�EHWZHHQ�WKHP��DQG�WKH�SULPDU\�GLIIHUHQFH�EHWZHHQ�WKH�WZR�JURXSV�LV�HYLGHQWO\�WKH�ODUJH��RUDQJH�IORZHUV�RI�A. aristata��7KLV�IORZHU�W\SH�LV�DVVRFLDWHG�ZLWK�ELUG�SROOLQD-WLRQ�DQG�KDV�DULVHQ�VHYHUDO�WLPHV�ZLWKLQ�WKH�VXEIDPLO\��$ORQH��LW�LV�WKHUHIRUH�QRW�QHFHVVDULO\�DQ�LQGLFDWLRQ�RI�UHODWLRQVKLSV�

8QOLNH� WKH� RWKHU� WZR� VXEJHQHUD� RI�Haworthia�� VXEJ��Hexangulares LV�SRVVLEO\�SRO\SK\OHWLF�ZLWK�WKH�LQFOXVLRQ�RI�H. koelmaniorum��ZKLFK�RFFXSLHV�DQ�LVRODWHG�SRVLWLRQ�VLVWHU�WR�H. VXEJ��Robustipedunculares�$VWURORED࣠�Aloe aristata EXW�ZLWK�RQO\�ZHDN�VXSSRUW.�7KH�JHRJUDSKLFDO�GLVWULEXWLRQ�RI�WKLV�VSHFLHV�LV�ZHOO�QRUWK�RI�PRVW�RWKHU�VSHFLHV�RI�Haworthia ZLWK�WKH�H[FHSWLRQ�RI�H. limifolia��:KHQ�ILUVW�GHVFULEHG��2EHUPH\HU���������H. koelmaniorum ZDV�WUHDWHG�LQ�H. VHFW��Margariti-fera �QRZ�H. VXEJ��Robustipedunculares��EXW�ODWHU�WUDQVIHUUHG�WR�H. VXEJ��Hexangulares��ZKHUH�LWV�UHODWLRQVKLSV�DSSHDU�WR�OLH�ZLWK�H. limifolia �%D\HU�������. )XUWKHU�HYLGHQFH�IRU�D�ILQDO�GHFLVLRQ�RQ�LWV�SRVLWLRQ�LV�UHTXLUHG��6SHFLHV�RI�H. VXEJ��Hex-angulares�GLVSOD\�WKH�ODUJHVW�YHJHWDWLYH�GLYHUVLW\�LQ�Haworthia��ZLWK�VRPH�VSHFLHV�FORVHO\�UHVHPEOLQJ�PHPEHUV�RI�Astroloba

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Evolution of selected characters in Alooideae. —�2XU�UHFRQVWUXFWLRQ�RI�WKH�QLQH�PRUSKRORJLFDO�FKDUDFWHUV�RQWR�WKH�%,�PDMRULW\�UXOH�FRQVHQVXV�WUHH�UHYHDOHG�WKDW�QRQH�LV�XQLTXH�WR�D�VLQJOH�FODGH�LGHQWLILHG�LQ�WKLV�VWXG\�

Habit. —�3UHYLRXV FODVVLILFDWLRQV��H�J���5H\QROGV��������+ROODQG��������DUH�LPSOLFLW�LQ�WUHDWLQJ�DUERUHVFHQFH�LQ�Aloe DV�D�GHULYHG�VWDWH��%UDQGKDP��������SURSRVHG�WKDW�VFDQGHQW�Aloe VSHFLHV�ZLWK�XVXDOO\�UHODWLYHO\�PHVRSK\WLF�OHDYHV��H�J���A��tenu-ior�DQG�A. ciliaris��$��VHFW��Macrifoliae���UHSUHVHQW�WKH�SULPLWLYH�VWDWH�LQ�Aloe VSHFLHV�EXW�6PLWK��9DQ�:\N��������DUJXHG�WKDW�ERWK�VPDOO��KLJKO\�VXFFXOHQW�WD[D�DQG�DUERUHVFHQW�IRUPV�ZHUH�GHULYHG�IURP�D�PHVRSK\WLF��FRPSDUDWLYHO\�DFDXOHVFHQW�WD[RQ��2XU�DQDO\VLV�VXSSRUWV�WKLV�K\SRWKHVLV��LQGLFDWLQJ�WKDW�DUERUHV-FHQFH�LQ�Aloe LV�IRXQG�QRW�RQO\�LQ�HDUO\�GLYHUJLQJ�OLQHDJHV�EXW�DOVR�LQ�RWKHUV�GHHSO\�HPEHGGHG�ZLWKLQ�Aloe��DQG�WKDW�WKH�VPDOO��VWHPOHVV�JUDVV�Aloe �$��VHFW��Leptoaloe��DUH�GHULYHG�

Leaf insertion. —�'LVWLFK\�LV�HYLGHQWO\�WKH�MXYHQLOH�FRQGL-WLRQ��SUHVHQW�LQ�DOO�Aloe�DQG�Gasteria�VHHGOLQJV��DQG�LWV�SHUVLV-WHQFH�LQ�DGXOW�SODQWV�LV�EHVW�LQWHUSUHWHG�DV�QHRWHQ\�

Leaf tuberculation. — 7XEHUFXODWLRQ�LV�FHUWDLQO\�D�GHULYHG�FRQGLWLRQ��DV�K\SRWKHVLVHG�E\�6PLWK��9DQ�:\N���������,Q�Gasteria��9DQ�-DDUVYHOG��������KDV�SURSRVHG�WKDW�LWV�HYROXWLRQ�ZDV�GULYHQ�E\�WKH�DEVHQFH�RI�WKH�ELWWHU�FRQVWLWXHQW�W\SLFDO�RI�Aloe VSHFLHV��LPSO\LQJ�WKDW�WKH�ULJLG�WXEHUFOHV�PD\�PDNH�WKH�OHDYHV�OHVV�SDODWDEOH�

Perianth colour. —�7KH�EULJKW�\HOORZ��RUDQJH�RU�UHGGLVK�IORZHUV�W\SLFDO�RI�PRVW�$ORRLGHDH�DUH�VWURQJO\�DVVRFLDWHG�ZLWK�RUQLWKRSKLO\��7KH�ZKLWLVK�RU�JUHHQLVK�SHULDQWK�FKDUDFWHULVWLF�RI�Chortolirion��Haworthia, Astroloba �H[FOXGLQJ�Poellnitzia) DQG�Aloe�VHFW��Leptoaloe��DSSHDUV�WR�EH�D�GHULYHG�DGDSWDWLRQ�WR�HQWRPRSKLO\��%RWHV��DO���������+DUJUHDYHV��DO���������

The Haworthia-type flower. —�6PDOO��VSUHDGLQJ�IORZHUV�ZLWK�D�ZKLWLVK��PRUH�RU�OHVV�ELODELDWH�SHULDQWK�DQG�LQFOXGHG�DQWKHUV�DUH�GLDJQRVWLF�RI�Haworthia DQG�Chortolirion �%D\HU���������%D\HU��������LGHQWLILHG�VPDOO�IORUDO�GLIIHUHQFHV�LQ�WKH�WKUHH�VXEJHQHUD�RI�Haworthia��ZKLFK�LV�FRQVLVWHQW�ZLWK�WKH�LQGHSHQGHQW�HYROXWLRQ�RI�WKLV�IORZHU�W\SH�LQ�WKLV�JHQXV��7KH�FRQYHUJHQFH�LQ�WKLV�IORZHU�V\QGURPH�LQ�Chortolirion SURYLGHV�FOHDU�HYLGHQFH�WKDW�VXFK�D�IORZHU�W\SH�FDQ�HYROYH�LQGHSHQGHQWO\�IURP�DQ�RUQLWKRSKLORXV�DQFHVWRU� 5HODWLYHO\�VKRUW�WXEHG��ZKLW-LVK�RU�FUHDP�FRORXUHG�IORZHUV�LQ�VRPH�Aloe�VSHFLHV�VXFK�DV�A. inconspicua KDYH�EHHQ�VKRZQ�WR�EH�DQ�DGDSWDWLRQ�WR�LQVHFW�SROOLQDWLRQ��%RWHV��DO����������$OWKRXJK�VWLOO�UHFRJQLVDEO\�µAloe-OLNH¶��WKH�IORZHUV�RI�EHH�SROOLQDWHG�Aloe VSHFLHV��VHQVX�%RWHV��DO���������VXFK�DV�A. linearifolia DQG�A. minima�GLV-SOD\�VHYHUDO�FKDUDFWHULVWLFV�RI�WKH�Haworthia�W\SH�IORZHU�DSDUW�IURP�UHGXFHG�VL]H��QDPHO\�WKHLU�QHDUO\�KRUL]RQWDO�RULHQWDWLRQ��ZKLWLVK�DQG�ZHDNO\�ELODELDWH�SHULDQWK��DQG�VRPHWLPHV�LQFOXGHG�VWDPHQV��6PLWK��9DQ�:\N��������VXJJHVWHG�WKDW�IORUDO�]\JR-PRUSK\�LQ�Astroloba, Chortolirion DQG�Haworthia�UHSUHVHQWV�DQ�DGYDQFHG�VWDWH�GHULYHG�IURP�WKH�SOHVLRPRUSKLF�DFWLQRPRU-SKLF�SDWWHUQ��ZKLFK�LV�VXSSRUWHG�E\�RXU�DQDO\VLV�

The Gasteria-type flower. —�7KH�VR�FDOOHG�JDVWHULIRUP�IORZHU��FXUYHG�DQG�IODVN�VKDSHG�ZLWK�DQ�RYRLG��LQIODWHG�WXEH�DW�OHDVW�KDOI�DV�ORQJ�DV�WKH�SHULDQWK��DQG�LQFOXGHG�RU�VKRUWO\�

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Ɣ� Scenario 3. –�$�WD[RQRPLF�µVSOLWWHU¶V¶�DSSURDFK�WKURXJK�WKH�UHFRJQLWLRQ�RI�DGGLWLRQDO�VPDOOHU�JHQHUD�ZLWKLQ�$ORRLGHDH�LQ�RUGHU�WR�UHWDLQ�DOO�RU�PRVW�RI�WKH�FXUUHQWO\�DFFHSWHG�JHQHUD�DV�PRQRSK\OHWLF��$OWKRXJK�ZH�DUH�QRZ�LQ�SRVVHVVLRQ�RI�D�ZHOO�VDPSOHG�WUHH��WKH�SRRU�VXSSRUW�DORQJ�WKH�EDFNERQH�SODFHV�WKH�VDPH�FRQVWUDLQWV�XSRQ�XV��8QWLO�WKHVH�EDVDO�QRGHV�DUH�EHWWHU�VXSSRUWHG�ZH�UHPDLQ�XQDEOH�WR�SURSRVH�DQ�DOWHUQDWLYH�FODVVLIL-FDWLRQ��Aloe aristata LV�FOHDUO\�PLVSODFHG�ZLWK�WKH�RWKHU�DORHV�

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Ɣ� Scenario 4. – $� WD[RQRPLF� µOXPSHU¶V¶� DSSURDFK�� LQ�ZKLFK�DOO�PHPEHUV�RI�$ORRLGHDH�DUH�LQFOXGHG�LQ�D�VLQJOH�ODUJH�JHQXV�Aloe��,Q�WKLV�VFHQDULR�DGGLWLRQDO�LQIUDJHQHULF�WD[D�DUH�QHFHVVDU\�WR�UHIOHFW�WKH�PRUSKRORJLFDO�GDWD�DQG�WKH�UHVXOWV�RI�WKH�SK\ORJHQHWLF�DQDO\VHV��$OWKRXJK�XQZLOOLQJ�WR�DGRSW�WKLV�VR-OXWLRQ�IRUPDOO\��7UHXWOHLQ��DO�������E��SURSRVHG�LQIRUPDO�WD[R-QRPLF�JURXSLQJV�WKDW�LQGLFDWH�XQHTXLYRFDOO\�WKDW�WKH\�IDYRXU�WKLV�VROXWLRQ��7KHLU�DUWLFXODWHG�REMHFWLRQ�WR�DGRSWLQJ�LW�WXUQV�RQ�WKH�SUDFWLFDO�GLIILFXOWLHV�RI�LPSOHPHQWLQJ�D�KLHUDUFKLFDO�FODV-VLILFDWLRQ�VKRXOG�WKH�HYROXWLRQ�RI�WKH�VXEIDPLO\�EH�VKRZQ�WR�EH�UHWLFXODWH��7KH�KLJK�FRQJUXHQFH�EHWZHHQ�WKH�SODVWLG�DQG�QXFOHDU�WUHHV��HVSHFLDOO\�DW�WKH�ORZHU�QRGHV��LQ�RXU�DQDO\VLV�LV�D�FOHDU�LQGLFDWLRQ�WKDW�UHWLFXODWH�HYROXWLRQ�LV�QRW�D�VLJQLILFDQW�SUREOHP�DW�WKHVH�OHYHOV��$OWKRXJK�5DPGKDQL��DO���������KDYH�VXJJHVWHG�WKDW�K\EULGLVDWLRQ�LV�ULIH�ZLWKLQ�Haworthia� GXH�WR�WKH�KLJK�OHYHOV�RI�LQFRQJUXHQFH�REVHUYHG�EHWZHHQ�WKH�SODVWLG�DQG�WKH�QXFOHDU�WUHHV��WKHLU�VWXG\�XWLOLVHG�PXOWLSOH�DFFHVVLRQV�ZLWKLQ�D�JHQXV�LQ�ZKLFK�VSHFLHV�ERXQGDULHV�DUH�QRWRULRXVO\�XQFHUWDLQ��7KHLU�FRQFOXVLRQV�WKHUHIRUH�KDYH�OLWWOH�EHDULQJ�RQ�WKH�LVVXH�KHUH��7KLV�RSWLRQ�DFKLHYHV�PD[LPXP�QRPHQFODWXUDO�VWDELOLW\�EXW�WKH�LQIRUPDWLRQ�FRQWHQW�LV�UHGXFHG�DW�WKH�JHQHULF�OHYHO��DOWKRXJK�WKH�UHFRJQLWLRQ�RI�LQIUDJHQHULF�WD[D�DW�WKH�OHYHO�RI�VXEJHQHUD�DQG�VHFWLRQV�ZLOO�UHWDLQ�WKLV�

2Q�PRUSKRORJLFDO�JURXQGV�WKHUH�LV�OLWWOH�WR�SUHFOXGH�LP-SOHPHQWLQJ�WKLV�RSWLRQ�VLQFH�Aloe�DOUHDG\�LQFOXGHV�PRVW�RI�WKH�YDULDWLRQ�HYLGHQW�LQ�WKH�VXEIDPLO\��%UDQFK�VXSSRUW�DW�WKH�ORZHU�QRGHV�LQ�RXU�DQDO\VLV�LV�QRW�KLJK�HQRXJK�WR�SUHGLFDWH�WKH�DGRS-WLRQ�RI�WKLV�RSWLRQ��DQG�VHTXHQFLQJ�RI�DGGLWLRQDO�JHQH�UHJLRQV�PLJKW�VWLOO�JURXS�WKH�WUHH�DQG�UDPEOLQJ�DORHV�ZLWK�WKH�WUXH�DORHV��8QWLO�D�VXLWDEO\�ZHOO�VXSSRUWHG�WRSRORJ\�LV�DYDLODEOH�RQ�ZKLFK�WR�EDVH�D�SK\ORJHQHWLF�FODVVLILFDWLRQ�LW�VHHPV�EHVW�WR�UHIUDLQ�IURP�PDMRU�IRUPDO�WD[RQRPLF�DQG�QRPHQFODWXUDO�DGMXVWPHQWV�

NOMENCLATURE

Aloe VHFW� Chortolirion��$��%HUJHU��%RDWZU���-�&��0DQQLQJ��comb. & stat. nov.�Ł�Chortolirion�$��%HUJHU�LQ�(QJOHU��3IODQ]HQU��,9�����,,,������������±�7\SH��Chortolirion an-golense (%DNHU��$��%HUJHU� [= Aloe subspicata� �%DNHU��%RDWZU���-�&��0DQQLQJ@

��

'DUX��DO����Phylogeny of AlooideaeTAXON����������)HEUXDU\���������±��

��9HUVLRQ�RI�5HFRUG��LGHQWLFDO�WR�SULQW�YHUVLRQ��

Adams, S.P., Leitch, I.J. & Bennett, M.D. ������5LERVRPDO�'1$�HYROXWLRQ�DQG�SK\ORJHQ\�LQ�Aloe �$VSKRGHODFHDH���Amer. J. Bot. ��������±�����

APG II ������$Q�XSGDWH�RI�WKH�$QJLRVSHUP�3K\ORJHQ\�*URXS�FODV-VLILFDWLRQ�IRU�WKH�RUGHUV�DQG�IDPLOLHV�RI�IORZHULQJ�SODQWV��$3*�,,��Bot. J. Linn. Soc.���������±����

Baldwin, B.G., Sanderson, M.J., Porter, J.M., Wojciechowski, M.F., Campbell, C.S. & Donoghue, M.J. ������7KH�,76�UHJLRQ�RI�QX-FOHDU�ULERVRPDO�'1$�$�YDOXDEOH�VRXUFH�RI�HYLGHQFH�RQ�$QJLR-VSHUP�SK\ORJHQ\��Ann. Missouri Bot. Gard.��������±����

Bayer, M.B. ������Haworthia handbook. .LUVWHQERVFK��1DWLRQDO�%R-WDQLF�*DUGHQV�RI�6RXWK�$IULFD�

Bayer, M.B. ������The new Haworthia handbook��.LUVWHQERVFK��&DSH�7RZQ��1DWLRQDO�%RWDQLF�*DUGHQV�RI�6RXWK�$IULFD�

Bayer, M.B. ������Haworthia revisited: A revision of the genus��+DW-ILHOG��8PGDXV�3UHVV�

Bayer, M.B. ������Haworthia update: Essays on Haworthia��YRO�����+DWILHOG��8PGDXV�3UHVV�

Bayer, M.B. ������Haworthia supplement��/RQGRQ��7KH�+DZRUWKLD�6RFLHW\�

���Aloe aestivalis %RDWZU���-�&��0DQQLQJ��nom. nov.�Ł�Chor-tolirion latifolium�=RQQ���*�3�-��)ULW]�LQ�%UDGOH\D���������ILJV���±����������QRQ�Aloe latifolia��+DZ���+DZ��±�+ROR-W\SH��6RXWK�$IULFD��>)UHH�6WDWH@��%ORHPIRQWHLQ�QHDU�DLUSRUW��������Fritz 1025��35(��

���Aloe subspicata �%DNHU��%RDWZU���-�&��0DQQLQJ��comb. nov.� Ł� Haworthia subspicata� %DNHU� LQ� %XOO�� +HUE��%RLVVLHU��VHU������������������±�+RORW\SH��6RXWK�$IULFD��>*DXWHQJ@��0RGGHUIRQWHLQ����6HS�������Conrath 645��=��LVRW\SH��.��

= Haworthia angolensis %DNHU�LQ�7UDQV��/LQQ��6RF��/RQGRQ��%RW���������������Ł�Chortolirion angolense (%DNHU��$��%HUJHU�LQ�(QJOHU��3IODQ]HQU��,9�����,,,�������������QRQ�Aloe angolen-sis %DNHU�±�+RORW\SH��$QJROD��+XLOOD��1RY�������Welwitsch 3756��%0��

���Aloe tenuifolia��(QJO���%RDWZU���-�&��0DQQLQJ��comb. nov. Ł�Haworthia tenuifolia�(QJO��LQ�%RW��-DKUE��6\VW���������W����������Ł�Chortolirion tenuifolium��(QJO���$��%HUJHU�LQ�(QJOHU��3IODQ]HQU��,9�����,,,������������±�+RORW\SH��6RXWK�$IULFD��>1RUWKHUQ�&DSH@��QHDU�.XUXPDQ��)HE�������Marloth 1049��%��LVRW\SH��35(��

ACKNOWLEDGEMENTS

:H�ZRXOG�OLNH�WR�WKDQN�%UXFH�0��%D\HU��6HDQ�*LOGHQKX\V��(UQVW�YDQ�-DDUVYHOG��DQG�WKH�0ROHFXODU�6\VWHPDWLFV�VHFWLRQ�RI�-RGUHOO�/DER-UDWRU\�DW� WKH�5R\DO�%RWDQLF�*DUGHQV��.HZ��/DV]OR�&VLED�DQG�)HOL[�)RUHVW��IRU�VDPSOHV��7KH�1DWLRQDO�5HVHDUFK�)RXQGDWLRQ��15)��6RXWK�$IULFD�DQG�WKH�5R\DO�6RFLHW\�RI�/RQGRQ�DUH�JUHDWO\�DFNQRZOHGJHG�IRU�IXQGLQJ��7KLV�SURMHFW�ZDV�DOVR�SDUWO\�IXQGHG�E\�WKH�*RYHUQPHQW�RI�&DQDGD�WKURXJK�*HQRPH�&DQDGD�DQG�WKH�2QWDULR�*HQRPLFV�,QVWLWXWH�������2*,�,&,������:H�DSSUHFLDWH�WZR�DQRQ\PRXV�UHIHUHHV�ZKR�SUR-YLGHG�XVHIXO�FRPPHQWV�RQ�DQ�HDUOLHU�GUDIW�RI�WKH�PDQXVFULSW�

LITERATURE CITED

Botes, C., Johnson, S.D. & Cowling, R.M. ������&RH[LVWHQFH�RI�VXF-FXOHQW�WUHH�DORHV��3DUWLWLRQLQJ�RI�ELUG�SROOLQDWRUV�E\�IORUDO�WUDLWV�DQG�IORZHULQJ�SKHQRORJ\��Oikos ��������±����

Botes, C., Wragg, P.D. & Johnson, S.D. ������1HZ�HYLGHQFH�IRU�EHH�SROOLQDWLRQ�V\VWHPV�LQ�Aloe �$VSKRGHODFHDH��$ORLGHDH���D�SUHGRPL-QDQWO\�ELUG�SROOLQDWHG�JHQXV��S. African J. Bot. �������±����

Brandham, P.E. ������(YROXWLRQ�LQ�D�VWDEOH�FKURPRVRPH�V\VWHP��,Q��%UDQGKDP��3�(���%HQQHWW��0�'���HGV����Kew Chromosome Confer-ence II. /RQGRQ��*HRUJH�$OOHQ��8QZLQ�

Bremer, K. ������7KH�OLPLWV�RI�DPLQR�DFLG�VHTXHQFH�GDWD�LQ�DQJLRVSHUP�SK\ORJHQHWLF�UHFRQVWUXFWLRQ��Evolution��������±����

CBOL Plant Working Group ������$�'1$�EDUFRGH�IRU�ODQG�SODQWV��Proc. Natl. Acad. Sci. U.S.A.�����������±������

Chase, M.W., De Bruijn, A.Y., Cox, A.V., Reeves, G., Rudall, P.J., Johnson, M.A.T. & Eguiarte, L.E. ������3K\ORJHQHWLFV�RI�$VSKR-GHODFHDH��$VSDUDJDOHV���$Q�DQDO\VLV�RI�SODVWLG�rbcL�DQG�trnL-F '1$�VHTXHQFHV��Ann. Bot. (Oxford)��������±����

DeSalle, R. & Brower A.V.Z. ������3URFHVV�SDUWLWLRQV��FRQJUXHQFH�DQG�WKH�LQGHSHQGHQFH�RI�FKDUDFWHUV��,QIHUULQJ�UHODWLRQVKLSV�DPRQJ�FORVHO\�UHODWHG�+DZDLLDQ�Drosophila�IURP�PXOWLSOH�JHQH�UHJLRQV��Syst. Biol.��������±����

Devey, D.S., Leitch, I., Rudall, P.J., Pires, J.C., Pillon, Y. & Chase, M.W. ������6\VWHPDWLFV�RI�;DQWKRUUKRHDFHDH�VHQVX�ODWR��ZLWK�HPSKDVLV�RQ�Bulbine��Aliso��������±����

Doyle, J.J. & Doyle, J.L. ������$�UDSLG�'1$�LVRODWLRQ�SURFHGXUH�IRU�VPDOO�TXDQWLWLHV�RI�IUHVK�OHDI�WLVVXH��Phytochem. Bull.�������±���

Duval, H.A. ������Plantae succulentae in horto Alenconio��3DULV�Edgar, R.C. ������086&/(��0XOWLSOH�VHTXHQFH�DOLJQPHQW�ZLWK�KLJK�

DFFXUDF\�DQG�KLJK�WKURXJKSXW��Nucl. Acids Res.���������±�����Felsenstein, J. ������&RQILGHQFH�OHYHOV�RQ�SK\ORJHQLHV��$Q�DSSURDFK�

XVLQJ�WKH�ERRWVWUDS��Evolution��������±����Fritz, G.P.J. ������5HYLHZ�RI�WKH�WKUHH�VSHFLHV�DFFHSWHG�LQ�Chortolirion

$��%HUJHU��;DQWKRUUKRHDFHDH��$VSKRGHORLGHDH���Aloe������±���Germishuizen, G., Meyer, N.L., Steenkamp, Y. & Keith, M. (eds.)

������A checklist of South African plants. 6RXWKHUQ�$IULFDQ�%R-WDQLFDO�'LYHUVLW\�1HWZRUN�5HSRUW�1R������3UHWRULD��6$%21(7�

Gildenhuys, S.D. ������Haworthia marxii �$VSKRGHODFHDH�$ORDFHDH���D�QHZ�VSHFLHV�IURP�WKH�/LWWOH�.DURR��6RXWK�$IULFD��Aloe �����±��

Glen, H.F. & Hardy, D.S. ������$ORDFHDH��Aloe��3S���±���LQ��*HUPLV�KXL]HQ��*���HG����Flora of Southern Africa��YRO�����SW�����IDVF���� Aloaceae (First part): Aloe��3UHWRULD��1DWLRQDO�%RWDQLFDO�,QVWLWXWH�

Glen, H.F. & Smith, G.F. ������1RWHV�RQ�WKH�W\SLILFDWLRQ�RI�VRPH�VSHFLHV�RI�Aloe �$VSKRGHODFHDH�$ORDFHDH���Bothalia ������±���

Goldblatt, P. & Manning, J.C. ������5DGLDWLRQ�RI�SROOLQDWLRQ�V\VWHPV�LQ�WKH�,ULGDFHDH�RI�VXE�6DKDUDQ�$IULFD��Ann. Bot. (Oxford) �������±����

Hargreaves, A.L., Harder, L.D. & Johnson, S.D. ������Aloe incon-spicua��7KH�ILUVW�UHFRUG�RI�DQ�H[FOXVLYHO\�LQVHFW�SROOLQDWHG�DORH��S. African J. Bot. �������±����

Hillis, D.M. & Dixon, M.T. ������5LERVRPDO�'1$��0ROHFXODU�HYROX-WLRQ�DQG�SK\ORJHQHWLF�LQIHUHQFH��Quart. Rev. Biol. �������±����

Holland, P.G. ������$Q�HYROXWLRQDU\�ELRJHRJUDSK\�RI�WKH�JHQXV�Aloe. J. Biogeogr. ������±����

Huelsenbeck, J.P. & Ronquist, F. ������0U%D\HV��%D\HVLDQ�LQIHUHQFH�RI�SK\ORJHQHWLF�WUHHV��Bioinformatics��������±����

Jeppe, B. ������South African aloes��&DSH�7RZQ��3XUQHOO�Jombart, T. & Dray, S. ������DGHSK\OR��([SORUDWRU\�DQDO\VHV�IRU�WKH�

SK\ORJHQHWLF�FRPSDUDWLYH�PHWKRG��Bioinformatics ��������±�����Klopper, R.R. & Smith, G.F. ������7KH�JHQXV�Aloe /���$SVKRGHODFHDH��

$ORRLGHDH��LQ�1DPDTXDODQG��6RXWK�$IULFD��Haseltonia �����±���Klopper, R.R., Van Wyk, A.E. & Smith, G.F. ����� 3K\ORJHQHWLF�

UHODWLRQVKLSV�LQ�WKH�IDPLO\�$VSKRGHODFHDH��$VSDUDJDOHV���Biodiv-ers. & Ecol.�����±���

Linnaeus, C. ������Species plantarum���QG�HG��6WRFNKROP�Maddison, W.P. & Maddison, D.R. ������0HVTXLWH��$�PRGXODU�V\VWHP�

IRU�HYROXWLRQDU\�DQDO\VLV��YHUVLRQ�������KWWS���PHVTXLWHSURMHFW�RUJManning, J.C. & Smith, G.F. ������$VSKRGHODFHDH��$ORRLGHDH��7KH�

JHQXV�Poellnitzia�LQFOXGHG�LQ�Astroloba��Bothalia��������

��

TAXON����������)HEUXDU\���������±��'DUX��DO����Phylogeny of Alooideae

�� 9HUVLRQ�RI�5HFRUG��LGHQWLFDO�WR�SULQW�YHUVLRQ��

Meyer, N.L. & Smith, G.F. ������Astroloba corrugata��'HVFULSWLRQ�RI�D�ORQJ�NQRZQ�VSHFLHV�LQ�D�VRXWKHUQ�$IULFDQ�HQGHPLF�$ORRLG�JHQXV��Bothalia ������±���

Meyer, N.L. & Smith, G.F. ������$ORDFHDH��3S�����±����LQ��(JJOL��8���HG����Illustrated handbook of succulent plants: Monocotyledons. %HUOLQ��6SULQJHU�

Mössmer, M., Smith, G.F., Van Wyk, B.E. & Condy, G. ������Hawor-thia fasciata ID��fasciata��Fl. Pl. Africa ������±���

Obermeyer, A.A. ������$ORDFHDH��Haworthia koelmaniorum�2EHUP���+DUG\. Fl. Pl. Africa ���������

Obermeyer, A.A. ������/LOLDFHDH��Aloe, Chamaealoe� Haworthia� Astroloba� Poellnitzia DQG�Chortolirion. Bothalia ��������

Palumbi, S.R. ������1XFOHLF�DFLGV�,,��7KH�SRO\PHUDVH�FKDLQ�UHDFWLRQ��3S�����±����LQ��+LOOLV��'�0���0RULW]��&���0DEOH��%�.���HGV����Molecular systematics���QG�HG��6XQGHUODQG��6LQDXHU�

Paradis, E., Claude, J. & Strimmer, K. ������$3(��$QDO\VHV�RI�SK\OR-JHQHWLFV�DQG�HYROXWLRQ�LQ�5�ODQJXDJH��Bioinformatics��������±����

Posada, D. & Crandall, K.A. ������02'(/7(67��7HVWLQJ�WKH�PRGHO�RI�'1$�VXEVWLWXWLRQ��Bioinformatics �������±����

R Development Core Team ������5��$�ODQJXDJH DQG�HQYLURQPHQW�IRU�VWDWLVWLFDO�FRPSXWLQJ��5�)RXQGDWLRQ�IRU�6WDWLVWLFDO�&RPSXWLQJ��9LHQQD��$XVWULD��KWWS���ZZZ�5�SURMHFW�RUJ�

Ramdhani, S., Barker, N.P. & Cowling, R.M. ������5HYLVLWLQJ�PRQR-SK\O\�LQ�Haworthia 'XYDO��$VSKRGHODFHDH���,QFRQJUXHQFH��K\-EULGL]DWLRQ�DQG�FRQWHPSRUDU\�VSHFLDWLRQ��Taxon ��������±�����

Ratnasingham, S. & Herbert, P.D. ����� %2/'��7KH�%DUFRGH�RI�/LIH�'DWD�6\VWHP��KWWS���ZZZ�EDUFRGLQJOLIH�RUJ���Molec. Ecol. Notes �� ���±����

Reeves, G., Chase, M.W., Goldblatt, P., Fay, M.F., Cox, A.V., Lejeune, B. & Souza-Chies, T. ������0ROHFXODU�V\VWHPDWLFV�RI�,ULGDFHDH��(YLGHQFH�IURP�IRXU�SODVWLG�'1$�UHJLRQV��Amer. J. Bot. ��������±�����

Reynolds, G.W. ������The aloes of tropical Africa and Madagascar. 0EDEDQH��6ZD]LODQG��7KH�$ORHV�%RRN�)XQG�

Reynolds, G.W. ������The aloes of South Africa.�&DSH�7RZQ��$�$��%DONHPD�

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

Salm-Dyck, J.D. ����±������Monographia generum Aloe et Mesem-bryanthemi����YROV��%RQQ�

Sang, T., Crawford, D.J. & Steussy, T.F. ������&KORURSODVW�'1$�SK\-ORJHQ\��UHWLFXODWH�HYROXWLRQ�DQG�ELRJHRJUDSK\�RI�Paeonia��3DHRQL�DFHDH���Amer. J. Bot.���������±�����

Seelanan, T., Schnabel, A. & Wendel, J.F. ������&RQJUXHQFH�DQG�FRQVHQVXV�LQ�WKH�FRWWRQ�WULEH��0DOYDFHDH���Syst. Bot. �������±����

Small, R.L., Cronn, R.C. & Wendel, J.F. ������8VH�RI�QXFOHDU�JHQHV�IRU� SK\ORJHQ\� UHFRQVWUXFWLRQ� LQ�SODQWV��Austral. Syst. Bot. �������±����

Smith, G.F. ������+LVWRULFDO�UHYLHZ�RI�WKH�WD[RQRP\�RI�Chortolirion %HUJHU��$VSKRGHODFHDH��$ORRLGHDH�. Aloe ������±���

Smith, G.F. ����D��)6$ FRQWULEXWLRQV����$VSKRGHODFHDH�$ORDFHDH����������Poellnitzia. Bothalia ������±���

Smith, G.F. ����E��)6$ FRQWULEXWLRQV����$VSKRGHODFHDH�$ORDFHDH����������Chortolirion. Bothalia ������±���

Smith, G.F. & Steyn, E.M.A. ������7D[RQRP\�RI�$ORDFHDH��3S����±���LQ��5H\QROGV��7���HG����Aloes: The genus Aloe.�/RQGRQ��&5&�3UHVV�

Smith, G.F. & Steyn, E.M.A. ������1RWHV�RQ�WKH�SKHQRORJ\��QDWXUDO�JHRJUDSKLFDO�GLVWULEXWLRQ�UDQJH�DQG�WD[RQRP\�RI�Aloe dichotoma �$ORDFHDH���Bradleya ������±���

Smith, G.F. & Van Wyk, B.E. ������*HQHULF�UHODWLRQVKLSV�LQ�WKH�$OR�RLGHDH��$VSKRGHODFHDH���Taxon �������±����

Smith, G.F. & Van Wyk, A.E. ����� 7KH�JHQHULF�VWDWXV�RI�Chortolirion �$ORDFHDH���(YLGHQFH�IURP�OHDI�DQDWRP\��Kew Bull. �������±����

Smith, G.F. & Van Wyk, B.E. ������$VSKRGHODFHDH��3S�����±����LQ��.XELW]NL��.���HG��� The families and genera of vascular plants��YRO�����Flowering plants: Monocotyledons; Lilianae (except Orchid-aceae).�1HZ�<RUN��6SULQJHU�

Smith, G.F., Van Wyk, B.E. & Condy, G. ������Poellnitzia rubriflora. Fl. Pl. Africa ������±���

Smith, G.F., Van Wyk, B.E., Steyn, E.M.A. & Breuer, I. ������,QIUD-JHQHULF�FODVVLILFDWLRQ�RI�Haworthia �$ORDFHDH���3HUVSHFWLYHV�IURP�QHFWDU�VXJDU�DQDO\VLV��Syst. Geogr. Pl. �������±����

Sorenson, M.D. & Franzosa, E.A. ������7UHH5RW��YHUVLRQ����%RVWRQ��0DVVDFKXVHWWV��%RVWRQ�8QLYHUVLW\�

Swofford, D.L. ������3$83 ��3K\ORJHQHWLF�DQDO\VLV�XVLQJ�SDUVLPRQ\�� �DQG�RWKHU�PHWKRGV���YHUVLRQ����E����6XQGHUODQG��0DVVDFKXVHWWV��6LQDXHU�

Taylor, W.R. ������&\WRORJLFDO�VWXGLHV�RQ�Gasteria�,,��$�FRPSDULVRQ�RI�WKH�FKURPRVRPHV�RI�Gasteria��Aloe�DQG�Haworthia��Amer. J. Bot. �������±����

Treutlein, J., Smith, G.F., Van Wyk, B.E. & Wink, M. ����D��(YL-GHQFH�IRU�WKH�SRO\SK\O\�RI�Haworthia��$VSKRGHODFHDH�VXEIDPLO\�$ORRLGHDH��$VSDUDJDOHV��LQIHUUHG�IURP�QXFOHRWLGH�VHTXHQFHV�RI�rbcL��matK��,76��DQG�JHQRPH�ILQJHUSULQWLQJ�ZLWK�,665�3&5��Pl. Biol.�������±����

Treutlein, J., Smith, G.F., Van Wyk, B.E. & Wink, M. ����E��3K\-ORJHQHWLF�UHODWLRQVKLSV�LQ�$VSKRGHODFHDH��VXEIDPLO\�$ORRLGHDH��LQIHUUHG�IURP�FKORURSODVW�'1$�VHTXHQFHV��rbcL��matK��DQG�IURP�JHQRPLF�ILQJHU�SULQWLQJ��,665���Taxon �������±����

Uitewaal, A.J.A. ������$�ILUVW�DWWHPSW�WR�VXEGLYLGH�WKH�JHQXV�Hawor-thia�EDVHG�RQ�IORUDO�FKDUDFWHUV��Desert Pl. Life �������±����

Van Jaarsveld, E.J. ������7KH�JHQXV�Gasteria��$�V\QRSWLF�UHYLHZ��Aloe������±���

Van Jaarsveld, E.J. ������Gasterias of South Africa: A new revision of a major succulent group. &DSH�7RZQ��)HUQZRRG�3UHVV�LQ�DVVRFLD-WLRQ�ZLWK�WKH�1DWLRQDO�%RWDQLFDO�,QVWLWXWH��6RXWK�$IULFD�

Van Jaarsveld, E.J. ������$�QHZ�WD[RQ�DQG�QHZ�FRPELQDWLRQV�LQ�WKH�Gasteria carinata FRPSOH[��Cact. Succ. J. (Los Angeles) ������±���

Van Jaarsveld, E.J. ������Gasteria polita YDQ�-DDUVY���D�QHZ�VSHFLHV�IURP�WKH�:HVWHUQ�&DSH��Cact. Succ. J. (Los Angeles) �������±����

Van Jaarsveld, E.J. ������7KH�JHQXV�Gasteria��$�V\QRSWLF�UHYLHZ��QHZ�WD[D�DQG�FRPELQDWLRQV���Aloe�������±����

Van Jaarsveld, E.J. & Van Wyk, A.E. ������Gasteria doreeniae� D�QHZ�VSHFLHV�IURP�WKH�(DVWHUQ�&DSH��Aloe ������±���

Van Jaarsveld, E.J. & Van Wyk, A.E. ������Gasteria tukhelensis� D�QHZ�VSHFLHV�IURP�.ZD=XOX�1DWDO��6RXWK�$IULFD��Bothalia �������±����

Van Jaarsveld, E.J. & Van Wyk, A.E. ������Aloe challisii� D�QHZ�FOLII�GZHOLLQJ�DORH�IURP�0SXPDODQJD��DQG�D�FKHFNOLVW�RI�WKH�REOLJDWH�FOLII�GZHOOLQJ�DORHV�LQ�6RXWK�$IULFD�DQG�1DPLELD��Aloe ������±���

Van Jaarsveld, E.J., Smith, G.F. & Van Wyk, B.E. ������$�FODGLVWLF�DQDO\VLV�RI�Gasteria �$ORDFHDH���S. African J. Sci. �������±����

Van Wyk, B.E. & Smith, G.F. ������Guide to aloes of South Africa��3UHWRULD��%UL]D�3XEOLFDWLRQV�

Van Wyk, B.E., Whitehead, C.S., Glen, H.F., Hardy, D.S., Van Jaarsveld, E.J. & Smith, G.F. ������1HFWDU�VXJDU�FRPSRVLWLRQ�LQ�WKH�VXEIDPLO\�$ORRLGHDH��$VSKRGHODFHDH���Biochem. Syst. Ecol. �������±����

Viljoen, A.M. ������A chemotaxonomic study of phenolic leaf com-pounds in the genus Aloe. 3K�'��WKHVLV��5DQG�$IULNDDQV�8QLYHUVLW\��-RKDQQHVEXUJ��6RXWK�$IULFD�

Viljoen, A.M., Van Wyk, B.E. & Van Heerden, F.R. ������'LVWUL-EXWLRQ�DQG�FKHPRWD[RQRPLF�VLJQLILFDQFH�RI�IODYRQRLGV�LQ�Aloe �$VSKRGHODFHDH���Pl. Syst. Evol. �������±���

Wiens, J.J. ������&RPELQLQJ�GDWD�VHWV�ZLWK�GLIIHUHQW�SK\ORJHQHWLF�KLV-WRULHV��Syst. Biol. �������±����

Yoder, A.D., Irwin, J.A. & Payseur, B.A. ������)DLOXUH�RI�WKH�,/'�WR�GHWHUPLQH�GDWD�FRPELQDELOLW\�IRU�VORZ�ORULV�SK\ORJHQ\��Syst. Biol. �������±����

Zonneveld, B.J.M. & Van Jaarsveld, E.J. ������7D[RQRPLF�LPSOLFD-WLRQV�RI�JHQRPH�VL]H�IRU�DOO�VSHFLHV�RI�WKH�JHQXV�Gasteria 'XYDO��$ORDFHDH���Pl. Syst. Evol. ��������±����

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