Molecular phylogenetics of Alooideae (Asphodelaceae)
by
Barnabas Haruna Daru
Dissertation submitted
in fulfilment of the requirements
for the degree
MAGISTER SCIENTIAE
in
BOTANY
in the
FACULTY OF SCIENCE
at the
UNIVERSITY OF JOHANNESBURG, SOUTH AFRICA
Supervisor: Prof. M. van der Bank
Co-supervisor: Dr. O. Maurin
January 2012
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DECLARATION
I declare that this dissertation has been composed by myself and the work contained
within, unless otherwise stated, is my own.
Barnabas Haruna Daru (January 05, 2012)
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TABLE OF CONTENTS
Declaration .......................................................................................................................... ii
Table of Contents ............................................................................................................... iii
Index to tables .................................................................................................................... vi
Index to Figures ............................................................................................................... viii
Acknowledgements ........................................................................................................... xii
Foreword .......................................................................................................................... xiii
List of Abbreviations ..................................................................................................... xivv
Abstract ............................................................................................................................ xvi
Chapter One: General Introduction and Objectives
1.
GENERAL INTRODUCTION AND OBJECTIVES ............................................... 1
1.1 Family Asphodelaceae .................................................................................................. 1
1.1.1 Subfamily Asphodeloideae ........................................................................................ 2
1.1.2 Subfamily Alooideae ................................................................................................. 2
1.2 Review of the genera within Alooideae ........................................................................ 4
1.2.1 Aloe L. (1753) ............................................................................................................ 4
1.2.2 Astroloba Uitew. (1947) ............................................................................................ 6
1.2.3 Chortolirion A.Berger (1908) .................................................................................... 7
1.2.4 Gasteria Duval (1809) ............................................................................................... 8
1.2.5 Haworthia Duval (1809) ............................................................................................ 9
1.3
Objectives of the study ............................................................................................ 10
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Chapter Two: Material and Methods
2. MATERIAL AND METHODS .................................................................................... 19
2.1 Choice of gene regions................................................................................................ 19
2.2 Taxon sampling........................................................................................................... 19
2.3 DNA extraction, amplification and sequencing .......................................................... 20
2.4 Phylogenetic analyses and tree construction............................................................... 21
2.5 Coding of morphological characters ........................................................................... 23
Chapter Three: Results
3. RESULTS ..................................................................................................................... 49
3.1 Molecular Evolution ................................................................................................... 49
3.2 Combined plastid data................................................................................................. 49
3.3 ITS1 data ..................................................................................................................... 50
3.4 Combined plastid and nuclear dataset (rbcLa + matK + trnH-psbA + ITS1) ........... 550
3.5 Evolution of morphological traits ............................................................................... 53
Chapter Four: Discussion and Conclusions
4. DISCUSSION AND CONCLUSIONS ........................................................................ 70
4.1 Phylogenetic Relationships within Alooideae ............................................................ 70
4.1.1 Relationships within Aloe ........................................................................................ 71
4.1.2 Relationships within Haworthia .............................................................................. 72
4.1.3 Relationships within smaller segregate genera ........................................................ 74
4.2 Evolution of selected morphological characters in Alooideae ................................... 74
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4.3 Conclusions ................................................................................................................. 79
Chapter Five: References
REFERENCES ................................................................................................................. 82
Appendix I Abstracts of all papers presented at conferences ........................................... 91
Appendix II Paper submitted to Taxon – 15 December 2011 (under review) .................. 94
Appendix III Taxonomic changes in Alooideae resulting from the current study ........... 95
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INDEX TO TABLES
Chapter One: General Introduction and Objectives
Table 1.1 Alooideae genera (Van Wyk et al., 1993; Smith & Steyn, 2004; Klopper et al.,
2010) ................................................................................................................................. 11
Table 1.2 Proposed families within Asparagales from 1981 to 2009. Names in brackets
are the combination of groups belonging to that family ................................................... 12
Table 1.3 History of genera recognised within Asphodelaceae from 1981 to 2010 ......... 13
Table 1.4 Historical bibliographical notes on genus Haworthia (Author dates are based
on Scott (1985) and Bayer (1999) .................................................................................... 14
Chapter Two: Material and Methods
Table 2.1 Table of taxa, voucher information, and accession numbers from GenBank.
DNA sequences obtained from GenBank are indicated by 1Treutlein et al., 2003a,
2
Treutlein et al., 2003b, and 3Kim et al., 2010. Original publication dates and synonyms
were obtained from The Plant List Version 1 (2010) ...................................................... 25
Table 2.2 Gene regions and primers used for amplifying the cpDNA and nrDNA in the
current study...................................................................................................................... 39
Table 2.3 Characters, character states, and explanatory notes on characters used in the
cladistic analyses of the morphological data matrix. ........................................................ 40
Table 2.4 Character states for the 20 morphological characters scored for accessions
included in the moelcular analyses. .................................................................................. 41
Chapter Three: Results
Table 3.1 Statistics from MP analyses obtained from separate and combined datasets. .. 56
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Table 3.2 Characters states for nine selected autapomorphic traits which have been
employed in previous Alooideae systematics. ................................................................ 567
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INDEX TO FIGURES
Chapter One: General Introduction and Objectives
Figure 1.1 (a) Global distribution of all Alooideae genera highlighted in red. Distributions
of (b) Gasteria; (c) Haworthia; (d) Astroloba; (e) Aloe; (f) Chortolirion. Adapted from
Smith & van Wyk (1991), Bayer (1999) and Viljoen (1999). .......................................... 15
Figure 1.2 (a) Aloe peglarae Schönland, and (b) A. plicatilis with distichous leaf
arrangement. Photo: O. Maurin. ....................................................................................... 16
Figure 1.3 (a) Astroloba rubriflora (L.Bolus) G.F.Sm. & J.C.Manning, (b) Astroloba
corrugata N.L.Mey. & Gideon F.Sm.and (c) A. spiralis (L.) Uitewaal showing the
densely packed triangular leaves (Photo: M.B.Bayer)...................................................... 16
Figure 1.4 (a) Chortolirion angolense showing leaves and underground bulb, and (b)
zygomorphic flowers (b). Photo: Sean Gildenhuys (a) and www.lifestyleseeds.co.za (b)16
Figure 1.5 (a) Gasteria croucheri (Hook.f) leaves with bands of whitish spots and (b)
secund inflorescence of G. carinata var. retusa van Jaarsv. Photo: O. Maurin. .............. 18
Figure 1.4 Haworthia mirabilis (Haw.) Haw. showing (a) succulent leaves and, (b)
bilabiate flower. Photo: O. Maurin. .................................................................................. 18
Chapter Two: Material and Methods
Figure 2.1 Organisation of the ITS region. Green area represents ITS region sequenced in
the study. Arrows indicate orientation and approximate position of primer sites. Adapted
from Baldwin et al. (1995)................................................................................................ 47
Figure 2.2 Distribution of Alooideae collected in the wild and grown at Sheilam Nursery,
Gariep Nursey and Kirstenbosch Botanical Garden, used in this study (yellow circles). 47
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Chapter Three: Results
Figure 3.1 Simplified tree topologies showing relationships among Alooideae for the
plastids, nuclear and combined dataset. ............................................................................ 58
Figure 3.2 One of the most parsimonious trees obtained from the maximum parsimony
(MP) analyses of the combined plastid data set (rbcLa + matK + trnH-psbA) for
Alooideae plus outgroups (TL = 752; CI = 0.72; RI = 0.91). Numbers above the branches
are MP bootstrap support. ................................................................................................. 60
Figure 3.3 One of the most parsimonious trees obtained from the maximum parsimony
(MP) analyses of the ITS1 data set for Alooideae (TL = 346; CI = 0.64; RI = 0.92)....... 62
Figure 3.4 Majority-rule consensus tree obtained from the Bayesian analysis of the
combined plastid and nuclear internal transcribed spacer (ITS) data set for Alooideae plus
outgroup (TL = 1159; CI = 0.63; RI = 0.89). Numbers above the branches are MP
bootstrap support and ones below are BI posterior probabilities. ..................................... 64
Figure 3.5 Distribution of morphological traits along phylogeny of Alooideae. All trait
values are centred and scaled, resulting in circles of different colours. The colours of the
symbols are proportional to the absolute value of each trait ............................................ 65
Figure 3.6 Parsimony-based reconstructions of a) habit character (character 1), b) leaf
insertion (character 2), and c) leaf tubercles (character 5) on the majority-rule consensus
tree from the combined data. ............................................................................................ 66
Figure 3.7 Parsimony-based reconstruction of a) flower orientation (character 11), b)
perianth symmetry (character 14), and c) perianth tube (character 17) on the majority-rule
consensus tree from combined molecular data. ................................................................ 65
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Figure 3.8 Parsimony-based reconstruction of a) perianth shape (character 18), b) stamen
inclusion (character 20), and c) tepal fusion (character 19) on the majority-rule consensus
tree from combined molecular data. ................................................................................. 65
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Acknowledgments
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ACKNOWLEDGEMENTS
First and foremost I would like to thank my supervisor, Prof. Michelle van der Bank for
her constant support and insightful guidance. She made sure I had the practical, financial
and intellectual tools to accomplish this work. I thank my co-supervisor Dr. Olivier
Maurin, University of Johannesburg, South Africa for ensuring that this thesis is well
written and the project carried out very well. Thank you very much.
Ledile Mankga, Kowiyou Yessoufou, Jephris Gere, Philip Rousseau, Ronny
Kabongo, Bezeng Simeon, Theresa Sethusa, Mark Cooper and Bruce Lububi as well as
Salome Malgas have been my colleagues, office, lab companions and friends for 24
months and have provided honest, precise, and always helpful critical comments.
I like to thank Erin Corstorphine and Maria (Masha) Kuzmina of the International
Barcode of Life Project, Biodiversity Institute of Ontario, University of Guelph Canada
for helping me with DNA sequencing. I am very grateful.
The following people supplied plant samples and are greatly acknowledged: Dr
Felix Forest of Royal Botanic Gardens Kew, Bruce M. Bayer, Sean Gildenhuys of Gariep
Plants Pretoria, Wesley Chin and his father, Drs John Manning, Ernst van Jaarsveld and
Stephen Boatwright of Kirstenbosch Gardens, Cape Town.
Profs Vincent Savolainen and Mark Chase; Drs Martyn Powell, Luis Valente and
Jan Schnitzler; Abel Gizaw and Adam Britton provided useful comments and provided
some research articles for the thesis.
The following people helped in one way or the other especially in terms of
logistics: Flip Minaar, Anne Fourie (Librarian of SANBI, Pretoria) and Tinus Fourie.
Oageng Kale, Pamela Sekoto and Francois Joshua helped me particularly in the
lab – thank you very much.
My parents Mr. Haruna Daru and Mrs. Christiana Daru for the ‘good genes’ and
my siblings; Patrick, Rifkat, John, Rita, Nanbam and Lydia as well as my nieces and
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Acknowledgments
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nephews (Nash P, Sekyen, Retji, Leyitnaan, Katlong, Terkuma, Durnaan, Naanman,
Joshua, Jighitnaan). Thanks for visiting me during the spring break here in Johannesburg:
Patrick, Pamela, Nash P, Sekyen, Retji, Leyitnaan – I love you all.
I appreciate the friendship of Halimat Suberu, Wussi Turaki, Toyin Ojo, families
of Dr Esther Akinlabi, Dr Peter and Mrs. Oluwatoyin Olubambi who have been my
source of motivation and encouragement throughout this study.
Staff and students of the African Centre for DNA Barcoding, University of
Johannesburg, are greatly acknowledged as well as staff of University of Jos, Nigeria,
particularly Prof H. B. Mafuyai, Drs Georgina Mwansat, Shiiwua Manu and Fidelis
Tiseer, Mrs. Cecelia and Mr. Nannim Nanvyat (Zoology Dept). Thanks for believing in
me.
Funding for this project was provided by the Royal Society of London and the
National Research Foundation (NRF), South Africa. This project was also partly funded
by the Government of Canada through Genome Canada and the Ontario Genomics
Institute (2008-OGI-ICI-03).
The Almighty God through Jesus Christ has been my source of inspiration and
help throughout this study (John 1:3). To You I dedicate this project.
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Foreword
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FOREWORD
This study is a contribution to the phylogenetic understanding of Alooideae
(Asphodelaceae) and follows on the efforts of Reynolds (1966), Van Jaarsveld (1994),
Bayer (1999), Treutlein et al. (2003a, b), and Vosa (2004). These taxonomic treatments
provided the baseline for the evaluation of hypotheses presented in this study.
The dissertation is presented in five chapters. Chapter one presents the general
synopses, motivation and objectives of the study. Chapter two provides details of the
sampling, methodology and phylogenetic analyses used. In Chapter three, the results of
the phylogenetic relationships within the subfamily Alooideae is presented. Appendices
include abstracts of papers presented at conferences as well as paper submitted to Taxon.
Preliminary findings of the study was presented at the 2011 Congress of the Southern
African Society of Systematic Biology (SASSB) and earned me the award for the Best
MSc Presentation. Appendix III resulted from collaborative research with John Manning
and Stephen Boatwright and form part of the paper submitted to Taxon (Appendix II).
All analyses, tables, images and figures used in the dissertation are by the author,
unless otherwise stated.
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List of abbreviations
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LIST OF ABBREVIATIONS
°C = Degrees Celsius
A.Berger = Alwin Berger (1871-1931)
ABI = Applied Biosystems, Inc
ACCTRAN = Accelerated transformation optimisation
ACDB = African Center for DNA Barcoding
AIC = Akaike Information Criterion
Aloes = Aloe
APG = Angiosperm Phylogeny Group
Baijnath = Baijnath, Himansu (1943-)
BI = Bayesian inference
Bp = base pair
BP = Bootstrap Percentage (support)
ca. = approximately
CBOL PG = Consortium for the Barcode of Life Plant Working Group
CI = Consistency Index
Corp. = Corporation
CTAB = Hexadecyltrimethylammonium bromide
DELTRAN = Delayed transformation optimisation.
DMSO = Dimethyl Sulfoxide
DNA = Deoxyribonucleic acid
EtOH = Ethanol
F = Forward primer
GenBank = National Center for Biotechnology Information
GTR+I+G = General Time Reversible + Gamma + Proportion Invariant
i.e. = id est (that is)
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List of abbreviations
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Inc. = Incorporated
iso. = Isotype
ITS = Internal Transcribed Spacer
JRAU = Herbarium of the University of Johannesburg (UJ), Johannesburg, South Africa
K = Herbarium of the Royal Botanic Gardens, Kew, Richmond, United Kingdom
Kunth = Kunth, Karl Sigismund (1788-1850)
L. = Linnaeus (von Linné), Carl (1707-1778)
M.Bieb. = Marschall von Bieberstein, Friedrich August (1768-1826)
m = meter
MCMC = (Bayesian) Markov Chain Monte Carlo
min. = Minimum
min = Minutes
mm = millimeter
Moench. = Moench, Conrad (1744-1805)
MP = Maximum Parsimony
MRCA = Most recent common ancestor
MulTrees = Multiple equally parsimonious trees
Mya = Millions years ago
NaCl = Sodium Chloride
No. = Number
nom. cons. = nomen conservandum
nrDNA = nuclear ribosomal deoxyribonucleic acid
NRF = National Research Foundation (South Africa)
PAUP = Phylogenetic Analysis Using Parsimony software program
PCR = Polymerase chain reaction
Poelln. = von Poellnitz, Karl (1896-1945)
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List of abbreviations
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PP = Posterior probabilities
PVP = Polyvinyl pyrolidone
R = Reverse primer
rbcL = ribulose-bisphosphate carboxylase gene
RBG Kew = the Royal Botanic Gardens Kew
RI = Retention Index
s.l. = sensu lato
s.s. = sensu stricto
SANBI = South African National Biodiversity Institute
sect. nov. = section novis
SEM = Scanning Electron Microscope
sp. nov. = species novis (new species)
Syn = Synonymous
TBR = Tree-bisection-reconnection
TIM + G = Transitional Model + Gamma
TL = Tree length
TReeBoL Africa = DNA Barcoding Africa Trees
trnH-psbA = spacer between trnH and psbA genes
Uitew. = Uitewaal, Antonius Josephus Adrianus (1899-1963)
UK = United Kingdom
USA = United States of America
UV = Ultraviolet
µl = microliter
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Abstract
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ABSTRACT
Alooideae (Asparagales–Asphodelaceae) currently comprises five genera, four of which
are endemic to southern Africa. Despite their importance in commercial horticulture,
evolutionary relationships among the genera are still incompletely understood, and the
generic taxonomy remains unresolved. This study addresses generic delimitation in
subfamily Alooideae using an expanded molecular dataset from three plastid regions
(rbcLa, matK and trnH-psbA) and the first subunit of the nuclear ribosomal internal
transcribed spacer (ITS1). Sequence data were analysed using maximum parsimony and
bayesian statistics, and selected morphological traits were mapped onto the molecular
phylogeny. Among the polytypic genera, only Astroloba and Gasteria were retrieved as
monophyletic. Haworthia is shown to be polyphyletic and to comprise three main clades
largely correllated with current subgeneric circumscriptions. The taxonomic implications
of these findings are discussed.
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General Introduction and Objectives
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CHAPTER(ONE(
General(Introduction(and(Objectives(
Photo: O. Maurin
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General Introduction and Objectives
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1.#GENERAL INTRODUCTION AND OBJECTIVES#
In this chapter, I present an overview of Alooideae taxonomy and phylogenetics and state
the objectives of my study. The classification system of APG III (2009) placed members
of Alooideae i.e. Asphodelaceae (sensu Mabberley, 2008) within the family
Xanthorrhoeaceae yet, most molecular studies in Alooideae systematics retained
Alooideae as a subfamily within Asphodelaceae (e.g. Chase et al., 2000; Treutlein et al.,
2003a, b; Klopper et al., 2010; Ramdhani et al., 2011). I have adopted the latter
classification system throughout the dissertation.
1.1 Family Asphodelaceae
Asphodelaceae is a monocotyledon family in the order Asparagales and comprises about
13 genera and ± 800 species (Klopper et al., 2010). Members of this family are
distributed in xeric and mesic regions of the temperate, subtropical and tropical zones of
the Old World (including Africa, Eurasia, Australia, New Zealand) with the main centre
of distribution in southern Africa (Van Wyk et al., 1993; Smith & van Wyk, 1998;
Treutlein et al., 2003a, b; Klopper et al., 2010).
Many species of Asphodelaceae are cultivated as ornamentals with some listed on
the IUCN Redlist of species (IUCN, 2001). For example, Aloe bowiea Schult. &
Schult.f., Bulbine frutescens (L.) Willd. var. chalumnensis Baijnath ined., Gasteria
baylissiana Rauh, G. ellaphieae Van Jaarsv., G. glomerata Van Jaarsv., Haworthia
bryunsii M.B.Bayer, H. springbokvlakensis C.L.Scott, H. woolleyi Poelln. and Kniphofia
acrea Codd have been listed as threatened in the wild with extinction by version 3.1 of
the IUCN categories and criteria (Victor & Dold, 2003).
Asphodelaceae is divided into two subfamilies, based on vegetative and
reproductive characters: Asphodeloideae and Alooideae (Treutlein et al., 2003a, b;
Klopper et al., 2010).
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General Introduction and Objectives
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1.1.1 Subfamily Asphodeloideae
Subfamily Asphodeloideae consists of the genera Trachyandra Kunth., Asphodelus L.,
Asphodeline Rchb., Eremurus M. Bieb., Bulbine Wolf., Jodrellia Baijnath, Bulbinella
Kunth and Kniphofia Moench (Smith & van Wyk, 1998; Klopper et al., 2010). These
genera are widespread throughout the temperate and subtropical regions of Africa, SaudiArabia, Madagascar and some of the Mascarene Islands off the east coast of Africa. The
subfamily has varying proportions of small to large chromosomes with a basic set of six
chromosomes (2n = 12).!
1.1.2 Subfamily Alooideae
Subfamily Alooideae (Aloaceae sensu Smith & Steyn, 2004) of the family
Asphodelaceae is an Old World group (Fig. 1.1a) often characterised by conspicuous leaf
succulence, cresentiform or cymbiform leaf outline in cross-section, and a rarely altered
diploid karyotype n = 7 (Taylor, 1925; Smith & van Wyk, 1998). Cladistically this
character represents a synapomorphy for all taxa of Alooideae. It also represents a sharp
discontinuity between the Asphodeloideae and Alooideae, and suggests a distinct barrier
for gene interchange (Smith & van Wyk, 1991, 1998). Representatives of the subfamily
also share some chemical characters, notably the presence of anthrone-C-glycosides in
their leaves and of 1-methyl-8-hydroxyanthraquinones in their roots (Smith & van Wyk,
1998).
Modern taxonomy of the Alooideae begins with Linnaeus (1753), whose concept
of the genus Aloe L. was rather heterogeneous one. Of the 16 names that he included in
this genus, four are not members of Alooideae (three are now in Sansevieria and one is in
Kniphofia). The remainder, grouped by flower size and shape, are currently segregated
among Aloe (four species), Gasteria (one species), Astroloba (one species) and
Haworthia (five species). Linnaeus’ preliminary groupings were subsequently
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General Introduction and Objectives
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formalised, first at sectional level within Aloe (Linnaeus, 1774) and later by the
recognition of the segregate genera Gasteria, defined by the moderately large, curved and
often gasteriform flowers, and Haworthia, with very much smaller, whitish flowers
(Duval, 1809). Haworthia was later further split when those species with often
actinomorphic flowers were removed from those with bilabiate flowers into the small
genus Astroloba Uitewaal (Roberts, 1965). Three additional small genera have since been
recognised: Chortolirion, Lomatophyllum and Poellnitzia. Lomatophyllum consists of
about 14 species from Madagascar and the Mascarene islands and it is now included in
the genus Aloe (Klopper et al., 2010) and the monotypic Poellnitzia rubriflora (L.Bolus)
Uitewaal was recently included in Astroloba as A. rubriflora (L.Bolus) Gideon F.Sm. &
J.C.Manning, (Manning & Smith, 2000; Germishuizen et al., 2006). Thus the current
classification of the subfamily (reviewed in Klopper et al., 2010) recognises the
following five genera: Aloe, Astroloba, Chortolirion, Gasteria, and Haworthia (Table
1.1).
Alternative classification systems are briefly discussed below.
Classification system suggested by Cronquist (1981) ! The classification
system employed by Cronquist (1981) for flowering plants is based solely on
morphological features. He placed Alooideae genera in the order Liliales within families
Aloeaceae and Liliaceae (Table 1.2). The systematic framework for Liliales laid down by
Cronquist (1981) was based on the unifying synapomorphy of seed-coat micromorphology (the characteristic black seeds caused by phytomelan incrustation of the seed
coat in most capsular and berry-fruited taxa). Cronquist (1981) maintained families such
as Alliaceae, Amaryllidaceae, Agavaceae, Iridaceae, and Orchidaceae within Liliales, but
lumped most of the rest into a single family Liliaceae, which included taxa from both
orders Asparagales and Liliales. Other authors over the years adopted this system but
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General Introduction and Objectives
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modified the spelling of the family name from “Aloeaceae” sensu Cronquist (1981) to
“Aloaceae” (e.g. Glen & Hardy, 2000; Smith & Steyn, 2004).
Classification system suggested by Mabberley (2008) ! The classification
upheld by Mabberley (2008) recognised the family Asphodelaceae (Asparagales) to
consist of 14 genera (Table 1.3) with 800 species. Mabberley (2008) noted a superficial
resemblance between Asphodelaceae, Hyacinthaceae and Anthericaceae. In his system,
two subfamilies are recognised: Alooideae (pachycaul trees with secondary growth to
succulent rosettes) and Asphodeloideae (herbaceous plants).
Classification system by APG III (2009) and Chase et al. (2009) ! The
Angiosperm Phylogeny Group (APG III, 2009) reduced the numbers of families within
Asparagales to 15 (Table 1.2). All members of Alooideae i.e. Asphodelaceae (sensu
Mabberley, 2008) were placed within Xanthorrhoeaceae. Chase et al. (2009) however, in
a subfamilial classification of Asparagales (Amaryllidaceae, Asparagaceae and
Xanthorrhoeaceae), recommended that Xanthorrhoeaceae be split into three subfamilies
(Asphodeloideae, Hemerocallidoideae and Xanthorrhoeoideae) to simplify taxonomic
delimitation in this family.
1.2 Review of genera within Alooideae
1.2.1 Aloe L. (1753)
The genus Aloe, with about 400 species native to Africa, the Arabian Peninsula, and
Madagascar plus other islands in the western Indian Ocean (Fig. 1.1e), is the largest
within the Alooideae (Reynolds, 1966; Viljoen, 1999; Glen & Hardy, 2000; Klopper &
Smith, 2007). Its species are characterised by tough, spiked/toothed leaves with
astringent/unpalatable juice, sunken pores, vivid (red, white, or sometimes bicoloured)
flowers, and wind-dispersed seeds. There are however numerous exceptions. Some Aloe
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General Introduction and Objectives
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species, have tall inflorescences with reddish-toned flowers with erect to spreading
pedicels. Polystichous leaf arrangements in spiral rosettes with spiny margins are typical
of most Aloe species (Fig. 1.2a) others have leaves arranged distichously in two vertical
rows on opposite sides of an axis. Aloe plicatilis (L.) Mill. (Fig. 1.2b) for example, often
called “fan aloe”, is characterised by its leaf clusters resembling an open fan with
margins almost smooth, except for some small teeth in the upper part and an arborescent
growth habit. Other exceptions within the genus include a wide variation in growth form
(trees, shrubs and stemless, sometimes geophytic perennials), margin almost entire to
toothed) and surface (smooth to verrucose), flower size, shape, symmetry and colour
(small to large, campanulate to tubular or gasteriform, actinomorphic to bilabiate, and
pallid to brightly coloured), and fruit characteristics (small to large and dry to fleshy).
Aloe is one of the genera within Alooideae that has been studied the most
comprehensively. Recent studies include phytochemical analysis (Viljoen et al., 1998;
Viljoen, 1999; Grace et al., 2010), leaf morphology (Grace et al., 2009), molecular
evidence (Adams et al., 2000a, b) and ecology (Botes et al., 2008) all of which are based
on the work done by Reynolds (1966).
Ecologically, due to the ease of hybridisation between Aloe species, often
flowering simultaneously and mostly ornithophilous, Botes et al. (2008) indicated that
co-flowering Aloe species partitioned flower-visiting birds through differences in floral
morphology and nectar traits. Species that share bird pollinators tend to flower
sequentially or utilise different pollen placement sites on the same birds. Aloe
inconspicua Plowes is however pollinated exclusively by insects (Amegilla fallax,
Apidae) that visit its flowers for nectar and pollen (Hargreaves et al., 2008). It has white
flowers, strongly zygomorphic with perianth tubes curved upwards. Recent molecular
evidence suggests that insect-pollination is the ancestral state for Alooidea, with the
ancestral Aloe species postulated to originate in the highlands of south-east Africa and
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spreading during the Tertiary era (Holland, 1978). These differences in pollination
system, flowering phenology, and biogeography may suggest taxonomic affinities within
the Aloe genus and also enable a greater number of Aloe species to coexist. However,
detailed pollination and biogeographic studies of Aloe species, just like other Alooideae
genera, is lacking.
1.2.2 Astroloba Uitew. (1947)
Astroloba is a clump forming, Haworthia-like perennial succulent endemic to the
Western and Eastern Cape regions of South Africa (Mabberley, 2008; Fig. 1.1d). This
genus contains six species plus Poellnitzia rubriflora L.Bolus from Western Cape
recently transferred to Astroloba (as A. rubriflora (L.Bolus) Gideon F.Sm. &
J.C.Manning) by Manning & Smith (2000) due to its close vegetative similarity to some
species of Astroloba, as well as similarities in lipophilic anthronoid aglycones. It was
essentially distinguished from Astroloba by its inclined racemes of secund, orange-red
flowers with connivent tepals, apparently an adaptation to sunbird pollination (Manning
& Smith, 2000). The six species of Astroloba are typically multi-stemmed at ground level
with stems totally covered by hard, spirally arranged, stiff, pungent leaves (Fig. 1.3). The
abaxial side of the leaf is tuberculate or smooth. The inflorescences are axillary racemes
with simple or branched peduncles. The flowers are pedicellate with a straight tubular
perianth (Smith, 1995b; Manning & Smith 2000).
Astroloba is vegetatively very similar to some species of Haworthia and the two
are distinguished solely by floral symmetry. Flowers of Astroloba are actinomorphic with
tepals spreading at the tips (star-shape) whereas the perianth in Haworthia is irregular
and nearly always bilabiate.
6
!
General Introduction and Objectives
!
!
!
1.2.3 Chortolirion A.Berger (1908)
Chortolirion comprises a single species, C. angolense (Baker) A.Berger (Mabberley,
2008). This monotypic genus has an underground bulb (Fig. 1.4a) and narrow grass-like
leaves weakly armed with small, white marginal teeth (Smith & Van Wyk, 1993; Smith,
1995b) which die back to ground level during fire or in winter, a character unique to this
genus. Its flowers closely resemble those of Haworthia species (Fig. 1.4b) and it has been
included in that genus in the past (Obermeyer, 1973) but has generally been retained as
distinct from it on the basis of the distinct, bulb-like swelling of the leaf bases (Smith,
1995a). Chortolirion is also characterised by spirally twisted, graminoid, deciduous
leaves (Smith & van Wyk, 1993). Furthermore, Chortolirion has a widespread northern
distribution in Angola, Botswana, Namibia and the summer rainfall areas in South Africa
(Fig. 1.1f), whereas Haworthia is predominantly restricted to the winter rainfall regions
in the Western and Northern Cape provinces (Fig. 1.1c).
Chortolirion, just like most Alooideae genera, have witnessed some taxonomic
controversy over the years so that a considerable number of synonyms have been
attributed to it. Smith & van Wyk (1993) using leaf anatomy found that the distribution of
tissues in the leaves of C. angolense is similar to that described for grass-like Aloe
species in sect. Leptoaloe Berger (e.g. Aloe verecunda Pole-Evans, A. boylei Baker, and
the graminifoliate Haworthia blackburniae Baker). This relationship was confirmed by a
recent molecular study (Treutlein et al., 2003b). Remarkably, it shares a xerophytic
character with other xerophytes such as sunken and protected stomata. A distinctive
palisade layer immediately inside the epidermis is conspicuous, although this has not
been accepted as a generic marker to delimit this monotypic genus (Smith et al., 1996). It
however shares the presence of inner bundle sheath-caps consisting of thin-walled
parenchymatous cells with other alooid genera.
7
!
General Introduction and Objectives
!
!
!
1.2.4 Gasteria Duval (1809)
Linnaeus (1753) in Species Plantarum included Gasteria under the name Aloe disticha
(Gasteria disticha sensu Van Jaarsveld, 1994). In 1809 Duval formally described
Gasteria, recognising 43 species. Since then many new species have been described.
Finally in 2007, Van Jaarsveld arranged Gasteria into two sections, Longiflorae and
Gasteria, based on the shape and epidermal texture of the leaf and recognised 23 species.
This classification system is widely accepted and the most comprehensive work on the
genus to date. Gasteria is endemic to South Africa with the greatest concentration
occurring in the Eastern Cape Province (Fig. 1.1b). Only one species, Gasteria pillansii
Kensit, extends beyond South Africa’s borders and crosses the Orange River into
southern Namibia (Van Jaarsveld, 2007).
Gasteria is defined as a monophyletic group within Alooideae based on floral
morphology, vegetative characters and molecular data (Zonneveld & Van Jaarsveld,
2005; Treutlein et al., 2003b). It is distinguished from most, but not all aloes by its
inclined racemes of pendulous, curved flowers (Fig. 1.5b) sometimes swollen at the base
(gasteriform). Sunbirds (Nectariniidae) and insects pollinate the flowers of Gasteria.
Bimodal karyotype is uniform in all Gasteria species with eight large and six small
chromosomes (Vosa & Bennett, 1990). The leaves of Gasteria are triangular in section
and firm-textured and mostly dark green with bands of whitish spots (Fig. 1.5a), and the
margins are horny but never spiny (Van Jaarsveld, 2007). Brandham (1977) evaluated the
inheritance of leaf pigmentation in Gasteria and showed that despite their wide
morphological diversity the ‘spotted’ Gasteria species have identical genetic systems
controlling the distribution of leaf pigmentation, which underlines their basic genetic
similarities. Today Gasteria species are threatened in the wild due to succulent collectors
and agricultural practices.
8
!
General Introduction and Objectives
!
!
!
1.2.5 Haworthia Duval (1809)
Haworthia includes approximately 61 species and numerous infra-specific taxa (Bayer,
1999; 2002; Mabberley, 2008). Most species are highly localised and are largely
restricted to the southern regions of South Africa, with outliers in the Mpumalanga,
KwaZulu-Natal and Free State provinces of South Africa, Swaziland, Mozambique and
Namibia (Fig. 1.1c). Haworthia species are small succulent herbs (Fig. 1.6a) with their
leaves arranged in a rosette. The size and shape of the leaves are variable, even within
species. The flowers are small, tubular, bilabiate and white.
Haworthia has a long history of taxonomic changes (Table 1.4) since Linnaeus
(1753) broadly placed it within “aloes” and Duval formally described it in 1809. In 1999,
Bayer subdivided Haworthia into three subgenera namely H. subgenus Haworthia, H.
subgenus Hexangulares, and H. subgenus Robustipendunculares (see also Vosa, 2004).
This was based on morphological characters as well as continuity of characters in relation
to their geographical distribution. Subgenus Haworthia has a wide range of distribution
from the Western to the Eastern Cape Provinces including the Free State. Flowers with
elongated triangles petals as well as perianths that are triangular in cross-section (e.g.
Haworthia mirabilis; Fig. 1.6b) unified members of this subgenus. Haworthia subgenus
Hexangulares occurs in the Eastern Cape Province with a disjunct distribution of H.
limifolia Marloth in the Swaziland-Mpumalanga area. Flowers are six-sided at the base.
Only four species (Haworthia marginata (Lam.) Stearn, H. pumila (L.) Duval, H. minor
(Aiton) Duval and H. kingiana Poelln.) are recognised within H. subgenus
Robustipendunculares. They are distributed exclusively in the Western Cape Province
and it is characterised by robust tubular perianths. Two recent molecular studies
(Treutlein et al., 2003a; Ramdhani et al., 2011) recovered the above-mentioned
subgenera but demonstrated that Haworthia was paraphyletic. Sampling in both studies
were however limited.
9
!
General Introduction and Objectives
!
!
!
1.3 Objectives of the study
From the above it is evident that evolutionary relationships among the Alooideae genera
are still incompletely understood and the generic taxonomy remains uncertain. As yet, no
adequately sampled and well-supported phylogenetic analysis exists on which to base a
classification for Alooideae. Treutlein et al. (2003b) offered three alternative scenarios
for resolving this problem but however refrained from implementing any of them. This
study was thus aimed at addressing previous morphological and molecular hypotheses on
generic circumscriptions and affinities among the five genera of Alooideae using
molecular sequence data.
The specific objectives of the study were to:
1) Reconstruct a phylogeny for Alooideae (154 species representing all five genera
and 20 sections within Aloe, incorporating a wide range of vegetative and floral
diversity in the subfamily) using DNA sequences from the internal transcribed
spacers (ITS1) of nuclear ribosomal DNA and three plastid regions (rbcLa, trnHpsbA and matK).
2) Mapped morphological trait data onto the phylogeny to determine the distribution
of these characters in order to investigate whether these traits are phylogenetically
structured.
3) Data from 1 and 2 were then used to: a) clarify phylogenetic relationships among
the genera, b) assess the monophyly of various groups within Alooideae, and c)
reconcile the current taxonomy with the molecular data.
10
!
General Introduction and Objectives
!
!
!
Table 1.1 Alooideae genera (Van Wyk et al., 1993; Smith & Steyn, 2004; Klopper et al.,
2010)
!
Currently recognised
Alooideae genera
References
Names not upheld
Aloe Linnaeus: 319
Reynolds (1966,
Catevala Medikus: 67 (1786) pro parte;
(1753)
1969), Glen &
Kumara Medikus: 69 (1786); Rhipidodendrum
Hardy (2000),
Willdenow 164 (1811); Pachidendron
Germishuizen et al.
Haworth: 35 (1821); Bowiea Haworth: 299
(2006)
(1824) non J.D. Hooker: t. 5619 (1867);
Agriodendron Endlicher: 144 (1836);
Succosaria Rafinesque: 137 (1840); Busipho
Salisbury: 76 (1866); Ptyas Salisbury: 76
(1866); Chamaealoe A.Berger: 43 (1905);
Leptaloe Stapf: t. 9300 (1933) Aloinella
Lemee: 27 (1939) non Cardot: 76 (1909);
Guillauminia Bertrand: 41 (1956); P.V.Heath:
153 (1993); Lomatophyllum Willdenow: 5
(1811)
Gasteria Duval: 6
Van Jaarsveld (1992,
Atevala Rafinesque: 136 (1840); Papilista
(1809)
1994, 2007)
Rafinesque: 137 (1840)
Haworthia Duval: 7
Bayer (1999)
Catevala Medikus: 67 (1786) pro parte;
(1809) nom. cons.
Apicra Willdenow: 167 (1811) non Haworth:
61 (1819); Kumaria Rafinesque: 137 (1840);
Tulista Rafinesque: 137 (1840)
Chortolirion A.Berger:
Smith (1995a)
!
Astroloba Uitewaal: 53
Roberts (1965),
Apicra Haworth: 61 (1819) non Willdenow:
(1947)
Smith (1995b),
167 (1811); Poellnitzia Uitewaal Succulenta
Manning & Smith
22:61 (1940)
72 (1908)
(2000)
11
!
General Introduction and Objectives
!
!
!
Table 1.2 Proposed families within Asparagales from 1981 to 2009. Names in brackets are the combination of ‘groups’ belonging to the family.
!
Cronquist (1981)
Order: Liliales
!
!
Agavaceae
Mabberley (2008)
Order: Asparagales
Alliaceae
Agapanthaceae
Amaryllidaceae
Aloeaceae
Cyanastraceae
Dioscoreaceae
Haemodoraceae
Hanguanaceae
Liliaceae
!
!
Iridaceae
Philydraceae
Pontederiaceae
Smilacaceae
Asparagaceae (Agavaceae, Anthericaceae, Aphyllanthaceae,
Behniaceae, Convallariaceae, Dracaenaceae,
Eriospermaceae, Herriaceae, Hyacinthaceae)
Asphodelaceae
Asteliaceae
Blandfordiaceae
Boryaceae
Doryanthaceae
Hemerocallidaceae
Hypoxidaceae
Iridaceae
Ixioliriaceae
Lanariaceae
Orchidaceae
Stemonaceae
Taccaceae
Velloziaceae
Xanthorrhoeaceae
!
Tecophilaeceae
!
Xanthorrhoaceae
!
Xeronemataceae
APG II (2003)
Order: Asparagales
!
!
Agavaceae (Anemarrhenaceae,
Anthericaeae, Behniaceae,
Herreriaceae)
Asparagaceae
!
!
!
!
!
!
!
!
!
!
Ruscaceae (Convallariaceae,
Dracaenaceae, Eriospermaceae,
Nolinaceae)
!
!
!
Xanthorrhoeaceae (Asphodelaceae,
Hemerocallidaceae, Phormiaceae).
!
APG III (2009)
Order: Asparagales
!
!
Amaryllidaceae (Agapanthaceae)
Asparagaceae (Agavaceae, Aphyllanthaceae,
Hesperocallidaceae, Hyacinthaceae,
Laxmanniaceae, Ruscaceae, Themidaceae)
!
Asteliaceae
Blandfordiaceae
Boryaceae
Doryanthaceae
!
Hypoxidaceae
Iridaceae
Ixioliriaceae
Lanariaceae
Orchidaceae
Ruscaceae
Tecophilaeceae
!
Xanthorrhoaceae (Asphodelaceae,
Hemerocallidaceae)
Xeronemataceae
!
12
!
General Introduction and Objectives
!
!
!
Table 1.3 History of genera recognised within Asphodelaceae from 1981 to 2010.
Cronquist (1981)
Smith & van Wyk (1991)
Mabberley (2008)
Klopper et al. (2010)
Family: Aloeaceae
Family: Asphodelaceae
Family: Asphodelaceae
Family: Asphodelaceae
Aloe L.
Aloe L.
Aloe L.
Aloe L.
!
!
Asphodeline Reichb.
Asphodeline Reichb.
!
!
Asphodelus L.
Asphodelus L.
!
Astroloba Uitew.
Astroloba Uitew.
Astroloba Uitew.
!
!
Bulbine Wolf.
Bulbine Wolf.
!
!
Bulbinella Kunth.
Bulbinella Kunth.
!
Chortolirion A.Berger
Chortolirion A.Berger
Chortolirion A.Berger
!
!
Eremurus M.Bieb.
Eremurus M.Bieb.
Gasteria
Gasteria
Gasteria Duval
Gasteria Duval
Haworthia (including
Astroloba, Chortolirion,
and Poellnitzia)
Haworthia Duval
Haworthia Duval
Haworthia Duval
!
!
Jodrellia Baijnath
Jodrellia Baijnath
KniphofiaMoench.
KniphofiaMoench.
KniphofiaMoench.
KniphofiaMoench.
LomatophyllumWilld.
LomatophyllumWilld.
LomatophyllumWilld.
!
!
PoellnitziaUitew.
!
!
!
!
Trachyandra Kunth.
Trachyandra Kunth.
13
!
General Introduction and Objectives
!
!
!
Table 1.4 Historical bibliographical notes on genus Haworthia (Author dates are based
on Scott (1985) and Bayer (1999).
!
Author (Date)
Placement of Haworthia
Reference
Linnaeus (1753)
“Aloes”
In Species Plantarum Part I (first edition) (1753)
Linnaeus (1762)
“Aloes”
In Species Plantarum Part I (second edition) (1762)
Burman (1768)
“Aloes”
In Appendix: Florae Capensis Prodromus in Flora
Indica (1768)
Miller (1768)
“Aloes”
In Gardeners’ Dictionary, ed. 8 (1768)
Aiton (1789)
“Aloes”
Hortus Kewensis Vol. 1, pp. 466-471 (1789).
Willdenov (1799)
“Aloes”
Species Plantarum 2: 184-192 (1799)
De Candolle (17991801)
“Aloes”
“Aloes” illustrations: Plantarum historia
succulentarum (Histoire des plantes grasses)
Jacquin (1804)
“Aloe”
“Aloes” illustrations (1804)
Thunberg (1785)
“Aloe”
Dissertation on the genus “Aloe” (1785)
Friedrich Medicus
(1786)
“Aloe” genus Catevala
First to subdivide and separate genus Aloe (today’s
Aloe + Haworthia) from genus Catevala
Haworth (1804)
“Aloe” section Paviflorae
“A new arrangement of the genus Aloe” in Trans.
Linn. Soc. 7:1-28 (1804).
Duval (1809)
Haworthia
Plantae succulentae in Horto Alenconio, separated
genera Haworthia and Gasteria from Aloe (1809).
Willdenov (1811)
Generic name: Apicra for
Paviflorae
Ges. Naturf. Fr. Berl. Mag. 5: 167 (1811)
Aiton (1811)
“Aloes”
Hortus Kewensis ed. 2, 2:292-301 (1811)
Haworth (1812-27)
Haworthia
In Synopsis plantarum succulentarum pp. 90-99,
accepted Duval’s genera Haworthia and Gasteria
Schultes & Schultes
(1829)
Reverted to “Aloe” in
broad Linnaean sense
Roemer and Schultes, Systema Vegetabilium 7: 682715 (1829)
Salm-Dyck (1836-63)
“Aloe” in the broad sense
Monographiae Generum Aloes et Mesembryantheni
Kunth (1843)
“Aloe” in a broad sense
Enumeratio Plantarum, Vol 4, gave Apicra and
Haworthia separate status as sections (1843).
Baker (1870-71; 1896)
Haworthia
Baker (1880)
Aloineae
Saunders, Refugium Botanicum Vol. 4; Flora Capensis
Vol.6: 332-355 (edited by Sir W. T. Thiselton-Dyer)
(1896)
“A synopsis of Aloineae and Yuccoideae” in J. Linn.
Soc. Bot. Vol. 18 (1880);
Berger (1908)
Haworthia
Engler’s Pflanzenreich 4, 38: 74-114 (1908).
Smith (1942)
Haworthia
J. S. Afr. Bot. 8: 247. Collections donated to
Kirstenbosch in 1957
Uitewaal (1938-1951)
Haworthia
Possibly the greatest European student of Haworthia.
Published widely in Succulenta.
Bayer (1962, 1976;
1982; 1999, 2002)
Haworthia
Haworthia Handbook (1976), and The New Haworthia
Handbook (1982, 1999, 2002); Haworthia revisited
(1999); Haworthia update. (1999)
14
!
General Introduction and Objectives
!
!
!
a
b
c
e
f
d
Figure 1.1 (a) Global distributions of all Alooideae genera highlighted in red. Distributions of (b)
Gasteria; (c) Haworthia; (d) Astroloba; (e) Aloe; (f) Chortolirion. Adapted from Smith & van
Wyk (1991), Bayer (1999) and Viljoen (1999).
15
!
!
General Introduction and Objectives
!
!
!
Figure 1.2 (a) Aloe peglarae Schönland and (b) A. plicatilis with distichous leaf arrangement.
Photo: O. Maurin
Figure 1.3 (a) Astroloba rubriflora (L.Bolus) G.F.Sm. & J.C.Manning, (b) Astroloba corrugata
N.L.Mey. & Gideon F.Sm. and (c) A. spiralis (L.) Uitewaal showing the densely packed
triangular leaves (Photo: M.B.Bayer).
Figure 1.4 (a) Chortolirion angolense showing leaves and underground bulb and (b) zygomorphic
flowers. Photo: Sean Gildenhuys (a) and www.lifestyleseeds.co.za (b)
16
!
General Introduction and Objectives
!
!
!
Figure 1.5 (a) Gasteria croucheri (Hook.f) leaves with bands of whitish spots and (b) the secund
inflorescence of G. carinata var. retusa van Jaarsv. Photo: O. Maurin
Figure 1.6 (a) Haworthia mirabilis (Haw.) Haw. Showing succulent leaves and (b) bilabiate
flower. Photo: O. Maurin
17
!
!
Chapter two: Material and Methods
!
!
CHAPTER(TWO(
Material(and(Methods(
!
!
!
!
!
!
18
!
Chapter two: Material and Methods
!
!
2. MATERIAL AND METHODS
2.1 Choice of gene regions
The first subunit of the nuclear ribosomal internal transcribed spacer (ITS1; Fig. 2.1) and three
plastid regions (matK, rbcLa and trnH-psbA) were chosen to reconstruct a phylogeny of
Alooideae in order to add to the already existing phylogenies of Chase et al. (2000) and
Treutlein et al. (2003b). Also, these regions have been shown to be useful in resolving
phylogenetic relationships at various taxonomic levels in other plant groups (e.g. Hillis &
Dixon, 1991; Baldwin et al., 1995; Small et al., 2004).
2.2 Taxon sampling
Representatives of all five genera accepted in subfamily Alooideae (including 154 species and
subspecies) were analysed for the four gene regions mentioned above. I included 20 species of
Gasteria, 69 Haworthia, 60 Aloe from 20 sections (including Lomatophyllum), four Astroloba
(including Poellnitzia) and one Chortolirion species in the analyses. Samples were collected
from living material in private and national collections in South Africa (Sheilam Nursery,
Robertson; Gariep Nursery, Pretoria; University of Johannesburg (JRAU); Kirstenbosch
Botanical Gardens, Cape Town). Most of these accessions were originally wild-collected
(Figure 2.2). Representatives of Anthericaceae (Anthericum liliago L.), Asphodelaceae:
Asphodeloideae (Asphodeline lutea (L.) Reichb., Bulbine frutescens (L.) Willd., Bulbine
semibarbata (R.Br.) Haw., Eremurus spectabilis M.Bieb., Bulbine fistulosa (Chiov.) Baijnath,
Kniphofia galpinii Baker, and Kniphofia uvaria (L.) Oken), Tecophilaeaceae (Tecophilaea
cyanocrocus Leyb. and Zephyra elegans D.Don), and Xanthorrhoeaceae (Xanthorrhoea
resinosa Pers. and Xanthorrhoea sp.) were selected as outgroups based on previous molecular
and morphological studies within Asparagales (Smith & van Wyk, 1991; Chase et al., 2000;
Treutlein et al., 2003a, b; Devey et al., 2006). These samples were obtained from the DNA
19
!
Chapter two: Material and Methods
!
!
Bank at the Royal Botanic Gardens, Kew (UK). Voucher specimen information and GenBank
accession numbers are listed in Table 2.1. Taxonomic concepts in Gasteria and Haworthia
follow Van Jaarsveld (2007) and Bayer (1999), respectively.
2.3 DNA extraction, amplification and sequencing
Total genomic DNA was extracted from either fresh or silica gel dried leaf material using the
10× CTAB method described by Doyle & Doyle (1987). Polyvinyl pyrolidone (2% PVP) was
added to reduce the effect of high polysaccharide concentrations in the samples. All samples
were purified using QIAquick purification columns (QIAgen, Inc., Hilden, Germany)
according to the manufacturer’s protocol.
Primers used for the polymerase chain reaction (PCR) amplification of the cpDNA
rbcLa, matK, and trnH-psbA, were rbcLa-F: rbcLa-R, Kim Ki-Joong-3F: Kim Ki-Joong-1-R
(CBOL Plant Working Group, 2009), and psbAF: trnH-R (Sang et al., 1997), respectively. The
ITS1 region was amplified using the primer combination ITS18: ITS5 (Treutlein et al.,
2003a). The PCR amplification primers were also used as cycle sequencing primers (Table
2.2).
PCR amplification for rbcLa and matK was carried out at the Canadian Centre for
DNA Barcoding (CCDB), Biodiversity Institute of Ontario, University of Guelph, Canada.
Details of the project including voucher information, GPS coordinates, pictures and DNA
barcodes are available on BOLD (http://www.boldsystems.org) (Ratnasingham & Herbert,
2007) within the project file ‘Alooideae of Africa’ (ALOAF). Sequencing of ITS1 and trnHpsbA as well as some additional matK and rbcLa was carried out at the African Centre for
DNA Barcoding (ACDB) of the University of Johannesburg, South Africa. All PCR
amplifications were performed using ReadyMix Master mix (Advanced Biotechnologies,
Epsom, Surrey, UK). Bovine serum albumin (3.2%) was added to both nuclear and plastid
20
!
Chapter two: Material and Methods
!
!
reactions, whereas 4.5% dimethyl sulfoxide (DMSO) was added only to matK and ITS1
amplifications. These additives serve as stabilisers for enzymes, reduce problems with
secondary structure and improve annealing (Palumbi, 1996). PCR reaction amplification was
performed using the following programs: for rbcLa and trnH-psbA, pre-melt at 94°C for 3
min, denaturation at 94°C for 1 min, annealing at 48°C for 1 min, extension at 72°C for 1 min
(for 28 cycles), followed by a final extension at 72°C for 7 min; for matK, the protocol
consisted of pre-melt at 94°C for 1 min, denaturation at 94°C for 30 s, annealing at 50°C for
40 s, extension at 72°C for 40 s (for 35 cycles), and a final extension at 72°C for 5 min. The
ITS1 protocol consisted of pre-melt at 94°C for 3 min, denaturation at 94°C for 1 min,
annealing at 48°C for 1 min, extension at 72°C for 3 min (for 26 cycles), followed by a final
extension at 72°C for 7 min. Prior to cycle sequencing, PCR products were visualised on a
1.5% agarose gel and subsequently purified using QIAquick (Qiagen Inc.) silica columns
according to the manufacturer’s protocol. The PCR products were purified using QIAquick
(Qiagen Inc.) silica columns according to the manufacturer’s protocol.
Cycle sequencing reactions for all genes used in this study were performed using ABI
PRISM® BigDye® Terminator v3.1 Cycle Sequencing Kit (Applied Biosystems, Inc.,
California, USA). Cycle sequenced products were precipitated in ethanol and sodium acetate
to remove excess dye terminators before sequencing on an ABI 3130x1 genetic analyser.
2.4 Phylogenetic analyses and tree construction
Complementary strands were assembled and edited using Sequencher 4.8 (Gene Codes, Ann
Arbor, Michigan, USA). The sequences were aligned using Multiple Sequence Comparison by
Log-Expectation (MUSCLE v. 3.8.31; Edgar, 2004) and the alignment finally adjusted
manually in PAUP* (v. 4.0b.10; Swofford, 2002) without difficulties because of low levels of
insertions/deletions except for the trnH-psbA region of which 15.5% regions were excluded
21
!
Chapter two: Material and Methods
!
!
from analyses due to alignment difficulties at positions 1-24, 123-202; 272-290; 830-839 of
the
aligned
matrix.
The
aligned
matrices
are
available
from
the
author
(darunabas@gmail.com).
The separate datasets were assessed for congruence by means of visual inspection of
the individual bootstrap consensus trees. The bootstrap trees were declared incongruent in
cases where they exhibit “hard” (high bootstrap support) instead of “soft” (low bootstrap
support; Seelanan et al., 1997) incongruence. Maximum parsimony analyses (MP) were
performed on the ITS1, combined plastid, and total combined datasets whereas Bayesian
inference (BI) analysis was employed only on the combined data set. Maximum parsimony
analyses were performed using PAUP* version 4.0b.10 (Swofford, 2002). Tree searches were
conducted using 1 000 random sequence additions, retaining 10 trees at each step, with treebisection-reconnection (TBR) branch swapping and MulTrees in effect. Resulting trees were
then used in a second search with the same parameters, however, without a limit for the
number of trees per replicate (swopping to completion). Delayed transformation (DELTRAN)
character optimisation was used instead of acceleration of transformation (ACCTRAN) for
calculating branch lengths, because of reported errors with version 4.0b.10 of PAUP*
(http://paup.csit.fsu.edu/problems.html). Branch support was estimated using bootstrap
analysis (Felsenstein, 1985) with 1 000 replicates, simple sequence addition, no swapping,
with MulTrees in effect saving 10 trees per replicate. The following scale for evaluating
bootstrap support (BP) was applied: weak (50-74%), moderate (75-84%) or high (85-100%).
Bayesian inference analysis (BI; Huelsenbeck & Ronquist, 2001; Ronquist &
Huelsenbeck, 2003) was performed using MRBAYES v. 3.1.2. For each matrix (ITS1, rbcLa,
matK, trnH-psbA) the most appropriate model was selected based on Akaike Information
Criterion (AIC) implemented in MODELTEST v. 3.06 (Posada & Crandall, 1998). The model
22
!
Chapter two: Material and Methods
!
!
TRN+G was selected for ITS1, TVM+G for matK, HKY+G for rbcLa and TVM+1+ G for
trnH-psbA. I used 2 000 000 generations with a sample frequency of 200. Partition analysis
was run for the combined dataset. The log-likelihood scores were plotted to determine the
point of stationarity, and all trees prior to stationarity were discarded as the “burn-in” phase
(1000 trees). All remaining trees were imported into PAUP* 4.0b.10 and a majority-rule
consensus tree was produced showing the frequencies (posterior probabilities or PP) of all
observed bi-partitions. The following scale was used to evaluate the PPs: below 0.95, weakly
supported; 0.95-1.0, strongly supported. To map the BP and PP values (MrBayes tree) onto
the tree, the nexus tree file from the Bayesian analysis was rescaled using the ape vs. 2.0-1
(Paradis et al., 2004) and adephylo vs. 1.1 (Jombart & Dray, 2010) packages implemented in
R (R Development Core Team, 2011).
2.5 Coding of morphological characters
A matrix of 20 morphological characters was prepared for the 154 species of Alooideae
included in the analyses. Most morphological studies in Alooideae systematics employed
these diagnostic characters (often chosen at species level) to infer relationships within the
subfamily (Germishuizen et al., 2006; Gildenhuys, 2007; Glen & Hardy, 2000; Glen & Smith,
1995; Jeppe, 1969; Bayer, 1982, 1999, 2002, 2009; Klopper & Smith, 2007; Meyer & Smith,
1998, 2001; Mössmer et al., 1995; Reynolds, 1966, 1969; Smith et al., 1995; Smith & Steyn,
2005; Smith, 1995a, b; Van Jaarsveld et al., 1994; Van Jaarsveld & van Wyk, 2004, 2005,
2006; Van Jaarsveld, 1992, 1994, 1998, 2001; Van Wyk & Smith, 2003). To test for
phylogenetic structure in morphological characters, the R function ‘table.phylo4d’
implemented in the R package adephylo vs. 1.1 (Jombart & Dray, 2010) was used. By plotting
phylogeny and traits simultaneously, this function allows a direct visualisation of phylogenetic
structure in morphological traits.
23
!
Chapter two: Material and Methods
!
!
The patterns of evolution of these characters were examined by reconstructing them
onto the majority-rule consensus tree produced by the BI analysis using Mesquite vs 2.75
(Maddison & Maddison, 2011). Morphological characters and character-states are defined in
Table 2.3 and the data matrix used for character reconstructions is shown in Table 2.4. All trait
values are centred and scaled into binary values. These values are represented using coloured
symbols, with the different colours proportional to the absolute value of each trait.
24
!
Chapter two: Material and Methods
!
!
Table 2.1 Table of taxa, voucher information, and accession numbers from GenBank. DNA sequences obtained from GenBank are indicated by
1
Treutlein et al., 2003a, 2Treutlein et al., 2003b, and 3Kim et al., 2010. Original publication dates and synonyms were obtained from The Plant
List Version 1 (2010).
!
Original Publication Details
Aloe L.
Section Aloe
Aloe arenicola Reynolds
Aloe pearsonii Schönland
Aloe perfoliata L. (TYPE)
Synonym(s)
Distribution
GenBank Accession Number
Voucher
(herbarium)
trnH-psbA
ITS
rbcLa
matK
J. S. African Bot. 4: 21 (1938)
Rec. Albany Mus. 2: 229 (1911)
Sp. Pl. 319 (1753)
—
—
Aloe albispina Haw., Aloe brevifolia Haw. [nom.
illegit.], Aloe commelyni Willd., Aloe comptonii
Reynolds, Aloe depressa Salm-Dyck ex Steud. [nom.
inva.], Aloe distans Haw., Aloe flavispina Haw., Aloe
mitriformis Mill., Aloe mitriformis DC. [nom. illegit.],
Aloe mitriformis Willd. [nom. illegit.], Aloe mitriformis
var. albispina (Haw.) A.Berger, Aloe mitriformis var.
angustior Lam., Aloe mitriformis var. commelyni
(Willd.) Baker, Aloe mitriformis var. comptonii
(Reynolds) Zonn., Aloe mitriformis subsp. distans
(Haw.) Zonn., Aloe mitriformis var. elatior Haw., Aloe
mitriformis var. flavispina (Haw.) Baker, Aloe
mitriformis var. humilior Haw.,Aloe mitriformis var.
humilior Willd., Aloe mitriformis var. pachyphylla
Baker, Aloe mitriformis var. spinosior Haw., Aloe
mitriformis var. spinulosa (Salm-Dyck) Baker, Aloe
mitriformis var. xanthacantha (Willd.) Baker, Aloe
nobilis Haw., Aloe parvispina Schönland, Aloe
perfoliata var. brevifolia Aiton, Aloe perfoliata var.
mitriformis (Mill.) Aiton, Aloe reflexa Marum ex Steud.,
Aloe spinulosa Salm-Dyck, Aloe xanthacantha Willd.
Western Cape
Northern Cape, Namibia
Eastern and Western Cape
OM3018 (JRAU)
OM2955 (JRAU)
BHD391
(JRAU)BHD461
(JRAU) OM2819
(JRAU)
JQ024863
JQ024868
—
JQ025268
JQ025269
JQ025315
JQ024528
JQ024487
JQ024526
JQ024527
JQ024111
JQ024154
JQ024155
JQ024156
Gard. Chron. n.s., 1874(1): 566
(1874)
Aloe bainesii Dyer, Aloe bainesii Dyer var. barberae
(Dyer) Baker, Aloe zeyheri Baker [nom. illegit.]
BHD192 (JRAU)
JQ024864
JQ025262
JQ024489
JQ024113
Aloe eminens Reynolds and
P.R.O.Bally
J. S. African Bot. 24: 187
(1958)
—
Eastern Cape, KwazuluNatal, Mpumalanga,
Swaziland
Northern Somalia
BHD474 (JRAU)
JQ039258
JQ025369
—
—
Section Anguialoe
Aloe alooides (Bolus) Druten
Bothalia 6: 544 (1956)
Mpumalanga
BHD476 (JRAU)
JQ039243
JQ025325
—
—
Aloe spicata L.f.
Suppl. Pl. 205 (1782)
Aloe recurvifolia Groenew., Notosceptrum alooides
(Bolus) Benth., Urginea alooides Bolus
Aloe sessiliflora Pole-Evans
OM1522 (JRAU)
JQ039282
JQ025290
—
—
Aloe vryheidensis Groenew.
Tijdschr. Natuurk. Wetensch.
Kunsten 38 (1935)
Kwazulu-Natal, Limpopo,
Mpumalanga, Swaziland
Kwazulu-Natal, Limpopo,
Mpumalanga
BHD467 (JRAU)
JQ039288
JQ025308
—
—
Section Aloidendron
Aloe barberae Dyer
Aloe dolomitica Groenew.
Section Arborescentes
25
!
Chapter two: Material and Methods
!
!
Aloe arborescens Mill.
Gard. Dict. ed. 8 3 (1768)
Aloe perfoliata var. arborescens (Mill.) Aiton, Catevala
arborescens (Mill.) Medik.
Eastern and Western Cape,
Gauteng, Kwazulu-Natal,
Limpopo, Mpumalanga,
North-West, Botswana,
Swaziland
BHD420 (JRAU)
JQ039246
JQ025326
JQ024486
JQ024110
Section Aristatae
Aloe aristata Haw.
Philos. Mag. J. 67: 280 (1825)
Aloe aristata Haw. var. leiophylla Baker, Aloe aristata
Haw. var. parvifolia Baker, Aloe ellenbergeri
Guillaumin, Aloe longiaristata Schult. and Schult.f.
Eastern and Western Cape,
Free State, Kwazulu-Natal,
Limpopo
BHD459 (JRAU)
BG HeidelbergBG
Jena
JQ039247
JQ025312
AY3236511
AY3236521
AJ5123191
AY3236341
AJ5114071
AY3237131
Aloes S. Afr. Veld 287 (1971)
—
Namibia
BHD386 (JRAU)
JQ039254
JQ025303
JQ024500
JQ024125
Bot. Jahrb. Syst. 10: 2 (1889)
—
Free State, Northern Cape,
Namibia
BHD388 (JRAU)
JQ039266
JQ025305
JQ024514
JQ024140
Gard. Chron. III, 1905(2): 102
(1905)
—
Kwazulu-Natal, Limpopo,
Mpumalanga, Botswana,
Swaziland
OM1625 (JRAU)
JQ039249
JQ025299
—
—
Philos. Trans. 66: 310 (1776)
Rhipidodendrum dichotomum (Masson) Willd.
Northern Cape, Namibia
OM2953 (JRAU)
—
JQ025368
JQ024501
JQ024126
Aloe dichotoma Masson subsp.
pillansii (L.Guthrie) Zonn.
Bradleya 20: 10 (2002)
Aloe pillansii L.Guthrie
Northern Cape, Namibia
BHD390 (JRAU)
JQ039255
JQ025372
JQ024502
JQ024127
Aloe dichotoma Masson subsp.
ramosissima (Pillans) Zonn.
Bradleya 20: 10 (2002)
Aloe dichotoma Masson var. ramosissima (Pillans) Glen
and D.S.Hardy, Aloe ramosissima Pillans
Namibia
OM2954 (JRAU)
JQ039256
JQ025367
JQ024503
JQ024128
Bot. Jahrb. Syst. 36: 63 (1905)
Bothalia 12: 62 (1976)
—
—
Northern and Western Cape
Eastern Cape
BHD462 (JRAU)
BHD381 (JRAU)
JQ039271
JQ039277
JQ025267
JQ025324
—
JQ024530
—
JQ024158
Gard. Dict. ed. 8 7 (1768)
Aloe disticha L. var. plicatilis L., Aloe flabelliformis
Salisb., Aloe lingua Thunb., Aloe linguaeformis L.f.
[nom. illegit.], Aloe tripetala Medik., Kumara disticha
Medick., Rhipidodendron distichum (Medick.) Willd.,
Rhipidodendron plicatile (L.) Haw.
Western Cape
BHD193 (JRAU)
BG Heidelberg,
Germany 9286
JQ039278
JQ025373
AY3236621
JQ024531
AY3236131
JQ024159
AY3236931
Fl. Pl. South Africa 18: 707
(1938)
—
Limpopo
RL1193 (JRAU)
JQ039270
JQ025348
—
—
J. S. African Bot. 13: 101
(1947)
Aloe kraussii Baker var. minor Baker, Aloe myriacantha
(Haw.) Schult. and Schult.f. var. minor (Baker)
A.Berger, Leptaloe albida Stapf
Mpumalanga, Swaziland
BHD475 (JRAU)
JQ039242
JQ025366
—
—
—
Aloe challisii van Jaarsv. and
A.E.van Wyk
Aloe chortolirioides A.Berger
Aloe 43: 36 (2006)
—
Mpumalanga
BHD471 (JRAU)
JQ039250
JQ025355
—
—
Pflanzenr. IV, 38: 171 (1908)
—
BHD477 (JRAU)
JQ039251
JQ025374
—
—
Aloe ecklonis Salm-Dyck
Aloes Mesembr. 2: 21 (1849)
Aloe agrophila Reynolds
BHD465 (JRAU)
JQ039257
JQ025307
—
—
Aloe fouriei D.S.Hardy and
Glen
Limpopo, Mpumalanga,
Swaziland
Eastern Cape, Free State,
Kwazulu-Natal, Limpopo,
Mpumalanga
Limpopo, Mpumalanga
BHD469 (JRAU)
JQ039261
JQ025358
—
—
Fl. Pl. Africa 49: 1941 (1987)
Section Asperifoliae
Aloe dewinteri Giess ex
Borman and Hardy
Aloe hereroensis Engl.
Section Chabaudia
Aloe chabaudii Schönland
Section Dracoaloe
Aloe dichotoma Masson
Section Echinatae
Aloe melanacantha A.Berger
Aloe pictifolia Hardy
Section Kumara
Aloe plicatilis (L.) Mill.
Section Latebracteatae
Aloe lutescens Groenew.
Section Leptoaloe
Aloe albida (Stapf) Reynolds
—
26
!
Chapter two: Material and Methods
!
!
Aloe nubigena Groenew.
Aloe saundersiae (Reynolds)
Reynolds
Aloe verecunda Pole-Evans
Aloe vossii Reynolds
Section Macrifoliae
Aloe ciliaris Haw.
Aloe commixta A.Berger
Aloe gracilis Haw.
Aloe striatula Haw.
Aloe tenuior Haw.
Section Pachydendron
Aloe angelica Pole-Evans
Aloe excelsa A.Berger
Aloe ferox Mill.
Aloe petricola Pole Evans
Aloe rupestris Baker
Aloe thraskii Baker
Section Paniculatae
Aloe buhrii Lavranos
Aloe kouebokkeveldensis van
Jaarsv. and A.B.Low
Aloe reynoldsii Letty
Aloe striata Haw.
Aloe striata Haw. subsp.
karasbergensis (Pillans) Glen
and D.S.Hardy
Tijdschr. Natuurk. Wetensch.
Kunsten 14: 3 (1936)
J. S. African Bot. 13: 103
(1947)
—
Limpopo, Mpumalanga
BHD353 (JRAU)
JQ039274
JQ025356
—
—
Leptaloe saundersiae Reynolds
Kwazulu-Natal
BHD463 (JRAU)
JQ039281
JQ025345
—
—
Trans. Roy. Soc. South Africa
5: 703 (1917)
J. S. African Bot. 2: 65 (1936)
—
BHD444 (JRAU)
JQ039286
JQ025346
—
—
—
Gauteng, Limpopo,
Mpumalanga
Limpopo
BHD0464 (JRAU)
JQ039287
JQ025347
—
Philos. Mag. J. 67: 281 (1825)
Pflanzenr. IV, 38: 260 (1908)
—
—
Eastern and Western Cape
Western Cape
BHD431 (JRAU)
BHD405 (JRAU)
JQ024866
JQ039252
JQ025292
JQ025329
JQ024496
JQ024497
—
—
JQ024121
JQ024122
Philos. Mag. J. 67: 280 (1825)
Philos. Mag. J. 67: 281 (1825)
Philos. Mag. J. 67: 281 (1825)
Aloe laxiflora N.E.Br.
—
Aloe tenuior Haw. var. decidua Reynolds, Aloe tenuior
Haw. var. densiflora Reynolds, Aloe tenuior Haw. var.
glaucescens Zahlbr., Aloe tenuior Haw. var. rubriflora
Reynolds, Aloe tenuior Haw. var. viridifolia van Jaarsv.
Eastern and Western Cape
Eastern Cape, Limpopo
Eastern and Western Cape,
Kwazulu-Natal,
Mpumalanga
BHD399 (JRAU)
BHD191 (JRAU)
BHD393 (JRAU)
BHD415 (JRAU)
JQ039263
JQ024872
JQ039284
JQ025330
JQ025291
JQ025331
JQ024510
JQ024538
JQ024541
JQ024136
JQ024166
JQ024169
Fl. Pl. South Africa 14: 554
(1934)
Notizbl. Königl. Bot. Gart.
Berlin 4: 247 (1906)
Gard. Dict. ed. 8 22 (1768)
—
Limpopo
OM2960 (JRAU)
JQ039244
JQ025310
—
JQ024109
—
Limpopo, Botswana
OM1621 (JRAU)
JQ039259
JQ025301
—
—
Aloe candelabrum A.Berger [nom. illegit.], Aloe ferox
var. galpinii (Baker) Reynolds, Aloe ferox var. incurva
Baker, Aloe ferox var. subferox (Spreng.) Baker, Aloe
galpinii Baker, Aloe horrida Haw., Aloe muricata Haw.,
Aloe pallancae Guillaumin [nom. inva.], Aloe perfoliata
var. ferox (Mill.) Aiton, Aloe pseudoferox Salm-Dyck,
Aloe subferox Spreng., Aloe supralaevis var.
erythrocarpa Baker, Busipho ferox (Mill.) Salisb. [nom.
inva.], Pachidendron ferox (Mill.) Haw., Pachidendron
pseudoferox (Salm-Dyck) Haw., Pachidendron
supralaeve (Haw.) Haw.
Eastern and Western Cape,
Kwazulu-Natal, Limpopo
BHD407 (JRAU)
JQ039260
JQ025327
—
—
Trans. Roy. Soc. South Africa
5: 707 (1917)
Fl. Cap. 6: 327 (1896)
J. Linn. Soc., Bot. 18: 180
(1880)
—
Limpopo, Mpumalanga
OM2959 (JRAU)
JQ039276
JQ025300
JQ024529
JQ024157
Aloe nitens Baker [nom. illegit.]
—
Kwazulu-Natal, Swaziland
Kwazulu-Natal
BHD468 (JRAU)
BHD411 (JRAU)
JQ039280
JQ039285
JQ025317
JQ025319
—
JQ024542
—
JQ024170
J. S. African Bot. 37: 37 (1971)
Aloe 41: 36 (2004)
—
—
Northern Cape
Western Cape
BHD402 (JRAU)
BHD378 (JRAU)
JQ024865
JQ024867
JQ025263
JQ025264
JQ024494
JQ024518
JQ024118
JQ024144
Fl. Pl. South Africa 14: 558
(1934)
Trans. Linn. Soc. London 7: 18
(1804)
S. African J. Bot. 53: 491
(1987)
—
Eastern Cape
BHD379 (JRAU)
JQ024869
JQ025265
JQ024532
JQ024160
—
Eastern and Western Cape
BHD400 (JRAU)
JQ024870
JQ025260
JQ024534
JQ024162
Aloe karasbergensis Pillans
Northern Cape
BHD408 (JRAU)
JQ039283
JQ025306
JQ024536
JQ024164
27
!
Chapter two: Material and Methods
!
!
Aloe striata Haw. subsp.
komaggasensis (Kritz. and van
Jaarsv.) Glen and D.S.Hardy
S. African J. Bot. 53: 491
(1987)
Aloe komaggasensis Kritz. and van Jaarsv.
Northern Cape
BHD417 (JRAU)
JQ024871
JQ025261
JQ024537
JQ024165
S. African J. Bot. 53: 490
(1987)
Aloe barbertoniae Pole-Evans, Aloe comosibracteata
Reynolds, Aloe davyana Schönland, Aloe davyana var.
subolifera Groenew., Aloe graciliflora Groenew., Aloe
labiaflava Groenew., Aloe longibracteata Pole-Evans,
Aloe mutans Reynolds, Aloe verdoorniae Reynolds
Free State, Gauteng,
Kwazulu-Natal, Limpopo,
Mpumalanga, North-West,
Swaziland
BHD380 (JRAU)
JQ039264
JQ025304
JQ024512
JQ024138
Section Proliferae
Aloe brevifolia Mill.
Gard. Dict. Abr. ed. 6 8 (1771)
Aloe brevifolia Mill. var. postgenita (Schult. and
Schult.f.) Baker
Western Cape
BHD460 (JRAU)
JQ039248
JQ025314
—
—
Section Purpurascentes
Aloe microstigma Salm-Dyck
Aloes Mesembr. 2: 26 (1849)
—
BHD392 (JRAU)
JQ039272
JQ025323
JQ024525
JQ024152
Aloe succotrina Weston
Encycl. 1: 85 (1783)
Aloe perfoliata L. var. purpurascens Aiton, Aloe
perfoliata L. var. succotrina (Lam.) Aiton, Aloe
purpurascens (Aiton) Haw., Aloe sinuata Thunb., Aloe
sinuata Willd. [nom. illegit.], Aloe soccotorina Schult.
and Schult.f., Aloe soccotrina Garsault, Aloe soccotrina
DC. var. purpurascens (Aiton) Ker Gawl., Aloe
succotrina Lam. var. saxigena A.Berger, Aloe vera Mill.
[nom. illegit.]
Northern and Western Cape,
Namibia
Western Cape
BHD424 (JRAU)
JQ024873
JQ025266
JQ024539
JQ024167
Bot. Jahrb. Syst. 38: 86 (1905)
—
Western Cape
BHD385 (JRAU)
JQ039253
JQ025328
JQ024499
JQ024124
Gard. Dict. ed. 8 16 (1768)
Aloe glauca Mill. var. elatior Salm-Dyck, Aloe glauca
Mill. var. humilior Salm-Dyck, Aloe glauca Mill. var.
major Haw., Aloe glauca Mill. var. minor Haw., Aloe
glauca Mill. var. muricata (Schult.) Baker, Aloe glauca
Mill. var. spinosior Haw., Aloe muricata Schult. [nom.
illegit.], Aloe perfoliata var. glauca (Mill.) Aiton, Aloe
rhodacantha DC.
Aloe perfoliata var. lineata Aiton
Northern and Western Cape
BHD419 (JRAU)
JQ039262
JQ025313
JQ024508
JQ024134
Eastern and Western Cape
BHD416 (JRAU)
JQ039267
JQ025320
JQ024520
JQ024147
Aloe muirii Marloth
Eastern and Western Cape
BHD412 (JRAU)
BHD387 (JRAU)
JQ039269
JQ025322
JQ025321
JQ024521
JQ024522
JQ024148
JQ024149
Bradleya 16: 114 (1998)
Lomatophyllum anivoranoense Rauh and Hebding
North-Eastern Madagascar
BHD473 (JRAU)
JQ039245
JQ025371
—
—
J. Linn. Soc., Bot. 22: 529
(1887)
Bradleya 16: 114 (1998)
Aloinella haworthioides (Baker) Lemée, Leemea
haworthioides (Baker) P.V.Heath
Lomatophyllum propaguliferum Rauh and Razaf.
Central Madagascar
BHD190 (JRAU)
JQ039265
JQ025357
JQ024513
JQ024139
Central to Eastern
Madagascar
BHD472 (JRAU)
JQ039279
JQ025359
—
—
Fl. Pl. Africa 28: 1091 (1951)
—
Zimbabwe, Mozambique
(Chimanimani Mountains)
BHD470 (JRAU)
JQ039273
JQ025302
—
—
Section Pictae
Aloe greatheadii var. davyana
(Schönland) Glen and
D.S.Hardy
Section Rhodacanthae
Aloe comosa Marloth and
A.Berger
Aloe glauca Mill.
Aloe lineata (Aiton) Haw.
Aloe lineata (Aiton) Haw.var.
muirii(Marloth) Reynolds
Madagascan Aloes
Aloe anivoranoensis (Rauh and
Hebding) L.E.Newton and
G.D.Rowley
Aloe haworthioides Baker
Aloe propagulifera (Rauh and
Razaf.) L.E.Newton and
G.D.Rowley
Taxonomic Status Unknown
Aloe munchii Christian
Trans. Linn. Soc. London 7: 18
(1804)
Aloes S. Afr. 205 (1950)
Astroloba Uitewaal
28
!
Chapter two: Material and Methods
!
!
Astroloba corrugata N.L.Mey.
and Gideon F.Sm.
Bothalia 28: 61 (1998)
Apicra aspera var. major Haw., Haworthia aspera var.
major (Haw.) Parr, Haworthia corrugata (N.L.Mey. and
Gideon F.Sm.) M.Hayashi
Western Cape
BHD219 (JRAU)
JQ039290
JQ025350
JQ024545
JQ024173
Astroloba foliolosa (Haw.)
Uitewaal
Succulenta (Netherlands) 28:
54 (1947)
Aloe foliolosa Haw., Apicra foliolosa (Haw.) Willd.,
Astroloba smutsiana nom. prov. Reinecke, Astroloba
spiralis subsp. foliolosa (Haw.) L.E.Groen, Haworthia
foliolosa (Haw.) Haw.
Eastern Cape
BHD228 (JRAU)
JQ039291
JQ025351
JQ024547
JQ024175
Astroloba herrei Uitewaal
Desert Pl. Life 20: 37 (1948)
Astroloba dodsoniana Uitewaal, Haworthia dodsoniana
(Uitewaal) Parr, Haworthia harlandiana Parr
Western Cape
BHD225 (JRAU)
JQ039292
JQ025349
JQ024548
JQ024176
Astroloba rubriflora (L.Bolus)
Gideon F.Sm. and J.C.Manning
Bothalia 30: 53 (2000)
Aloe rubriflora (L.Bolus) G.D.Rowley, Apicra
jacobseniana Poelln., Apicra rubriflora L.Bolus,
Haworthia rubriflora (L.Bolus) Parr, Poellnitzia
rubriflora (L.Bolus) Uitewaal, Poelnitzia rubriflora var.
jacobseniana (Poelln.) Uitewaal
Western Cape
BHD229 (JRAU)
JQ039293
JQ025297
JQ024549
JQ024177
Pflanzenr. IV, 38: 73 (1908)
Catevala angolensis (Baker) Kuntze, Chortolirion
bergerianum Dinter, Chortolirion stenophyllum Berger,
Chortolirion subspicatum (Baker) A.Berger,
Chortolirion tenuifolium (Engl.) A.Berger, Chortolirion
tenuifolium Berger, Haworthia angolensis Baker,
Haworthia stenophylla Baker, Haworthia subspicata
Baker, Haworthia tenuifolia Engl.
Eastern and Northern Cape,
Free State, Gauteng,
Kwazulu-Natal, Limpopo,
Mpumalanga, North West,
Botswana, Namibia,
Swaziland
BHD466 (JRAU)
JQ039295
JQ025344
—
—
Philos. Mag. Ann. Chem. 2:
351 (1827)
Aloe 41: 81 (2004)
Aloe disticha L., Ptys disticha (L.) Salisb. [nom. inva.]
Western Cape
OM2778 (JRAU)
JQ024879
JQ025278
JQ024560
JQ024187
—
Eastern Cape
OM2793 (JRAU)
JQ024880
JQ025279
JQ024561
JQ024188
Bradleya 9: 100 (1991)
Pl. Succ. Horto Alencon. 6
(1809)
—
Aloe bicolor (Haw.) Schult. and Schult.f., Aloe boureana
Schult. and Schult.f., Aloe dictyodes Schult. and
Schult.f., Aloe formosa (Haw.) Schult. and Schult.f.,
Aloe guttata Salm-Dyck, Aloe lingua Ker Gawl. [nom.
illegit.], Aloe maculata Thunb. [nom. illegit.], Aloe
maculata var. obliqua Aiton, Aloe marmorata Steud.,
Aloe nigricans var. fasciata Salm-Dyck, Aloe obliqua
(Aiton) Haw. [nom. illegit.], Aloe planifolia Baker [nom.
illegit.], Aloe vittata Schult. and Schult.f., Aloe zeyheri
Salm-Dyck, Gasteria bicolor Haw., Gasteria bicolor
var. fallax (Haw.) van Jaarsv., Gasteria bicolor var.
liliputana (Poelln.) van Jaarsv., Gasteria biformis
Poelln., Gasteria caespitosa Poelln., Gasteria
chamaegigas Poelln., Gasteria colubrina N.E.Br.,
Gasteria fasciata (Salm-Dyck) Haw., Gasteria formosa
Haw., Gasteria herreana Poelln., Gasteria kirsteana
Poelln., Gasteria liliputana Poelln., Gasteria lingua (Ker
Gawl.) A.Berger [nom. illegit.], Gasteria loeriensis
Poelln., Gasteria longiana Poelln., Gasteria
longibracteata Poelln., Gasteria maculata Haw.,
Gasteria maculata var. dregeana A.Berger, Gasteria
maculata var. fallax Haw., Gasteria marmorata Baker,
Gasteria multiplex Poelln., Gasteria nigricans var.
Eastern Cape
Eastern Cape
OM2772 (JRAU)
OM2788 (JRAU)
JQ024884
JQ024886
JQ025283
JQ025274
JQ024566
JQ024569
JQ024193
JQ024196
Chortolirion A.Berger
Chortolirion angolense (Baker)
A.Berger
Gasteria Duval
Section Gasteria
Gasteria disticha (L.) Haw.
Gasteria doreeniae van Jaarsv.
and A.E.van Wyk
Gasteria glomerata van Jaarsv.
Gasteria obliqua (Aiton) Duval
29
!
Chapter two: Material and Methods
!
!
fasciata (Salm-Dyck) Haw., Gasteria picta Haw.,
Gasteria planifolia (Baker) Baker, Gasteria retata Haw.,
Gasteria multiplex Poelln., Gasteria nigricans var.
fasciata (Salm-Dyck) Haw., Gasteria picta Haw.,
Gasteria planifolia (Baker) Baker, Gasteria retata Haw.,
Gasteria salmdyckiana Poelln., Gasteria spiralis Baker,
Gasteria spiralis var. tortulata Baker, Gasteria
variolosa Baker, Gasteria zeyheri (Salm-Dyck) Baker
Gasteria pillansii Kensit var.
ernesti-ruschii (Dinter and
Poelln.) van Jaarsv.
Gasteria pillansii Kensit var.
pillansii
Gasteria rawlinsonii Oberm.
Section Longiflorae Haw.
Gasteria acinacifolia (J.Jacq.)
Haw.
Aloe 29: 17 (1992)
Gasteria ernesti-ruschii Dinter and Poelln.
Northern Cape, Namibia
OM2779 (JRAU)
—
JQ025285
JQ024570
JQ024197
Unknown
Gasteria neliana Poelln.
Northern Cape
OM2781 (JRAU)
JQ024874
JQ025284
JQ024553
JQ024180
Fl. Pl. Africa 43: t. 1701 (1976)
—
Eastern Cape
OM2775 (JRAU)
JQ024889
JQ025288
JQ024573
JQ024200
Suppl. Pl. Succ. 49 (1819)
Aloe acinacifolia J.Jacq., Aloe acinacifolia var. minor
Salm-Dyck, Aloe candicans (Haw.) Schult. and Schult.f.,
Aloe ensifolia (Haw.) Schult. and Schult.f., Aloe nitens
Schult. and Schult.f., Aloe pluripunctata Schult. and
Schult.f., Aloe venusta Schult. and Schult.f., Gasteria
acinacifolia var. ensifolia (Haw.) Baker, Gasteria
acinacifolia var. nitens (Haw.) Baker, Gasteria
acinacifolia var. pluripunctata (Haw.) Baker, Gasteria
acinacifolia var. venusta (Haw.) Baker, Gasteria
candicans Haw., Gasteria ensifolia Haw., Gasteria
fuscopunctata Baker, Gasteria huttoniae N.E.Br.,
Gasteria inexpectata Poelln., Gasteria linita Haw.,
Gasteria lutzii Poelln., Gasteria nitens Haw., Gasteria
pluripunctata Haw., Gasteria venusta Haw.
Eastern Cape
OM2790 (JRAU)
JQ024875
JQ025271
JQ024554
JQ024181
Gasteria batesiana var.
dolomitica van Jaarsv. and Van
Wyk
Aloe 36(4) 74 (1999)
—
Mpumalanga
OM2777 (JRAU)
JQ024876
JQ025273
JQ024555
JQ024182
30
!
Chapter two: Material and Methods
!
!
Gasteria carinata (Mill.) Duval
var. carinata
Unknown
Gasteria carinata var. retusa
van Jaarsv.
Aloe 29: 15 (1992)
Gasteria croucheri (Hook.f.)
Baker
Gasteria ellaphieae van Jaarsv.
J. Linn. Soc., Bot. 18: 196
(1880)
Cact. Succ. J. (Los Angeles) 63:
3 (1991)
J. Linn. Soc., Bot. 18: 195
(1880)
Cact. Succ. J. (Los Angeles) 70:
65 (1998)
Aloe 29: 12 (1992)
Gasteria excelsa Baker
Gasteria glauca van Jaarsv.
Gasteria nitida var. armstrongii
(Schönland) van Jaarsv.
Gasteria polita van
Jaarsv.(TYPE)
Gasteria pulchra (Aiton) Haw.
Gasteria tukhelensis van
Jaarsv.
Gasteria vlokii van Jaarsv.
Aloe angulata Willd., Aloe angulata var. truncata
Willd., Aloe carinata Ker Gawl., Aloe carinata var.
subglabra Haw., Aloe excavata Willd., Aloe glabra
(Haw.) Salm-Dyck, Aloe laetepunctata (Haw.) Schult.
and Schult.f., Aloe laevis Salm-Dyck, Aloe lingua var.
angulata Haw., Aloe lingua var. multifaria Haw., Aloe
linguiformis DC. [nom. illegit.], Aloe pseudoangulata
Salm-Dyck, Aloe pusilla Schult. and Schult.f., Aloe
subcarinata Salm-Dyck, Aloe sulcata Salm-Dyck, Aloe
tristicha Medik., Aloe undata Schult. and Schult.f.,
Gasteria angulata (Haw.) Duval, Gasteria angulata
(Willd.) Haw. [nom. illegit.], Gasteria angulata var.
truncata (Willd.) A.Berger, Gasteria bijliae Poelln.,
Gasteria carinata var. falcata A.Berger, Gasteria
carinata var. glabra (Salm-Dyck) van Jaarsv., Gasteria
carinata var. latifolia A.Berger, Gasteria carinata var.
parva (Haw.) Baker, Gasteria carinata var. strigata
(Haw.) Baker, Gasteria disticha var. angulata (Willd.)
Baker, Gasteria excavata (Willd.) Haw., Gasteria
glabra Haw., Gasteria humilis Poelln., Gasteria
laetepunctata Haw., Gasteria laevis (Salm-Dyck) Haw.,
Gasteria pallescens Baker, Gasteria parva Haw.,
Gasteria parvifolia Baker, Gasteria patentissima
Poelln., Gasteria porphyrophylla Baker, Gasteria
schweickerdtiana Poelln., Gasteria strigata Haw.,
Gasteria subcarinata (Salm-Dyck) Haw., Gasteria
sulcata (Salm-Dyck) Haw., Gasteria trigona var.
kewensis A.Berger, Gasteria undata Haw.
Gasteria retusa (van Jaarsv.) van Jaarsv.
Western Cape
OM2780 (JRAU)
JQ039297
JQ025275
—
—
Western Cape
OM2798 (JRAU)
JQ024877
JQ025276
JQ024556
JQ024183
Aloe croucheri Hook.f., Gasteria disticha var. natalensis
Baker
—
Eastern Cape, KwazuluNatal
Eastern Cape
OM2773 (JRAU)
OM2791 (JRAU)
OM2782 (JRAU)
JQ024878
JQ025277
JQ024186
JQ024881
JQ025280
JQ024559
JQ024558
JQ024562
—
Eastern Cape
OM2789 (JRAU)
JQ024882
JQ025281
JQ024564
JQ024191
—
Eastern Cape
OM2771 (JRAU)
JQ024883
JQ025282
JQ024565
JQ024192
Gasteria armstrongii Schönland, Gasteria armstrongii
Schoenland
—
Eastern Cape
OM2792 (JRAU)
JQ024885
JQ025272
JQ024567
JQ024194
Western Cape
OM2784 (JRAU)
JQ024887
JQ025286
JQ024571
JQ024198
Aloe maculata var. pulchra Aiton, Aloe obliqua DC.,
Aloe pulchra (Aiton) Jacq., Gasteria poellnitziana
H.Jacobsen [nom. inva.]
Eastern Cape
OM2785 (JRAU)
JQ024888
JQ025287
JQ024572
JQ024199
Bothalia 35: 164 (2005)
—
Kwazulu-Natal
OM2774 (JRAU)
JQ024890
JQ025289
JQ024574
JQ024201
Cact. Succ. J. (Los Angeles) 59:
170 (1987)
—
Western Cape
OM2786 (JRAU)
JQ039298
JQ025298
JQ024575
JQ024202
Cact. Succ. J. (Los Angeles) 73:
127 (2001)
Syn. Pl. Succ. 86 (1812)
JQ024189
Haworthia Duval subgenus
Haworthia
31
!
Chapter two: Material and Methods
!
!
Haworthia angustifolia Haw.
Philos. Mag. J. 67:283 (1825)
Catevala angustifolia (Haw.) Kuntze, Haworthia
chloracantha var. angustifolia (Haw.) Halda, Haworthia
chloracantha subsp. angustifolia (Haw.) Halda
Eastern Cape
BHD234 (JRAU)
JQ039299
—
JQ024593
JQ024219
Haworthia arachnoidea (L.)
Duval (TYPE)
Pl. Succ. Horto Alencon. 7
(1809)
Aloe arachnoidea (L.) Burm.f., Aloe pumila var.
arachnoidea L., Apicra arachnoides (L.) Willd.,
Catevala arachnoidea (L.) Medik., Haworthia pallida
var. paynei L. Bolus.
Western Cape
BHD029 (JRAU)
BHD030 (JRAU)
JQ024891
JQ024892
—
JQ024601
JQ024602
JQ024226
Haworthia bayeri J.D.Venter
and S.A.Hammer
Haworthia blackburniae
W.F.Barker
Cact. Succ. J. (Los Angeles) 69:
75 (1997)
J. S. African Bot. 3: 93 (1937)
—
Eastern and Western Cape
BHD110 (JRAU)
JQ039301
JQ025360
JQ024615
JQ024239
Haworthia blackburniae Poelln. [nom. illegit.]
Western Cape
BHD198 (JRAU)
JQ024893
JQ025226
JQ024616
JQ024240
Haworthia blackburniae
W.F.Barker var. derustensis
M.B.Bayer
Haworthia Revisited 41 (1999)
Haworthia derustensis (M.B.Bayer) M.Hayashi
Western Cape
BHD243 (JRAU)
JQ039302
JQ025361
JQ024617
JQ024241
Haworthia blackburniae
W.F.Barker var. graminifolia
(G.G.Sm.) M.B.Bayer
Haworthia Revisited 42 (1999)
Haworthia blackburniae subsp. graminifolia (G.G.Sm.)
Halda, Haworthia graminifolia G.G.Sm.
Western Cape
BHD201 (JRAU)
JQ039303
JQ025362
JQ024618
JQ024242
Haworthia chloracantha Haw.
Saxifrag. Enum. 2: 57 (1821)
Western Cape
BHD075 (JRAU)
JQ039305
JQ025363
JQ024625
JQ024249
Haworthia cooperi Baker
Haworthia cooperi Baker var.
doldii M.B.Bayer
Haworthia cymbiformis (Haw.)
Duval
Haworthia cymbiformis (Haw.)
Duval var. ramosa (G.G.Sm.)
M.B.Bayer
Haworthia cymbiformis (Haw.)
Duval var. setulifera (Poelln.)
M.B.Bayer
Refug. Bot. 4: t. 233 (1870)
Haworthiad 16: 65 (2002)
Aloe chloracantha (Haw.) Schult. and Schult.f.,
Catevala chloracantha (Haw.) Kuntze
Catevala cooperi (Baker) Kuntze
Haworthia doldii (M.B.Bayer) M.Hayashi
Eastern Cape
Eastern Cape
BHD133 (JRAU)
BHD209 (JRAU)
JQ024895
JQ024896
JQ025227
JQ025228
JQ024631
JQ024634
JQ024255
JQ024258
Pl. Succ. Horto Alencon. 7
(1809)
Haworthia Revisited 60 (1999)
Aloe cymbiformis Haw., Catevala cymbiformis (Haw.)
Kuntze, Haworthia planifolia
Haworthia cymbiformis f. ramosa (G.G.Sm.)
M.B.Bayer, Haworthia ramosa G.G.Sm.
Eastern Cape
BHD322 (JRAU)
JQ024898
JQ025229
JQ024645
JQ024269
Eastern Cape
BHD321 (JRAU)
JQ024897
JQ025231
JQ024644
JQ024268
Haworthia Revisited 62 (1999)
Haworthia cymbiformis f. obesa (Poelln.) Pilbeam,
Haworthia cymbiformis var. obesa Poelln., Haworthia
planifolia var. setulifera Poelln.
Eastern Cape
BHD320 (JRAU)
BHD325 (JRAU)
JQ024899
JQ024900
JQ025230
JQ025232
JQ024648
JQ024649
JQ024272
JQ024273
Haworthia decipiens Poelln.
Repert. Spec. Nov. Regni Veg.
28: 103 (1930)
Haworthia Revisited 67(1999)
—
Eastern and Western Cape
BHD132 (JRAU)
JQ024901
JQ025233
JQ024652
JQ024276
Eastern Cape
BHD196 (JRAU)
JQ024902
JQ025250
JQ024654
JQ024278
Haworthiad 16: 63 (2002)
Haworthia bolusii var. pringlei (C.L. Scott) M.B.Bayer,
Haworthia pringlei C.L.Scott
Haworthia virella (M.B.Bayer) M.Hayashi
BHD328 (JRAU)
JQ024903
JQ025234
JQ024656
JQ024280
Repert. Spec. Nov. Regni Veg.
42: 271 (1937)
Haworthia retusa subsp. emelyae (Poelln.) Halda,
Haworthia retusa var. emelyae (Poelln.) Halda
Western Cape
BHD105 (JRAU)
JQ024904
JQ025236
JQ024661
JQ024285
Haworthia emelyae Poelln. var.
comptoniana (G.G.Sm.)
J.D.Venter and S.A.Hammer
Cact. Succ. J. (Los Angeles) 69:
77 (1997)
Haworthia comptoniana G.G.Sm., Haworthia retusa
var. comptoniana (G.G.Sm.) Halda
Eastern Cape
BHD220 (JRAU)
JQ039307
JQ025364
JQ024663
JQ024287
Haworthia floribunda Poelln.
Repert. Spec. Nov. Regni Veg.
40: 149 (1936)
Cact. Succ. J. Gr. Brit. 7: 40
(1938)
Haworthia Revisited 86 (1999)
Haworthia chloracantha var. floribunda (Poelln.) Halda
Western Cape
BHD077 (JRAU)
JQ024906
JQ025251
JQ024666
JQ024290
Aloe herbacea Mill.
Western Cape
BHD237 (JRAU)
—
—
JQ024685
JQ024307
Haworthia pallida var. flaccida (M.B.Bayer) M.Hayashi
Western Cape
BHD333 (JRAU)
JQ024907
JQ025252
JQ024686
JQ024308
Haworthia Revisited 87 (1999)
Haworthia pallida var. paynei (Poelln.) Poelln.,
Haworthia paynei Poelln.
Western Cape
BHD373 (JRAU)
JQ024908
JQ025254
JQ024687
JQ024309
Haworthia decipiens var.
pringlei (C.L.Scott) M.B.Bayer
Haworthia decipiens Poelln.
var. virella M.B.Bayer
Haworthia emelyae Poelln.
Haworthia herbacea (Mill.)
Stearn
Haworthia herbacea (Mill.)
Stearn var. flaccida M.B.Bayer
Haworthia herbacea (Mill.)
Stearn var. paynei (Poelln.)
32
!
Chapter two: Material and Methods
!
!
M.B.Bayer
Haworthia lockwoodii
Archibald
Haworthia maculata (Poelln.)
M.B.Bayer
Fl. Pl. South Africa 20: t. 792
(1940)
Haworthia Handb. 130 (1976)
Haworthia mucronata subsp. lockwoodii (Archibald)
Halda
Haworthia intermedia var. maculata (Poelln.)
Esterhuizen, Haworthia schuldtiana var. maculata
Poelln.
Haworthia dekenahii G.G.Sm., Haworthia retusa var.
dekenahii (G.G.Sm.) M.B.Bayer
Western Cape
BHD213 (JRAU)
—
JQ025378
JQ024711
JQ024336
Western Cape
BHD203 (JRAU)
JQ024911
JQ025237
JQ024715
JQ024340
Haworthia magnifica Poelln.
var. dekenahii (G.G.Sm.)
M.B.Bayer
Haworthia marumiana
Uitewaal var. archeri
(W.F.Barker ex M.B.Bayer)
M.B.Bayer
Haworthia marxii Gildenh.
Aloe 34: 6 (1997)
Western Cape
BHD092 (JRAU)
JQ024912
JQ025238
JQ024716
JQ024341
Haworthia Revisited 104
(1999)
Haworthia archeri W.F.Barker ex M.B.Bayer
Northern and Western Cape
BHD235 (JRAU)
JQ024913
JQ025248
JQ024727
JQ024352
Haworthia mirabilis (Haw.)
Haw.
Aloe 44: 4 (2007)
—
Western Cape
BHD339 (JRAU)
JQ024914
JQ025249
JQ024728
JQ024353
Syn. Pl. Succ. 95 (1812)
Aloe mirabilis Haw., Apicra mirabilis (Haw.) Willd.,
Catevala mirabilis (Haw.) Kuntze, Haworthia
beukmanii, Haworthia retusa var. mirabilis (Haw.)
Halda, Haworthia willowmorensis Poelln.
Western Cape
BHD032 (JRAU)
JQ039317
JQ025365
JQ024650
JQ024274
Haworthia mirabilis (Haw.)
Haw. var. paradoxa (Poelln.)
M.B.Bayer
Haworthia mirabilis var.
calcarea M.B.Bayer
Aloe 34: 6 (1997)
Haworthia magnifica var. paradoxa (Poelln.)
M.B.Bayer, Haworthia paradoxa Poelln.
Western Cape
BHD044 (JRAU)
JQ024915
JQ025254
JQ024749
JQ024373
Haworthia Revisited 110
(1999)
Western Cape
BHD232 (JRAU)
JQ024916
JQ025246
JQ024771
JQ024397
Haworthia mirabilis var.
triebneriana (Poelln.)
M.B.Bayer
Haworthia Revisited 113
(1999)
Haworthia calcarea (M.B.Bayer) M.Hayashi,
Haworthia rossouwii var. calcarea (M.B.Bayer)
M.B.Bayer
Haworthia mirabilis f. rubrodentata (Triebner and
Poelln.) Pilbeam, Haworthia nitidula Poelln., Haworthia
rossouwii Poelln., Haworthia triebneriana Poelln.,
Haworthia triebneriana var. depauperata Poelln.,
Haworthia triebneriana var. multituberculata Poelln.,
Haworthia triebneriana var. napierensis Triebner and
Poelln., Haworthia triebneriana var. pulchra Poelln.,
Haworthia triebneriana var. rubrodentata Triebner and
Poelln., Haworthia triebneriana var. subtuberculata
Poelln., Haworthia triebneriana var. turgida Triebner,
Haworthia willowmorensis Poelln.
Western Cape
BHD020 (JRAU)
JQ024917
—
JQ024773
JQ024399
Haworthia monticola Fourc.
var. asema M.B.Bayer
Haworthia mucronata Haw.
var. habdomadis (Poelln.)
M.B.Bayer
Haworthia mucronata Haw.
var. morrisiae (Poelln.)
M.B.Bayer
Haworthia Revisited 117
(1999)
Haworthia Revisited 120
(1999)
Haworthia asema (M.B.Bayer) M.Hayashi
Western Cape
BHD251 (JRAU)
JQ024918
JQ025255
JQ024780
JQ024405
Haworthia habdomadis Poelln., Haworthia inconfluens
var. habdomadis (Poelln.) M.B.Bayer
Western Cape
BHD347 (JRAU)
—
JQ025379
JQ024784
JQ024409
Haworthia altilinea var. morrisiae Poelln., Haworthia
altilinea f. subglauca Poelln., Haworthia habdomadis
var. morrisiae (Poelln.) M.B.Bayer, Haworthia
inconfluens var. morrisiae (Poelln.) M.B.Bayer,
Haworthia mucronata f. subglauca (Poelln.) Poelln.,
Haworthia sakaii M.Hayashi
Western Cape
BHD349 (JRAU)
JQ024920
JQ025240
JQ024785
JQ024410
Repert. Spec. Nov. Regni Veg.
49: 29 (1940)
33
!
Chapter two: Material and Methods
!
!
Haworthia mucronata var.
bijliana (Poelln.) ined.
Unknown
Haworthia altilinea f. inconfluens Poelln., Haworthia
bijliana Poelln., Haworthia bijliana var. joubertii
Poelln., Haworthia fergusoniae Poelln., Haworthia
habdomadis var. inconfluens (Poelln.) M.B.Bayer,
Haworthia inconfluens (Poelln.) M.B.Bayer, Haworthia
mucronata var. inconfluens (Poelln.) M.B.Bayer,
Haworthia mucronata f. inconfluens (Poelln.) Poelln.),
Haworthia setata var. bijliana (Poelln.) Poelln.
Western Cape
BHD350 (JRAU)
JQ024919
JQ025239
JQ024782
JQ024407
Haworthia mucronata var.
rycroftiana (M. B. Bayer) M.B.
Bayer
Haworthia mutica Haw.
Haworthia Revisited 124
(1999)
Haworthia rycroftiana M.B.Bayer
Western Cape
BHD345 (JRAU)
JQ024921
JQ025241
JQ024787
JQ024412
Saxifrag. Enum. 2: 55 (1821)
Western Cape
BHD102 (JRAU)
JQ024922
JQ025242
JQ024797
JQ024421
Haworthia mutica Haw. var.
nigra M.B.Bayer
Haworthia outeniquensis
M.B.Bayer
Haworthia pulchella
M.B.Bayer var. globifera
M.B.Bayer
Haworthia pygmaea Poelln.
Haworthia Revisited 126
(1999)
Haworthia Revisited 130
(1999)
Haworthia Revisited 136
(1999)
Aloe mutica (Haw.) Schult. and Schult.f., Haworthia
retusa var. mutica (Haw.) Halda
Haworthia silviae var. nigra (M.B.Bayer) M.Hayashi
Western Cape
BHD072 (JRAU)
JQ024923
JQ025243
JQ024798
JQ024422
—
Western Cape
BHD356 (JRAU)
JQ024924
JQ025256
JQ024807
JQ024431
Haworthia globifera (M.B.Bayer) M.Hayashi
Western Cape
BHD206 (JRAU)
JQ024925
JQ025257
JQ024813
JQ024437
Repert. Spec. Nov. Regni Veg.
27: 132 (1929)
—
Western Cape
BHD358 (JRAU)
JQ039320
JQ025333
JQ024816
JQ024440
Haworthia reticulata (Haw.)
Haw.
Syn. Pl. Succ. 94 (1812)
Western Cape
BHD117 (JRAU)
JQ024927
JQ025244
JQ024819
JQ024443
Haworthia retusa (L.) Duval
Pl. Succ. Horto Alencon. 7
(1809)
Haworthia Handb. 153 (1976)
Aloe reticulata Haw., Apicra reticulata (Haw.) Willd.,
Catevala reticulata (Haw.) Kuntze, Haworthia
haageana
Aloe retusa L., Apicra retusa (L.) Willd., Catevala
retusa (L.) Medik
Haworthia arachnoideae var. semiviva (Poelln.) Halda,
Haworthia bolusii var. semiviva Poelln.
JQ024928
JQ025245
JQ024832
Northern and Western Cape
BHD027 (JRAU)
BHD120 (JRAU)
BHD360 (JRAU)
JQ024929
JQ025247
JQ024844
JQ024456
JQ024455
JQ024467
Haworthia semiviva (Poelln.)
M.B.Bayer
Haworthia springbokvlakensis
C.L.Scott
Haworthia truncata Schönland
J. S. African Bot. 36: 288
(1970)
Trans. Roy. Soc. South Africa
1: 291 (1910)
Haworthia retusa var. springbokvlakensis (C.L.Scott)
Halda
—
Eastern Cape
BHD362 (JRAU)
—
—
JQ024847
JQ024470
Western Cape
BHD210 (JRAU)
JQ039323
JQ025375
JQ024848
JQ024471
Haworthia variegata L.Bolus
J. Bot. 67: 137 (1929)
Western Cape
BHD367 (JRAU)
JQ039324
JQ025376
JQ024850
JQ024473
Haworthia vlokii M.B.Bayer
Haworthia Revisited 160
(1999)
J. S. African Bot. 8: 245 (1942)
Haworthia chloracantha var. variegata (L.Bolus) Halda,
Haworthia chloracantha subsp. variegata (L.Bolus)
Halda
—
Western Cape
BHD249 (JRAU)
JQ024930
JQ025258
JQ024858
JQ024481
—
Western Cape
BHD200 (JRAU)
JQ024931
JQ025259
JQ024859
JQ024482
Repert. Spec. Nov. Regni Veg.
41: 217 (1937)
Haworthia chloracantha var. zantheriana (Poelln.)
Halda
Eastern Cape
BHD230 (JRAU)
JQ039326
JQ025370
JQ024860
JQ024483
Syn. Pl. Succ. 92 (1812)
Aloe attenuata Haw., Apicra attenuata (Haw.) Willd.,
Catevala attenuata (Haw.) Kuntze, Haworthia attenuata
var. britteniae Poelln., Haworthia attenuata var.
clariperla (Haw.) M.B.Bayer, Haworthia pumila subsp.
attenuata (Haw.) Halda
Eastern Cape
BHD253 (JRAU)
BHD264 (JRAU)
BHD265 (JRAU)
JQ039300
JQ025311
JQ024610
JQ024609
JQ024608
JQ024234
JQ024233
JQ024232
J. S. African Bot. 47: 789
(1981)
Haworthia retusa var. bruynsii (M.B.Bayer) Halda
Eastern Cape
BHD374 (JRAU)
JQ039304
JQ025334
JQ024622
JQ024246
Haworthia wittebergensis
W.F.Barker
Haworthia zantneriana Poelln.
Haworthia subgenus
Hexangulares Uitewaal ex
M.B.Bayer
Haworthia attenuata Haw.
Haworthia bruynsii M.B.Bayer
34
!
Chapter two: Material and Methods
!
!
Haworthia coarctata Haw.
Philos. Mag. J. 66: 301 (1824)
Aloe coarctata (Haw.) Schult. and Schult.f., Catevala
coarctata (Haw.) Kuntze, Haworthia reinwardtii var.
coarctata (Haw.) Halda, Haworthia reinwardtii subsp.
coarctata (Haw.) Halda
Eastern Cape
BHD327 (JRAU)
JQ024894
JQ025296
JQ024629
JQ024253
Haworthia coarctata var.
adelaidensis (Poelln.)
M.B.Bayer
Haworthia Revisited 172
(1999)
Haworthia coarctata subsp. adelaidensis (Poelln.)
M.B.Bayer, Haworthia coarctata f. bellula (G.G.Sm.)
Pilbeam, Haworthia reinwardtii var. adelaidensis
Poelln., Haworthia reinwardtii var. bellula G.G.Sm.,
Haworthia reinwardtii var. riebeekensis G.G.Sm.
Eastern Cape
BHD326 (JRAU)
JQ039306
JQ025335
JQ024630
JQ024254
Haworthia fasciata (Willd.)
Haw.
Saxifrag. Enum. 2: 54 (1821)
Eastern Cape
BHD330 (JRAU)
BHD331 (JRAU)
JQ024905
JQ025270
JQ024664
JQ024665
JQ024288
JQ024289
Haworthia glauca (Salm-Dyck)
Baker
J. Linn. Soc., Bot. 18: 203
(1880)
Aloe fasciata (Willd.) Salm-Dyck ex Schult. and
Schult.f., Aloe fasciata var. major Salm-Dyck, Aloe
subfasciata Salm-Dyck ex Schult. and Schult.f., Apicra
fasciata Willd., Catevala fasciata (Willd.) Kuntze,
Catevala subfasciata (Salm-Dyck ex Schult. and
Schult.f.) Kuntze, Haworthia browniana Poelln.,
Haworthia fasciata f. browniana (Poelln.) M.B.Bayer,
Haworthia fasciata var. major (Salm-Dyck) Haw.,
Haworthia fasciata f. ovatolanceolata Poelln.,
Haworthia fasciata f. sparsa Poelln., Haworthia fasciata
f. subconfluens (Poelln.) Poelln., Haworthia fasciata var.
subconfluens Poelln., Haworthia fasciata f.
vanstaadensis Poelln., Haworthia fasciata f. variabilis
Poelln., Haworthia pumila subsp. fasciata (Willd.)
Halda, Haworthia subfasciata (Salm-Dyck ex Schult.
and Schult.f) Baker
Catevala glauca (Baker)Kuntze, Haworthia reinwardtii
subsp. glauca (Baker) Halda, Haworthia reinwardtii var.
glauca (Baker) Halda
Eastern Cape
BHD061 (JRAU)
BHD062 (JRAU)
BHD370 (JRAU)
JQ039308
JQ025336
JQ024673
JQ024674
JQ024676
JQ024295
JQ024296
JQ024298
Haworthia koelmaniorum
Oberm. and D.S.Hardy var.
mcmurtryi (C.L.Scott)
M.B.Bayer
Haworthia limifolia Marloth
Haworthia Revisited 181
(1999)
Haworthia mcmurtryi C.L.Scott
Mpumalanga
BHD336 (JRAU)
BHD337 (JRAU)
JQ024909
JQ024910
JQ025293
JQ025294
JQ024690
JQ024689
JQ024312
JQ024311
Trans. Roy. Soc. South Africa
1: 409 (1910)
—
Kwazulu-Natal,
Mpumalanga
JQ039311
JQ039312
JQ025341
JQ025342
Haworthia limifolia Marloth
var. ubomboensis (I.Verd.)
G.G.Sm.
Haworthia longiana Poelln.
J. S. African Bot. 16: 3 (1950)
Haworthia ubomboensis I.Verd., Haworthia
ubomboensis Verdoorn
Swaziland
BHD135 (JRAU)
BHD159 (JRAU)
BHD173 (JRAU)
BHD175 (JRAU)
BHD183 (JRAU)
BHD221 (JRAU)
JQ039313
JQ025343
JQ024702
JQ024697
JQ024694
JQ024693
JQ024707
JQ024710
JQ024326
JQ024321
JQ024317
JQ024316
JQ024331
JQ024335
Repert. Spec. Nov. Regni Veg.
41: 203 (1937)
J. Linn. Soc., Bot. 18: 203
(1880)
Haworthia pumila subsp. longiana (Poelln.) Halda
Eastern Cape
JQ039314
JQ025316
Aloe nigra (Haw.) Schult. and Schult.f., Apicra nigra
Haw., Catevala nigra (Haw.) Kuntze
Eastern and Western Cape
BHD048 (JRAU)
BHD049 (JRAU)
BHD224 (JRAU)
JQ039318
JQ025352
JQ024714
JQ024712
JQ024799
JQ024339
JQ024337
JQ024423
Haworthia reinwardtii (SalmDyck) Haw.
Haworthia reinwardtii (SalmDyck) Haw. var. brevicula
G.G.Sm.
Saxifrag. Enum. 2: 53 (1821)
Aloe reinwardtii Salm-Dyck, Catevala reinwardtii
(Salm-Dyck) Kuntze
Haworthia reinwardtii var. diminuta G.G.Sm.
Eastern Cape
BHD254 (JRAU)
JQ039321
JQ025332
JQ024817
JQ024441
Eastern Cape
BHD359 (JRAU)
JQ024926
JQ025295
JQ024818
JQ024442
Haworthia sordida Haw.
Saxifrag. Enum. 2: 51 (1821)
Aloe sordida (Haw.) Schult. and Schult.f., Catevala
sordida (Haw.) Kuntze, Haworthia scabra subsp.
sordida (Haw.) Halda, Haworthia scabra var. sordida
(Haw.) Halda
Eastern Cape
BHD205 (JRAU)
JQ039322
JQ025354
JQ024845
JQ024468
Haworthia nigra (Haw.) Baker
J. S. African Bot. 10: 11 (1944)
35
!
Chapter two: Material and Methods
!
!
Haworthia venosa (Lam.) Haw.
Haworthia venosa Haw. subsp.
granulata (Marloth) M.B.Bayer
Haworthia subgenus
Robustipedunculares
M.B.Bayer
Haworthia kingiana Poelln.
Haworthia marginata (Lam.)
Stearn
Haworthia minor (Aiton) Duval
Saxifrag. Enum. 2: 44 (1821)
Haworthia Handb. 120 (1976)
Aloe venosa Lam., Catevala venosa (Lam.) Kuntze
Haworthia granulata Marloth, Haworthia scabra subsp.
granulata (Marloth) Halda
Western Cape
Northern and Western Cape
BHD246 (JRAU)
BHD088 (JRAU)
JQ039325
—
JQ025309
JQ025377
JQ024852
JQ024853
JQ024474
JQ024475
Repert. Spec. Nov. Regni Veg.
41: 203 (1937)
Cact. Succ. J. Gr. Brit. 7: 39
(1938)
Haworthia pumila var. kingiana (Poelln.) Halda,
Haworthia subfasciata var. kingiana (Poelln.) Poelln.
Aloe albicans Haw., Aloe laevigata Schult. and Schult.f.,
Aloe marginata Lam., Aloe ramifera Schult. and
Schult.f., Aloe virescens (Haw.) Schult. and Schult.f.,
Apicra albicans (Haw.) Willd., Catevala marginata
(Lam.) Kuntze, Haworthia albicans (Haw.) Haw.,
Haworthia albicans var. virescens (Haw.) Baker,
Haworthia laevis Haw., Haworthia marginata var. laevis
(Haw.) H.Jacobsen, Haworthia marginata var. ramifera
(Haw.) H.Jacobsen, Haworthia marginata var. virescens
(Haw.) Uitewaal, Haworthia ramifera Haw., Haworthia
virescens Haw., Haworthia virescens var. minor Haw.
Western Cape
BHD335 (JRAU)
BHD435 (JRAU)
BHD439 (JRAU)
BHD026 (JRAU)
JQ039309
JQ039310
JQ039316
JQ039315
JQ025339
JQ025340
JQ025337
JQ025338
JQ024688
JQ024310
JQ024719
JQ024344
Pl. Succ. Horto Alencon. 7
(1809)
Aloe brevis Schult. and Schult.f., Aloe erecta (Haw.)
Schult. and Schult.f., Aloe erecta var. laetivirens SalmDyck, Aloe granata Schult. and Schult.f., Aloe
margaritifera var. major Aiton, Aloe margaritifera var.
minima Aiton, Aloe margaritifera var. minor Aiton, Aloe
minor (Aiton) Schult. and Schult.f., Apicra granata
Willd., Apicra maxima (Haw.) Steud., Apicra minor
(Aiton) Steud., Catevala minima (Aiton) Kuntze,
Haworthia brevis Haw., Haworthia erecta Haw.,
Haworthia granata (Willd.) Haw., Haworthia granata
var. polyphylla Haw., Haworthia major (Aiton) Duval,
Haworthia margaritifera var. corallina Baker,
Haworthia margaritifera var. erecta (Haw.) Baker,
Haworthia margaritifera var. granata (Willd.) Baker,
Haworthia minima (Aiton) Haw., Haworthia minima
var. poellnitziana (Uitewaal) M.B.Bayer, Haworthia
mutabilis Poelln., Haworthia poellnitziana Uitewaal,
Haworthia pumila subsp. minima (Aiton) Halda,
Haworthia uitewaaliana Poelln.
Western Cape
BHD341 (JRAU)
—
—
JQ024733
JQ024358
Western Cape
36
!
Chapter two: Material and Methods
!
!
Haworthia pumila (L.) Duval
Pl. Succ. Horto Alencon. 7
(1809)
Aloe arachnoides var. pumila (L.) Aiton, Aloe granata
Salm-Dyck [nom. illegit.], Aloe margaritifera (L.)
Burm.f., Aloe margaritifera var. maxima Haw., Aloe
papillosa Salm-Dyck, Aloe pumila L., Aloe pumila var.
margaritifera L., Aloe semiglabrata (Haw.) Schult. and
Schult.f., Aloe semiglabrata Salm-Dyck, Aloe
semiglabrata var. major Salm-Dyck, Aloe semiglabrata
var. maxima (Haw.) Salm-Dyck, Aloe semiglabrata var.
minor Salm-Dyck, Aloe semiglabrata var. multipapillosa
Salm-Dyck, Aloe subalbicans Salm-Dyck, Aloe
subalbicans var. acuminata Salm-Dyck, Aloe
subalbicans var. laevior Salm-Dyck, Apicra
margaritifera (L.) Willd., Catevala margaritifera (L.)
Kuntze, Catevala semiglabrata (Haw.) Kuntze,
Haworthia corallina Baker, Haworthia margaritifera
(L.) Haw., Haworthia margaritifera var. laevior (SalmDyck) Uitewaal, Haworthia margaritifera var.
semimargaritifera (Salm-Dyck) Baker, Haworthia
margaritifera var. subalbicans (Salm-Dyck) A.Berger,
Haworthia maxima (Haw.) Duval, Haworthia
semimargaritifera var. major (Salm-Dyck) Haw.,
Haworthia semimargaritifera var. maxima (Haw.) Haw.,
Haworthia semimargaritifera var. multiperla Haw.,
Tulista margaritifera (L.) Raf.
Western Cape
BHD222 (JRAU)
JQ039319
JQ025353
JQ024815
JQ024439
Sp. Pl. 310 (1753)
Anthericum amoenum Salisb. [nom. illegit.], Anthericum
intermedium Willk. [nom. inva.], Anthericum liliago var.
australe Willk., Anthericum liliago subsp. macrocarpum
Boros, Anthericum liliago f. macrocarpum (Boros) Soó,
Anthericum liliago var. multiflorum P.Küpfer,
Anthericum liliago var. sphaerocarpum P.Küpfer,
Anthericum macrocarpum Boros, Anthericum nonramosum Gilib. [nom. inva.], Liliago vulgaris C.Presl.,
Ornithogalum gramineum Lam. [nom. illegit.],
Phalangites liliago (L.) Bubani, Phalangites
acuminatum Dulac, Phalangites lilaceum St.-Lag.,
Phalangites liliaginoides Schltdl., Phalangites liliago
(L.) Schreb., Phalangites renarnii Booth ex Schltdl.
Europe, Turkey
Chase515 (K)
—
—
—
—
Bonplandia (Hannover) 10:
370 (1862)
Tecophilaea cyanocrocus var. leichtlinii Regel,
Tecophilaea cyanocrocus var. regelii Baker,
Tecophilaea cyanocrocus var. violacea, Zephyra
cyanocrocus (Leyb.) Ravenna
Dicolus caerulescens Phil., Zephyra amoena Miers
Chile
Chase447 (K)
—
—
—
—
Chile
Chase1575 (K)
—
—
—
—
Asphodelus luteus L., Asphodelus sibiricus Schult. and
Schult.f. [nom. illegit.], Dorydium luteum (L.) Salisb.
[nom. inva.]
South-eastern Europe to
Turkey
UCI Arb.3440
JQ039289
—
—
—
OUTGROUP TAXA
Asparagaceae
Anthericum liliago L.
Tecophilaeaceae
Tecophilaea cyanocrocus Leyb.
Zephyra elegans D.Don
Edinburgh New Philos. J.
1832(Oct.): 236 (1832)
Xanthorrhoeaceae
Asphodeline lutea (L.) Reichb.
Fl. Germ. Excurs. 116 (1830)
37
!
Chapter two: Material and Methods
!
!
Eastern, Northern and
Western Cape, Free State,
Gauteng, Kwazulu-Natal,
Limpopo, Namibia,
Swaziland
Van Wyk4115
(JRAU)
JQ039294
—
AJ512323.12
AJ511414.12
Australia, except Northern
Territory
K.Dixon s.n.
(KPBG)
—
—
—
—
Israel, Lebanon, North
Caucasus, South European
Russia, Transcaucasia,
Armenia, Georgia, Asiatic
Turkey, Ukraine
Eritrea
Chase490 (K)
JQ039296
—
—
—
Chase3941 (K)
JQ039327
—
—
—
Kwazulu-Natal,
Mpumalanga, Swaziland
Northern and Western Cape
IPMB040340
(HEID)
IPMB040342
(HEID)
—
—
AJ512329.12
AJ511423.12
—
—
AJ512330.12
AJ511425.12
Acoroides resinifera Sol. ex Kite [nom. illegit.],
Xanthorrhoea hastilis R.Br., Xanthorrhoea hastilis Sm.
[nom. illegit.], Xanthorrhoea resinifera (Sol. ex Kite)
E.C.Nelson and D.J.Bedford [nom. illegit.]
Australia
Chase192 (NCU)
—
—
HM640546.13
HM6406633
—
—
Western Australia
Western Australia
Brummitt21391
Brummitt, George
and Oliver21376
JQ039328
JQ039329
—
—
—
—
—
—
Bulbine frutescens (L.) Willd.
Enum. Pl. 372 (1809)
Anthericum frutescens L., Anthericum fruticosum Salisb.
[nom. illegit.], Anthericum incurvum Thunb.,
Anthericum multiceps Poelln., Anthericum rostratum
Jacq., Bulbine caulescens L., Bulbine frutescens var.
incurva (Thunb.) Rowley, Bulbine frutescens var.
rostrata (Jacq.) Rowley, Bulbine incurva (Thunb.)
Spreng., Bulbine rostrata (Jacq.) Willd., Phalangium
frutescens (L.) Kuntze, Phalangium rostratum (Jacq.)
Kuntze
Amaryllis semibarbata (R.Br) Steud., Anthericum
semibarbatum R.Br., Bulbine floribunda Schrad. ex
Benth., Bulbine semibarbata f. gracilescens Domin,
Bulbinopsis semibarbata (R.Br.) Borzì, Phalangium
semibarbatum (R.Br.) Kuntze, Triglochin racemosa
Endl.
Ashodelus regius Heynh., Eremurus bachtiaricus Boiss.,
Eremurus caucasicus Steven, Eremurus libanoticus
Boiss., Eremurus sibiricus Weinm., Eremurus tauricus
Weinm.
Bulbine semibarbata (R.Br.)
Haw.
Saxifrag. Enum. 2: 33 (1821)
Eremurus spectabilis M.Bieb.
Cent. Pl. Ross. Merid. 2 (1810)
Jodrellia fistulosa (Chiov.)
Baijnath
Kew Bull. 32: 576 (1978)
Bulbine breviracemosa Poelln., Bulbine fistulosa Chiov.
Kniphofia galpinii Baker
Fl. Cap. 6: 281 (1896)
—
Kniphofia uvaria (L.) Oken
Allg. Naturgesch. 3(1): 566
(1841)
Aletris uvaria (L.) L., Aloe longifolia Lam., Aloe rigida
Salisb. [nom. illegit.], Aloe uvaria L., Kniphofia aloöides
Moench, Kniphofia bachmannii Baker, Kniphofia
burchellii (Sweet ex Lindl.) Kunth, Kniphofia
occidentalis A.Berger, Kniphofia odorata Heynh. [nom.
illegit.], Kniphofia uvaria (L.) Hook., Kniphofia uvaria
var. glaucescens G.Nicholson, Kniphofia uvaria var.
nobilis (Guillon) Baker, Kniphofia uvaria var. serotina
Baker, Triclissa uvaria (L.) Salisb. [nom. illegit.],
Tritoma burchellii Sweet ex Lindl., Tritoma canari
Carrière, Tritoma glauca H.Vilm., Tritoma nobilis
Guillon,Tritoma recurva H.Vilm., Tritoma saundersii
Carrière, Tritoma uvaria (L.) Ker Gawl., Tritomanthe
uvaria (L.) Link, Tritomium uvaria (L.) Link, Veltheimia
speciosa Roth, Veltheimia uvaria (L.) Willd.
Xanthorrhoea resinosa Pers.
Syn. Pl. 1: 370 (1805)
Xanthorrhoea sp. (stemless)
Xanthorrhoea sp. (with.trunk)
—
—
38
!
Chapter two: Material and Methods
!
!
Table 2.2 Gene regions and primers used for amplifying the cpDNA and nrDNA in the current study.
Region
Size (bp)
Primer sets (Sequence 5’-3’)
Reference
cpDNA regions
trnH-psbA
812
trnH Rev: CGC GCA TGG TGG ATT CAC AAT CC
Sang et al., 1997
psbA Fwd: GTT ATG CAT GAA CGT AAT GCT C
matK
rbcLa
945
552
3F KIM: CGT ACA GTA CTT TTG TGT TTA CGA G
CBOL Plant Working
1R KIM: ACC CAG TCC ATC TGG AAA TCT TGG TTC
Group, 2009
rbcL-barcode-F: ATG TCA CCA CAA ACA GAG ACT AAA GC
CBOL Plant Working
rbcL-barcode-R: GTA AAA TCA AGT CCA CCY CG
Group, 2009
Nuclear region
ITS1
482
ITS 18-F: GTC CAC TGA ACC TTA TCA TTT AGA GG
Treutlein et al.,
ITS S5: TTC GGG CGC AAC TTG CGT TC
2003a, b
39
!
Chapter two: Material and Methods
!
!
Table 2.3 Characters, character states, and explanatory notes on characters used in the cladistic
analyses of the morphological data matrix.
!
1. Habit: stemless = 0; caulescent = 1; arborescent (tree-like, with leafless, dichotomously branched
stems) = 2
2. Leaf insertion: spiral or polystichous = 0; distichous = 1
3. Leaf margins: smooth or entire = 0; toothed or bristled = 1
4. Leaf maculation (spots): immaculate = 0; maculate = 1
5. Leaf tubercles: concolorous or absent = 0; white = 1
6. Leaf apex: acute = 0; truncate = 1
7. Inflorescence: unbranched = 0; branched = 1
8. Inflorescence: raceme erect/decumbent and symmetrical = 0; raceme oblique and secund = 1
9. Inflorescence: flowers less than pedicel length apart or equal to = 0; flowers more than pedicel
length apart = 1
10. Inflorescence: raceme unicoloured = 0; raceme bicoloured = 1
11. Flower orientation at anthesis: spreading = 0; pendulous = 1
12. Flower arrangement: raceme elongate = 0; raceme capitate = 1
13. Flower shape: rotate = 0; campanulate = 1; tubular = 2
14. Perianth symmetry: actinomorphic = 0; weakly bilabiate = 1; strongly bilabiate = 2
15. Pedicel length: ¼ or more longer than perianth = 0; less than ¼ length of the perianth = 1 (perhaps
consider using 1/3)
16. Perianth colour: white/cream/greenish = 0; yellow/orange/red = 1
17. Perianth tube: straight = 0; curved upwards = 1
18. Perianth: cylindrical and tapering into pedicel = 0; inflated basally or truncated = 1
19. Tepals: both whorls connate basally = 0; both whorls connate ½ or more of length = 1; only outer
whorls connate for half or more of length = 2
20. Stamens: as long as/longer than perianth = 0; shorter than perianth = 1
40
!
Chapter two: Material and Methods
!
!
Table 2.4 Character states for the 20 morphological characters scored for accessions included in the molecular analyses
Taxon
Morphological characters
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
Aloe albida
0
0
1
0
0
0
0
0
1
0
1
1
2
2
1
0
1
0
0
0
Aloe alooides
2
0
1
0
0
0
0
0
1
0
0
0
1
0
1
1
0
0
0
0
Aloe angelica
2
0
1
0
0
0
1
0
0/1
1
0/1
1
2
1
0/1
1
1
0
0
0
Aloe anivoranoensis
1
0
1
0
0
0
0
0
0
0
1
0
2
0
0
1
0
0
0
0
Aloe arborescens
2
0
1
0
0
0
0
0
1
0
1
0
2
0
1
1
0
0
0
0
Aloe arenicola
1
0
1
1
0
0
1
0
0
0
1
1
2
0
0
1
0
0
2
0
Aloe aristata
0
0
1
1
1
0
1
1
1
0
1
0
2
0
1
1
0
0
2
0
Aloe barberae
2
0
1
0
0
0
1
0
1
0
0
0
2
1
1
1
0
0
0
0
Aloe brevifolia
0
0
1
0/1
0
0
0
0
1
0
1
0
2
1
1
1
0/1
0/1
0
0
Aloe buhrii
0
0
0/1
1
0
0
1
0
1
0
1
0/1
2
0
0/1
1
0
1
0
0
Aloe chabaudii
0
0
1
0/1
0
0
1
0
1
0
1
0/1
2
0
0
1
1
1
2
0
Aloe challisii
0
0
1
1
0
0
0
0
1
0
1
1
2
0
1
1
0
0
0
1
Aloe chortolirioides
1
0
1
1
0
0
0
0
1
0
1
1
2
0
1
1
0
0
0
0
Aloe ciliaris
1
0
1
0
0
0
0
0
1
0
1
0
2
0
1
1
0
0
0
0
Aloe commixta
1
0
1
0
0
0
0
0
1
0/1
1
1
2
0
1
1
0
0
0
0
Aloe comosa
2
0
1
0
0
0
0/1
0
1
1
1
0
2
0
1
0/1
0
0
0
0
Aloe dewinteri
0
0
1
0
0
0
1
0
1
1
1
0
2
0
1
1
0
0
0
0
Aloe dichotoma
2
0
1
0
0
0
1
0
1
0
0
0
2
1
1
1
0
0
0
0
Aloe dichotoma subsp. pillansii
2
0
1
0
0
0
1
1
1
0
1
0
2
1
1
1
0
0
0
0
Aloe dichotoma subsp. ramosissima
1
0
1
0
0
0
1
0
1
0
1
0
2
1
1
1
0
0
0
0
Aloe ecklonis
0
0
1
1
0
0
0
0
0
0
1
1
2
1
0
1
0/1
0
0
0
Aloe eminens
2
0
1
0
0
0
1
0
1
0
1
0/1
2
1
1
0
0
0
0
0
Aloe excelsa
2
0
1
0
0
0
1
0/1
1
0
1
0
2
1
1
1
0
0
0
0
Aloe ferox
2
0
1
0
0
0
1
0
1
0
1
0
2
1
1
1
0/1
0
0
0
Aloe fouriei
1
0
1
1
0
0
0
0
0
0
1
1
2
0
0
1
0
0
0
1
Aloe glauca
1
0
1
0
0
0
0
0
1
0
1
0
2
0
1
1
0/1
0/1
0
0
41
!
Chapter two: Material and Methods
!
!
Aloe gracilis
1
0
1
0
0
0
0/1
0
1
0
1
0
2
0
1
1
0
0
2
0
Aloe greatheadii subsp. davyana
0
0
1
1
0
0
1
0
1
0
1
0
2
0
1
1
1
1
2
0
Aloe haworthioides
0
0
1
0
1
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
Aloe hereroensis
0
0
1
1
0
0
1
0
0
0
1
1
2
1
0
1
1
0
2
0
Aloe kouebokkeveldensis
0
0
1
1
0
0
1
0
1
0
1
1
2
0
1
1
0
1
1
0
Aloe lineata
1
0
1
0
0
0
0
0
1
0
1
0
2
0
1
1
0
1
0
0
Aloe lineata var. muirii
1
0
1
0
0
0
0
0
1
0
1
0
2
0
1
1
0
1
0
0
Aloe lutescens
0
0
1
0
0
0
1
0
1
1
1
0
2
0
1
1
0
0
0
0
Aloe melanacantha
0
0
1
0
0
0
0
0
1
1
1
0
2
1
1
1
0
0
0
0
Aloe microstigma
0/1
0
1
1
0
0
0
0
1
1
1
0
2
1
0/1
1
0
0
0
0
Aloe munchii
2
0
1
0
0
0
1
0
1
0
1
0/1
2
1
1
1
1
0
0
0
Aloe nubigena
1
0/1
1
1
0
0
0
0
1
0
1
1
2
0
0/1
1
0
0
0
0
Aloe pearsonii
1
0
1
0
0
0
1
0
0
0
1
1
2
1
1
1
0
0
2
0
Aloe perfoliata
1
0
1
0/1
0
0
1
0
0
0
1
1
2
0
0
1
0
0
0
0
Aloe petricola
0
0
1
0
0
0
0/1
0/1
1
1
1
0
2
1
1
0/1
1
0
0
0
Aloe pictifolia
1
0
1
1
0
0
0
0
1
0
1
0
2
0
0
1
0
0
0
0
Aloe plicatilis
2
1
0
0
0
1
0
0
1
0
1
0
2
0
1
1
0
0
0
0
Aloe plicatilis
2
1
0
0
0
1
0
0
1
0
1
0
2
0
1
1
0
0
0
0
Aloe propagulifera
0
0
1
1
0
0
0
0
1
0
1
0
2
0
1
0
0
0
2
0
Aloe reynoldsii
0/1
0
1
1
0
0
1
0
1
0
0/1
1
2
0
1
1
0/1
1
2
0
Aloe rupestris
2
0
1
0
0
0
1
0
1
1
1
0
2
0
1
1
0
0
0
0
Aloe saundersiae
0
0
1
0/1
0
0
0
0
1
0
0/1
1
2
1
1
1
0
0
0
1
Aloe spicata
2
0
1
0
0
0
0
0
1
1
0
0
1
0
1
1
0
0
0
0
Aloe striata
0/1
0
0
0
0
0
1
0
1
0/1
1
1
2
0
1
1
0/1
1
2
0
Aloe striata subsp. karasbergensis
0
0
0
0
0
0
1
0/1
1
0/1
1
0/1
2
1
1
1
0/1
1
2
0
Aloe striata subsp. kommagasensis
0
0
0
0
0
0
1
0
1
0/1
1
1
2
1
1
1
0/1
1
2
0
Aloe striatula
1
0
1
0
0
0
0
0
1
1
1
0
2
0
1
1
0
0
0
0
Aloe succotrina
0/1
0
1
0
0
0
0
0
1
0
1
0
2
0
1
1
0
0
0
0
42
!
Chapter two: Material and Methods
!
!
Aloe tenuior
1
0
1
0
0
0
?
0
1
1
0/1
0
2
0
1
1
0
0
2
0
Aloe thraskii
2
0
1
0
0
0
1
0
1
0/1
0
0
2
0/1
1
1
1
0
0
0
Aloe verecunda
1
0/1
1
1
1
0
0
0
1
0
1
1
2
0
0/1
1
0
0
0
0
Aloe vossii
0/1
0
1
1
0
0
0
0
0
0
1
1
2
0
0
1
0
0
0
0
Aloe vryheidensis
1/2
0
1
0
0
0
0
0
1
1
0
0
1
0
1
1
0
1
0
0
Anthericum liliago
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Asphodeline lutea
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Astroloba corrugata
1
0
0
0
0
0
0
0
0
0
0
0
2
0
1
0
0
0
0
1
Astroloba foliolosa
1
0
0
0
0
0
0
0
1
0
0
0
2
0
1
0
0
0
0
1
Astroloba herrei
1
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
1
Astroloba rubriflora
1
0
1
0
0
0
0
1
1
0
0/1
0
2
0
1
1
1
0
1
1
Bulbine frutescens
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Bulbine semibarbata
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Chortolirion angolense
0
0
1
1
1
0
0
0
1
0
0
0
2
1
1
0
0
0
1
1
Eremurus spectabilis
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Gasteria acinacifolia
0
0
0/1
1
0/1
0
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria batesiana var. dolomitica
0
0
1
1
1
0
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria carinata var. carinata
0
0
0
0
1
0
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria carinata var. retusa
0
1
0
0
1
0/1
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria croucheri
0
0
0
1
0
0
0
1
1
0
1
0
2
1
1
2
1
1
1
0
Gasteria disticha
0
1
0
1
1
1
0/1
0
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria doreeniae
0
1
0
1
1
0
0
0
1
0
1
0
2
1
1
2
1
1
0/1
1
Gasteria ellaphieae
0
0
1
0
1
0
1
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria excelsa
0
0
0
1
0
0/1
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria glauca
0
0
1
0
1
0
0
1
1
0
1
0
2
1
1
2
1
?
?
?
Gasteria glomerata
0
1
0
0
1
1
0/1
0/1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria nitida var. armstrongii
0
1
0
0
1
1
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria obliqua
1
0/1
0
1
0/1
1
1
0/1
1
0
1
0
2
1
1
2
1
1
1
1
43
!
Chapter two: Material and Methods
!
!
Gasteria pillansii var. ernesti-ruschii
0
1
0
1
1
0/1
0
1
1
0
1
0
2
1
1
2
1
1
0
0
Gasteria pillansii var. pillansii
0
1
0
1
1
0/1
0
1
1
0
1
0
2
1
1
2
1
1
0
0
Gasteria polita
0
0
1
1
0
1
0
1
1
0
1
0
2
1
1
2
1
1
1
0
Gasteria pulchra
0
0
1
1
0
0
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria rawlinsonii
1
1
0/1
0/1
0
1
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Gasteria tukhelensis
0
0
0/1
1
0
0
1
1
1
0
1
0
2
1
1
2
1
1
1
0/1
Gasteria vlokii
0
0
1
1
0
1
0
1
1
0
1
0
2
1
1
2
1
1
1
1
Haworthia angustifolia
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
?
Haworthia arachnoidea
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia arachnoidea
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia attenuata
0
0
0
0
1
0
0/1
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia bayeri
0
0
0/1
0
0
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia blackburniae
0
0
0/1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia blackburniae var. derustensis
0
0
0/1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia blackburniae var. graminifolia
0
0
0/1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia bruynsii
0
0
0
0
0/1
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia chloracantha
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia coarctata var. adelaidensis
1
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia coarctata
1
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia cooperi
0
0
0/1
0
0
0/1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia cooperi var. doldii
0
0
0/1
0
0
0/1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia cymbiformis
0/1
0
0
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia cymbiformis var. ramosa
0/1
0
0
0
0
?
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia cymbiformis var. setulifera
0/1
0
0
0
0
?
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia decipiens
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia decipiens var. pringlei
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia decipiens var. virella
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia emelyae
0
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
44
!
Chapter two: Material and Methods
!
!
Haworthia emelyae var. comptoniana
0
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia fasciata
0/1
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia floribunda
0
0
0/1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia glauca
1
0
0/1
0
0/1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia herbacea var. flaccida
0
0
1
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia herbacea var. paynei
0
0
1
0
1
0
0
0
1
0/1
0
0
2
2
1
0
0
0
0
1
Haworthia kingiana
0
0
0
0
0/1
0
0/1
0
1
0
0
0
2
1
1
0
0
0
0
1
Haworthia kingiana
0
0
0
0
0/1
0
0/1
0
1
0
0
0
2
1
1
0
0
0
0
1
Haworthia koelmaniorum var. mcmurtryi
0
0
1
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia limifolia
0
0
0
0
0/1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia limifolia var. ubomboensis
0
0
0
0
0/1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia lockwoodii
0
0
0
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia longiana
0
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia maculata
0
0
1
1
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia magnifica var. dekenahii
0
0
0
0
0
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia marginata
0
0
0
0
0
0
1
0
1
0
0
0
2
1
1
0
0
0
0
1
Haworthia marginata
0
0
0
0
0
0
1
0
1
0
0
0
2
1
1
0
0
0
0
1
Haworthia marumiana var. archeri
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia marxii
0
0
0
1
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mirabilis
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mirabilis var. calcarea
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mirabilis var. paradoxa
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mirabilis var. triebneriana
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia monticola var. asema
0
0
1
1
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mucronata var. bijliana
0
0
0
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mucronata var. habdomadis
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mucronata var. morrisiae
0
0
0
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mucronata var. rycroftiana
0
0
0
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
45
!
Chapter two: Material and Methods
!
!
Haworthia mutica
0
0
0
0
0
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia mutica var. nigra
0
0
0
0
0
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia nigra
1
0
0
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia outeniquensis
0
0
1
0/1
0
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia pulchella var. globifera
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia pumila
0
0
0
0
1
0
1
0
1
0
0
0
2
1
1
0
0
0
0
1
Haworthia pygmaea
0
0
0
0
0
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia reinwardtii
1
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia reinwardtii var. brevicula
1
0
0
0
1
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia reticulata
0/1
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia retusa
0
0
0/1
0
0
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia semiviva
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia sordida
0
0
0
0
0
0/1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia truncata
0
1
0
0
1
1
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia variegata
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia venosa
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia venosa var. granulata
0/1
0
0
0
0
0
0
1
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia vlokii
0
0
1
1
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia wittebergensis
0
0
1
0
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Haworthia zantneriana
0
0
0
0/1
0
0
0
0
1
0
0
0
2
2
1
0
0
0
0
1
Jodrellia fistulosa
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Kniphofia galpinii
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Kniphofia uvaria
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Tecophilaea cyanocrocus
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Xanthorrhoea resinosa
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Xanthorrhoea sp.
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
Zephyra elegans
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
!
46
!
Chapter two: Material and Methods
!
!
!
Figure 2.1 Organisation of the ITS region. Green area represents ITS region sequenced in the
study. Arrows indicate orientation and approximate position of primer sites. Adapted from
Baldwin et al. (1995).
!
Figure 2.2 Distribution of Alooideae collected in the wild and grown at Sheilam Nursery, Gariep
Nursery and Kirstenbosch Botanical Garden, used in this study (yellow circles).
47
!
Chapter three: Results
!
!
CHAPTER(THREE(
Results(
H. cymbiformis var. ramosa
virella
H. decipiens var.
lei
s var. pring ii
H. decipien
i var. dold is
H. cooper
biform
H. cym
ra
setulife
is var. . marxii
biform
H
a
H. cym
miviv
H. se rcheri
a
var. . vlokiis
iana
H ensi
arum
erg ae
b
H. m
itte mely a
H. w H. e ibund a
tus
flor
H. H. re iana
ner tica
ieb mu ra
. tr H. . nig rea
va r
ar a xa
ilis
a v alc o ii
tic r. c rad ah a
rab
mi
mu va pa en id
H.
H. ilis var. dek lacc
.
b
ra is ar r. f
mi bil a v va
H. mira ific ea
c
n
H. ag rba
m e
H. H. h
●
H.
he
●
●
●
●
●
●
●
●
●
●
●
●
as
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
lir
io
A n
Al loe ang
ub
oe lu ol
sp
re te en
.k
om Alo yno scen se
Alo
ma e s lds s
ek
t
ou
eb Al gas riat ii
Alo okke oe b ensis a
e s vel uhr
d
a
Alo un ens ii
e v der is
si
er
A ecu ae
Alo loe vo nda
e ch ssii
Aloe allis
Aloe fourie ii
H. ko
elman
Aloe ecklon i
io
is
nu
ru
H. koe
lmanio m var. mc bigena
m
rum v
ar. mc urtryi
murt
Aloe ar ryi
ista
Aloe aris ta
tata
Aloe aristata
Astroloba rubriflo
ra
Astroloba foliolosa
Astroloba herrei
to
iat
str
Al
oe
●
●
●
●
●
●
● ●
●
or
Ch
●
lia
ifo
in sis
m
a
en
gr rust
r.
va r. de
a
ae
ni e v
i
ur ia
●
nsi
kb urn iae
c
illa
la kb rn
a
.p
p
. b lac kbu tilis
s
sim
H . b ac ca lis
ub
sis
mo
H . bl pli ati ma s
a
r
.
ic o
H e
● Aloloe pldichototoma a subsp
A oe ich om
t
l
d
o
A
h
●
e c
e
Alo oe di rbera scens
● AlAloe biane frute ibarbata
m
b
Bul ine se istulosa
●
f
ulb
● BJodrelhlioafia uvalpriainii
●
Knip hofia ga tabilis
c
● KEnreipmurus snpeelutea
●
hodeli a sp.
sp
A
● Xanthorrhoe
ea sp.
● Xanthoorrrrhhooea resinosa
●
Xanth
Zephyra elegans
● Tecophilaea cyanocrocus
●
Anthericum liliago
G. polita
●
G. doreeniae
G. disticha
tukhelensi
● GG.. pu
s
lchr
● GG. glomeraata
. glau
ca
● ● GG. .orawlinson
b
G. v liqua ii
G. lokii
●
G. excels
●
G. ellaph a
G bate ieae
● ● GG. a. cnitidasiana va
in va
r
.
c
G a aci r. ar . dolo
ms
mi
G . ca rina foli
●
● ● ● ● G G. p. crourinatata var a trongii tica
.
i
c
H . pi llan her var. retu
. c lla si i
ca sa
oa ns i v
rin
●
rc ii ar.
ata
ta va e
ta r. rn
pi est
lla i−
ns ru
sc
ii
hii
a
ul sis
ic
ev en
br laid
r.
va ade
.
ii
dt ar
ar a v
nw at ii
ei rct dt
. r a ar
H . co inw
H re ra
H. nig ca
i
u
H. la nsi
g
u
H. . bry ida
nsis
boe
H sord iana
bom
H. long ta
r. u
a
H. fasci lia va
H. limifo ia
H. limifol a
ulata
H. imifoli ar. gran
H. l enosa v
H. v nosa
H. ve enuata
H. att giana
H. kin ginata
H. mar
H. pumila
H. kingiana
H. marginata
Astroloba corrugata
ac H . m
ea .
a
Al va retic cula
oe r. u t
Al com pay lata a
●
Alo oe g mi nei
r x
e
●
Alo stri acili ta
●
Alo e cil atula s
ia
e
●
Alo
e c Aloe tenui ris
hor
o
tol albid r
●● ●
Aloe Aloe irioid a
mel emin es
●
a
e
n
a
Aloe
can ns
t
h
p
Aloe earsoniia
●
Aloe arenicola
●
●
perfo
li
ata
Aloe su
●
●
ccotrin
a
Aloe gl
au
Aloe brev ca
ifo
●
Aloe haworth lia
ioides
Aloe anivoranoens
is
Aloe dewinteri
Aloe propagulifera ●
●
Aloe hereroensis
●
a
yan
dav
.
var
ii
●
Aloe greathead
sbergensis
subsp. kara
●
eidensis
Aloe striata
Aloe vryh audii
●
ab
Aloe ch unchii
●
m
Aloe tricola ●
●
pe
Aloe excelsa
Aloe ngelica
●
a
s
e
e
Alo alooid a
s
Aloe e comotris ●
Alo rupes skii
●
e
a a ●
o
r
l
A
e th eat
Alo e lin uirii i
●
●
Alo ar. m uiri a
a v r. m gm
●
eat va sti lia
lin ata icro ctifo ens
e
e
x
i
o
Al e lin loe m e p resc fero ta
A Alo rbo oe ica
Alo
a Al sp
oe
oe
Al
Al
●
●
H
rb
H. cymbiformis var. setulifera
H. cooperi
H. mucronata var
. habdomadis
H. lockwoo
H. pulche dii
H. muc lla var. globifera
ronata
var. rycr
H. mu
oftiana
H. mu cronata va
r. morr
c
ro
nata
H. ar
is
v
a
a
r. bijli iae
H. a chnoid
a
na
H. d rachno ea
id
e
c
e
ipie
a
H.
H. chlor ns
H. pygm acanth
a
H. varieg aea
H. trun ata
H em cata
H. . mo elyae
va
H an nti
H . za gus cola r. c
H . b ntn tifo va omp
H . ou ayer eria lia r. as tonia
. m te i
em
na
na
a
ira niq
bi uen
lis
sis
!
●
●
● ●
●
●
●
●
●
●
48
!
Chapter three: Results
!
!
3. RESULTS
3.1 Molecular Evolution
The statistics from MP analyses for the single plastid analysis, combined plastid (rbcLa +
matK + trnH-psbA), ITS1 and the combined plastid and nuclear datasets are shown in Table
3.1. Of the plastid regions used, matK had a significantly higher number of variable sites
(33%; Table 3.1) than trnH-psbA (16%) or rbcLa (14%). The number of potentially
informative characters for matK (21%) is much higher than for the two other plastid regions,
rbcLa and trnH-psbA (10% and 9% respectively). The ITS1 region had significantly more
variable sites (40%) compared to the plastid regions. It also evolves at a much faster rate than
the plastid genes (2.1 changes per variable site) with a lower consistency index (CI).
Results from the following analyses are presented: simplified tree topologies of the
combined plastid regions (rbcLa + matK + trnH-psbA), ITS1 and combined plastid and
nuclear regions (Fig. 3.1), combined plastid regions (rbcLa + matK + trnH-psbA; Fig. 3.2),
ITS1 (Fig. 3.3) and combined plastid and ITS1 (rbcLa + matK + trnH-psbA + ITS1; Fig. 3.4).
The MP analyses were largely congruent with the Bayesian analyses, and therefore results are
displayed on the same tree (Fig. 3.4).
3.2 Combined Plastid Data
Individual plastid sequence analyses (results not shown) were topologically consistent
(negligible to zero incongruence), and for the purpose of the results and discussion were
combined and treated as a single analysis. The MP analysis yielded 661 most parsimonious
trees of 752 steps with a CI of 0.72 and a RI of 0.91 (Table 3.1). One of the most
parsimonious trees is illustrated in Fig. 3.2. Alooideae is strongly supported as monophyletic
(BP = 94; Fig. 3.2). Although resolution in the plastid tree were relatively poor, possibly due
to a lack of sufficient informative sequence variation, we observed eight clades (A, B, C, D, E,
49
!
Chapter three: Results
!
!
F, G and H) while the internal groups for each clade were retrieved as a polytomy.
3.3 ITS1 data
Analyses resulted in 314 equally most parsimonious trees with TL = 346, CI = 0.64, and RI =
0.92. One of the most parsimonious trees is illustrated in Fig. 3.3. Seven major clades were
recovered; Haworthia subg. Haworthia, H. subg. Robustipendunculares, Gasteria, Astroloba,
Aloe sect. Macrifoliae, tree Aloes, and remaining Aloe species. The tree Aloe species (A.
barberae, A. eminens and A. dichotoma) were retrieved as the early diverging lineage to the
subfamily Alooideae.
3.4 Combined plastid and nuclear dataset (rbcLa+matK+trnH-psbA+ ITS1)
Results of the combined plastid analysis are topologically congruent with those of the ITS
analysis, and all data were therefore combined. Congruence between the ITS and plastid
datasets was addressed by comparisons of bootstrap percentages from separate analyses.
Bootstrap trees were considered incongruent only when hard (i.e. with high bootstrap support)
instead of “soft” (with low bootstrap support) incongruence was displayed (See Wiens, 1998).
The aligned combined dataset (rbcLa + matK + trnH-psbA + ITS1) included 2518 characters
of which 604 (24%) are variable. The MP analysis yielded 5020 equally most parsimonious
trees of 1159 steps with a CI of 0.63 and a RI of 0.89 (Table 3.1). A monophyletic Alooideae
was recovered with high support (Fig. 3.4; BP = 93; PP = 1.00). It resolves eight major clades
represented in different colours in Fig. 3.4: Aloe plicatilis and the remaining tree Aloes,
Haworthia subg. Haworthia, the climbing Aloes, the true Aloe species, ‘Haworthioid’ clade
(H. subg. Robustipendunculares + Astroloba + Aloe aristata), Haworthia subg. Hexangulares,
and Gasteria. While the combined data recovered eight clades, the plastid data retrieved five
with clades A, B, D, and E, grouping together. The ITS1 data recovered seven clades
50
!
Chapter three: Results
!
!
compared to eight in the combined data.
The tree Aloe species — The analysis indicates an isolated position for Aloe plicatilis
in a polytomy with Haworthia subg. Haworthia but without BP support and weak support in
the BI analysis (PP = 0.58); and for A. eminens in the clade including A. perfoliata but without
BP support and weak support in the BI analysis (PP = 0.58). The other tree Aloe species (A.
barberae, A. dichotoma, and A. pillansii) were retrieved as a monophyletic clade with
moderate support in MP analysis and strong support in the BI analysis (BP = 76; PP = 1.00).
The A. plicatilis, A. eminens and A. barberae lineages constitute the early diverging elements
in subfamily Alooideae.
Haworthia subg. Haworthia — This clade is well supported in the MP and BI
analyses (BP = 98; PP = 1.00). This clade resolved (BP = 96; PP = 1.00) the grass-like
Haworthia blackburniae as the earliest-diverging lineage to the other Haworthia species,
which were unresolved.
Climbing Aloes — This group, representing Aloe sect. Macrifoliae (Aloe ciliaris, A.
commixta, A. gracilis, A. striatula, and A. tenuior) was recovered in both analyses as a
strongly supported clade (BP = 95; PP = 1.00).
The true Aloes — This group, which comprises the majority of Aloe species plus
Chortolirion (coloured brown; Fig. 3.4) was retrieved as a group with strong support (PP =
1.00) in the BI. Several internal clades, none of which correspond exactly to current sections
recognised in the taxonomy of the genus, were well supported in the BI analysis.
Among these, the majority of the grass Aloes (sect. Leptoaloe) grouped together (BP =
89; PP = 1.00) in a clade comprising Chortolirion angolense, A. kouebokkeveldensis, A.
buhrii, A. lutescens, A. reynoldsii, A. spicata, A. striata subsp. komaggasensis, and A. striata
subsp. striata. In the MP tree, it was recovered as part of a larger polytomy with no bootstrap
51
!
Chapter three: Results
!
!
support.
A second clade comprised several of the ‘single-stemmed Aloes’ (Aloe sect.
Pachydendron) plus other species, with weak MP support (BP = 61) but strong support in the
BI analysis (PP = 1.00).
A third clade comprising a heterogenous assemblage of Aloe species (A. excelsa, A.
petricola, A. munchii, A. chabaudii and A. vryheidensis) was well supported in the BI (BP =
53; PP = 0.99).
The fourth clade also recovered a heterogenous mix of Aloe species containing the type
of the genus, A. perfoliata with weak to moderate support (BP = 53; PP = 0.92).
‘Haworthioid’ clade (Haworthia subg. Robustipendunculares, Astroloba, Aloe
aristata, and H. koelmaniorum) — This clade was strongly supported (BP = 92, PP = 1.00) as
sister to the Gasteria–Hexangulares clade but was only moderately supported as monophyletic
in the BI (BP = 58; PP = 0.68). Haworthia koelmaniorum var. mcmurtryi (BP = 99; PP =
1.00), also Hexangulares was retrieved with weak BI support (PP = 0.68) as the earliestdiverging lineage in the clade. There is moderate to strong support (BP = 77, PP = 0.95) for a
sister relationship between Aloe aristata and H. subg. Robustipendunculares + Astroloba. The
remaining species of H. subg. Robustipendunculares (highlighted in chartreuse) were
recovered with strong support (BP = 100; PP = 1.00) but are only weakly supported in BI (PP
= 0.74) as sister to a moderately to well supported (BP = 79; PP = 1.00) Astroloba.
Haworthia subg. Hexangulares — H. subg. Hexangulares (Fig. 3.4) was recovered in
both MP and BI with moderate to strong support (BP = 80; PP = 1.00) in a clade with
Gasteria. Relationships between Haworthia attenuata, the rest of subg. Hexangulares and
Gasteria remain unresolved.
52
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Chapter three: Results
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!
Gasteria — Gasteria species form a well-supported monophyletic clade in both
analyses (BP = 97; PP = 1.00) (Fig. 3.4). The clade is recovered in all trees as one element of a
moderately to well supported trichotomy (BP = 80; PP = 1.00) that includes the species of
Haworthia subg. Hexangulares.
3.5 Evolution of morphological traits
Among 20 morphological traits that were analysed, habit, leaf arrangement, leaf maculation
and tuberculation, inflorescence branching and orientation, inflorescence shape and colour,
perianth orientation at anthesis, perianth symmetry and colour, perianth shape and curvature,
tepal connation, and stamen length are all shown to be homoplasious (Tables 2.3 and 2.4; Fig.
3.5).
Table 3.2 presents summary of the characters and character states constructed for nine of
the 20 morphological characters for Alooideae. These traits are selected because of their
importance in generic and infrageneric delimitation within Alooideae (see also Figs. 3.6Aabc,
3.7abc and 3.8abc ).
In Gasteria, only G. obliqua and G. rawlinsonii exhibit caulescence (character 1; Fig.
3.6a). Among members of Haworthia subg. Hexangulares almost equal numbers exhibit
caulescence and acaulescence. Members of subg. Robustipendunculares are all stemless but
all Astroloba species (including Poellnitzia) are caulescent. An arborescent habit is
characteristic of only a few species of Aloe, namely A. plicatilis (sect. Kumara) and members
of sect. Pachydendron and Dracoaloe.
Most members of Alooideae have a polystichous leaf arrangement (character 2; Fig.
3.6b). Exceptions include a few Gasteria species, which show a distichous leaf arrangement in
the juvenile stage. Most become polystichous later in the adult stage with the exception of
53
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Chapter three: Results
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Gasteria obliqua, which retains the distichous leaf arrangement into the adult stage. Other
species with a distichous leaf arrangement are Aloe nubigena, A. plicatilis, A. verecunda and
Haworthia truncata.
A toothed or bristled leaf margin (character 3) is characteristic of all species except
members of Haworthia subg. Hexangulares, in which the leaf margins are entire.
Maculate leaves (character 4) are consistently present in all Gasteria species and also in
several aloes, including Aloe aristata and the grass Aloes (sect. Leptoaloe).
Leaf tubercles (character 5; Fig. 3.6c) are also characteristic of all Gasteria species plus
Aloe aristata, and also all species of Haworthia subg. Hexangulares and Robustipedunculares.
Most members of Alooideae have acute leaf apices (character 6) except some Gasteria
and Haworthia species, which have truncate to blunt apices.
Five of the 20 Gasteria species (G. disticha, G. tukhelensis, G. glomerata, G. obliqua
and G. ellaphieae) have branched inflorescences (character 7) and this trait is also found in all
species of Astroloba and Haworthia subg. Robustipendunculares, but not in subg.
Hexangulares and Haworthia. Various species of Aloe also have branched inflorescences.
A secund inflorescence (character 8) is characteristic of all Gasteria species, and also of
Astroloba (Poellnitzia) rubriflora, Haworthia venosa var. granulata, Aloe excelsa, A.
petricola and A. striata var. karasbergensis.
Relatively lax inflorescences (character 9) are characteristic of most Alooideae. Only a
few Aloe species exhibit bicoloured racemes (character 10; e.g. A. striata, A. lutescens, A.
spicata, A. pictifolia, A. thraskii, A. rupestris, A. comosa). Pendulous flowers (character 11;
Fig. 3.7a) are found in most Aloe species and in Gasteria.
Capitate inflorescences (character 12) are found among relatively few, unrelated Aloe
species (sect. Leptoaloe plus A. chabaudii, A. munchii and A. striata subsp. karasbergensis).
54
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Chapter three: Results
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Strongly bilabiate flowers (character 14; Fig. 3.7b) are diagnostic for Haworthia subg.
Haworthia and Hexangulares plus Chortolirion angolense but H. subg. Robustipendunculares
and several Aloe species, notably A. albida, have ± zygomorphic flowers. A short pedicel
(character 15) is characteristic of all Alooideae. Flower colouring is strongly correlated with
current generic circumscriptions. All the Gasteria species have unusual, bicoloured flowers;
and all Haworthia species; Chortolirion and all but one Astroloba species have
white/cream/greenish flowers. The great majority of Aloe species have orange/red flowers but
isolated species have whitish flowers. Perianth curvature (character 17; Fig. 3.7c) is also
associated with current generic circumscriptions. All species of Chortolirion, Gasteria and
Haworthia subg. Haworthia and Hexangulares have curved perianth tubes whereas Astroloba,
H. subg. Robustipendunculares and the majority of Aloe species have straight perianth tubes.
Basally inflated flowers (character 18; Fig. 3.7a) are characteristic of Gasteria but also of
several Aloe species. All Gasteria species and some Aloe species have both tepal whorls
connate for half or more of their length (character 19; Fig. 3.7c) but a number of Aloe species
have flowers in which only the outer tepals are connate for half or more of length. Included
stamens (character 20; Fig. 3.7b) are typical of Chortolirion, Gasteria and Haworthia but are
also found in various Aloe species.
55
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Chapter three: Results
!
!
Table 3.1 Statistics from MP analyses obtained from separate and combined datasets.
rbcLa
matK
trnH-psbA
ITS1
157
Combined plastid (rbcLa
+ matK + trnH-psbA)
172
156
Combined plastid +
ITS1
172
No. of taxa included
172
172
No. of included characters
552
789
768
2109
409
2518
No. of constant characters
472
531
645
1648
244
1914
No. of variable sites
80
(14%)
258
(33%)
123
(16%)
461
(22%)
165
(40%)
604
(24%)
No. of parsimony
informative sites
55
(10%)
165
(21%)
70
(9%)
290
(14%)
92
(22%)
376
(15%)
No. of trees (Fitch)
523
408
101
661
314
5020
No. of steps (Tree length)
120
408
194
752
346
1159
CI
0.74
0.75
0.74
0.72
0.64
0.63
RI
0.91
0.92
0.92
0.91
0.92
0.89
Average number of changes
per variable site (number of
steps/ number of variable
sites)
Model selected by AIC
1.5
1.6
1.6
1.6
2.1
1.9
HKY+
G
TVM+G
TVM+I+ G
TRN+G
GTR+I+G
AIC = Akaike information criterion; CI = consistency index; RI = retention index
56
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Chapter three: Results
!
!
Table 3.2 Characters states for nine selected autapomorphic traits which have been employed
in previous Alooideae systematics.
Character
Character state
Species
Growth habit
Stemless
Most Gasteria; half of Hexangulares;
Robustipendunculares
Caulescent
Gasteria obliqua; half of Hexangulares; all
Astroloba (including Poellnitzia);
Arborescent
A. plicatilis (sect. Kumara); sect.
Pachydendron and Dracoaloe.
Most Alooideae;
Leaf insertion
Spiral/polystichous
Distichous
Leaf maculation
Immaculate
Maculate
Flower orientation at
anthesis
Spreading
Pendulous
Perianth symmetry
Actinomorphic
Weakly bilabiate
Strongly bilabiate
Perianth curvature
Straight
Curved upwards
Perianth
Stamen inclusion
Cylindrical and tapering
into pedicel
Inflated basally or
truncated
Exserted
Inserted
Tepal connation
Both whorls connate
basally
Both whorls connate ½ or
more of length
Only outer whorls connate
½ or more of length
Gasteria obliqua; Aloe nubigena, A. plicatilis,
A. verecunda and Haworthia truncata.
Most Alooideae
All Gasteria; several Aloe species e.g. A.
aristata and Aloe sect. Leptoaloe.
Chortolirion, Astroloba, Haworthia and a few
Aloe
Most Aloe and Gasteria species.
Robustipendunculares and several Aloe e.g. A.
albida
Gasteria species
Haworthia subg. Haworthia and
Hexangulares plus Chortolirion angolense
Astroloba, Robustipendunculares and majority
of Aloe
Chortolirion, Gasteria; Haworthia subg.
Haworthia and Hexangulares
Most Aloe; Astroloba; Chortolirion;
Haworthia subg. Haworthia and
Hexangulares
Gasteria and several Aloe species.
Most Aloe species
Chortolirion, Gasteria and the three
Haworthia subgenera
The three Haworthia subgenera; Astroloba
and some Aloe species
Gasteria and some Aloe species
A few Aloe species
57
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Chapter three: Results
!
!
(A) Plastids
(B) ITS1
(C) Combined
Gasteria (N = 20)
Gasteria (N = 20)
Gasteria (N = 20)
80
52
Hexangulares (N = 15)
1.00
Robustipendunculares (N = 5)
94
Aloe haworthioides
Astroloba (N = 4)
Hexangulares (N = 16)
92
1.00
Hexangulares (N = 4)
-
Haworthia attenuata
62
77
58 0.95
Aloe aristata (N = 3)
Robustipendunculares (N = 5)
0.68
Aloe aristata (N = 3)
65
Hexangulares (N = 14)
56
Haworthia koelmaniorum (N = 2)
68
76
Aloe aristata (N = 3)
Aloe sect. Leptoaloe (N = 7)
-
Aloe sect. Macrifoliae (N = 5)
Chortolirion angolense
Single-stemmed Aloe spp. (N = 12)
Haworthia subg. Haworthia (N = 49)
Aloe sect. Leptoaloe (N = 6)
Astroloba (N = 4)
Haworthia subg. Haworthia (N = 44)
1.00
92
Robustipendunculares (N = 5)
Haworthia koelmaniorum (N = 2)
0.97
Astroloba (N = 4)
0.74
61
-
True Aloe spp. (N = 46)
Other true Aloe spp. (N = 21)
0.53
96
Chortolirion angolense
Chortolirion angolense
Haworthia subg. Haworthia (N = 49)
93
1.00
Aloe plicatilis (N = 2)
Single-stemmed Aloe spp. (N = 12)
Aloe sect. Macrifoliae (N = 3)
Tree Aloe spp. (N = 5)
Outgroups (N = 13)
94
Aloe eminens
Aloe barberae
Aloe dichotoma (N = 3)
Aloe sect. Macrifoliae (N = 5)
Aloe plicatilis (N = 2)
Tree Aloe spp. (N = 5)
Outgroups (N = 13)
Figure 3.1 Simplified tree topologies showing relationships among Alooideae genera for the plastids, nuclear and combined dataset.
58
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Chapter three: Results
!
!
60
63
91
64
63
H
77
96
61
65
64
G
97
67
52
99
67
62
99
99
54
F
97
65
98
76
88
64
57
83
98
D
59
100
Gasteria tukhelensis
Gasteria pulchra
Gasteria glomerata
Gasteria glauca
Gasteria carinata var. retusa
Gasteria carinata var. carinata
Gasteria rawlinsonii
Gasteria obliqua
Gasteria vlokii
Gasteria polita
Gasteria excelsa
Gasteria ellaphieae
Gasteria doreeniae
Gasteria disticha
Gasteria croucheri
Gasteria batesiana var. dolomitica
Gasteria nitida var. armstrongii
Gasteria acinacifolia
Gasteria pillansii var. ernesti-ruschii
Gasteria pillansii var. pillansii
Haworthia nigra
Haworthia bruynsii
Haworthia coarctata var. adelaidensis
Haworthia reinwardtii
Haworthia sordida
Haworthia glauca
Haworthia coarctata
Haworthia reinwardtii var. brevicula
Haworthia longiana
Haworthia fasciata
Haworthia limifolia var. ubomboensis
Haworthia limifolia
Haworthia limifolia
Haworthia venosa var. granulata
Haworthia venosa
Aloe haworthioides
Haworthia attenuata
Haworthia pumila
Haworthia kingiana
Haworthia kingiana
Haworthia marginata
Haworthia marginata
Aloe aristata
Aloe aristata
Aloe aristata
Astroloba corrugata
Astroloba herrei
Astroloba foliolosa
Astroloba rubriflora
Haworthia koelmaniorum var. mcmurtryi
Haworthia koelmaniorum var. mcmurtryi
Haworthia mucronata var. habdomadis
Haworthia lockwoodii
Haworthia pulchella var. globifera
Haworthia monticola var. asema
Haworthia decipiens var. pringlei
Haworthia mucronata var. rycroftiana
Haworthia mucronata var. morrisiae
Haworthia mucronata var. bijliana
Haworthia arachnoidea
Haworthia arachnoidea
Haworthia maculata
Haworthia decipiens var. virella
Haworthia decipiens
Haworthia cymbiformis var. setulifera
Haworthia cymbiformis var. ramosa
Haworthia cymbiformis var. setulifera
Haworthia cymbiformis
Haworthia cooperi var. doldii
Haworthia cooperi
Haworthia herbacea var. paynei
Haworthia reticulata
Haworthia wittebergensis
Haworthia vlokii
Haworthia mirabilis var. paradoxa
Haworthia herbacea var. flaccida
Haworthia floribunda
Haworthia marxii
Haworthia marumiana var. archeri
Haworthia semiviva
Haworthia mirabilis var. calcarea
Haworthia mirabilis var. triebneriana
Haworthia retusa
Haworthia mutica var. nigra
Haworthia mutica
Haworthia magnifica var. dekenahii
Haworthia emelyae
Haworthia outeniquensis
Haworthia variegata
Haworthia truncata
Haworthia angustifolia
Haworthia zantneriana
Haworthia mirabilis
Haworthia emelyae var. comptoniana
Haworthia chloracantha
Haworthia bayeri
Haworthia pygmaea
Haworthia blackburniae var. graminifolia
Haworthia blackburniae var. derustensis
Haworthia blackburniae
Gasteria
Hexangulares I
Aloe haworthioides
Hexangulares I
Robustipendunculares
Aloe aristata
Astroloba
Hexangulares II
Haworthia subg.
Haworthia
[Fig 3.2b]
59
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Chapter three: Results
!
!
[Fig 3.2a]
85
75
50
65
96
85
96
91
79
80
61
50
81
70
92
C 62
B 95
81
64
83
100
100
90
100
98
58
100
79
100
100
Aloe kouebokkeveldensis
Aloe buhrii
Aloe striata subsp. kommagasensis
Aloe reynoldsii
Aloe striata
Aloe fouriei
Aloe nubigena
Aloe challisii
Aloe lutescens
Aloe vossii
Aloe verecunda
Aloe saundersiae
Chortolirion angolense
Aloe ecklonis
Aloe spicata
Aloe ferox
Aloe arborescens
Aloe microstigma
Aloe pictifolia
Aloe lineata var. muirii
Aloe lineata var. muirii
Aloe lineata
Aloe thraskii
Aloe rupestris
Aloe comosa
Aloe alooides
Aloe angelica
Aloe munchii
Aloe excelsa
Aloe petricola
Aloe chabaudii
Aloe vryheidensis
Aloe striata subsp. karasbergensis
Aloe hereroensis
Aloe greatheadii var. davyana
Aloe dewinteri
Aloe anivoranoensis
Aloe propagulifera
Aloe perfoliata
Aloe pearsonii
Aloe arenicola
Aloe melanacantha
Aloe tenuior
Aloe ciliaris
Aloe striatula
Aloe plicatilis
Aloe plicatilis
Aloe gracilis
Aloe commixta
Aloe brevifolia
Aloe glauca
Aloe chortolirioides
Aloe albida
Aloe succotrina
Aloe dichotoma subsp. pillansii
Aloe eminens
Aloe dichotoma
Aloe dichotoma subsp. ramosissima
Aloe barberae
Bulbine frutescens
Bulbine semibarbata
Jodrellia fistulosa
Kniphofia uvaria
Kniphofia galpinii
Eremurus spectabilis
Asphodeline lutea
Xanthorrhoea sp.
Xanthorrhoea sp.
Xanthorrhoea resinosa
Zephyra elegans
Tecophilaea cyanocrocus
Anthericum liliago
Section
Leptoaloe
Chortolirion
Single−
stemmed
Aloes
[E]
True Aloes
Section
Macrifoliae
[A]
Tree Aloes
Outgroups
Figure 3.2 One of the most parsimonious trees obtained from the maximum parsimony (MP) analyses
of the combined plastid data set (rbcLa + matK + trnH-psbA) for Alooideae plus outgroups (TL = 752;
CI = 0.72; RI = 0.91). Numbers above the branches are MP bootstrap support.
60
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Chapter three: Results
!
!
H58
88
63
62
50
62
61
99
58
94
68
58
63
52
76
74
52
62
59
64
79
84
100
D
98
68
89
C
96
66
Gasteria vlokii
Gasteria tukhelensis
Gasteria rawlinsonii
Gasteria pulchra
Gasteria polita
Gasteria pillansii var. ernesti-ruschii
Gasteria pillansii var. pillansii
Gasteria glomerata
Gasteria glauca
Gasteria excelsa
Gasteria ellaphieae
Gasteria doreeniae
Gasteria disticha
Gasteria croucheri
Gasteria carinata var. retusa
Gasteria carinata var. carinata
Gasteria obliqua
Gasteria batesiana var. dolomitica
Gasteria nitida var. armstrongii
Gasteria acinacifolia
Haworthia pumila
Haworthia kingiana
Haworthia kingiana
Haworthia marginata
Haworthia marginata
Astroloba corrugata
Astroloba herrei
Astroloba foliolosa
Astroloba rubriflora
Haworthia reinwardtii
Haworthia coarctata
Haworthia coarctata var. adelaidensis
Haworthia reinwardtii var. brevicula
Aloe aristata
Aloe aristata
Aloe aristata
Haworthia venosa subsp. granulata
Haworthia limifolia
Haworthia koelmaniorum var. mcmurtryi
Haworthia koelmaniorum var. mcmurtryi
Haworthia glauca
Haworthia fasciata
Haworthia sordida
Haworthia nigra
Haworthia limifolia var. ubomboensis
Haworthia limifolia
Haworthia bruynsii
Haworthia longiana
Haworthia attenuata
Haworthia venosa
Haworthia cymbiformis var. setulifera
Haworthia cymbiformis
Haworthia cooperi var. doldii
Haworthia bayeri
Haworthia decipiens var. pringlei
Haworthia decipiens var. virella
Haworthia cymbiformis var. ramosa
Haworthia cymbiformis var. setulifera
Haworthia cooperi
Haworthia pulchella var. globifera
Haworthia marxii
Haworthia mucronata var. habdomadis
Haworthia lockwoodii
Haworthia marumiana var. archeri
Haworthia semiviva
Haworthia mucronata var. rycroftiana
Haworthia mucronata var. morrisiae
Haworthia mucronata var. bijliana
Haworthia decipiens
Haworthia herbacea var. flaccida
Haworthia mutica var. nigra
Haworthia magnifica var. dekenahii
Haworthia mirabilis
Haworthia mirabilis var. paradoxa
Haworthia herbacea var. paynei
Haworthia mirabilis var. calcarea
Haworthia reticulata
Haworthia mutica
Haworthia maculata
Haworthia retusa
Haworthia chloracantha
Haworthia floribunda
Haworthia variegata
Haworthia pygmaea
Haworthia emelyae
Haworthia truncata
Haworthia emelyae var. comptoniana
Haworthia monticola var. asema
Haworthia zantneriana
Haworthia wittebergensis
Haworthia vlokii
Haworthia outeniquensis
Haworthia blackburniae var. graminifolia
Haworthia blackburniae var. derustensis
Haworthia blackburniae
Aloe tenuior
Aloe ciliaris
Aloe commixta
Aloe striatula
Aloe gracilis
Gasteria
Robustipendunculares
[F]
Astroloba
Hexangulares
Aloe aristata
[G]
Haworthia subg.
Hexangulares
Haworthia subg.
Haworthia
Aloe Section
Macrifoliae
[Fig 3.3b]
61
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Chapter three: Results
!
!
[Fig 3.3a]
85
75
50
65
96
85
96
91
79
80
61
50
81
70
92
C 62
B 95
81
64
83
100
100
90
100
98
58
100
79
100
100
Aloe kouebokkeveldensis
Aloe buhrii
Aloe striata subsp. kommagasensis
Aloe reynoldsii
Aloe striata
Aloe fouriei
Aloe nubigena
Aloe challisii
Aloe lutescens
Aloe vossii
Aloe verecunda
Aloe saundersiae
Chortolirion angolense
Aloe ecklonis
Aloe spicata
Aloe ferox
Aloe arborescens
Aloe microstigma
Aloe pictifolia
Aloe lineata var. muirii
Aloe lineata var. muirii
Aloe lineata
Aloe thraskii
Aloe rupestris
Aloe comosa
Aloe alooides
Aloe angelica
Aloe munchii
Aloe excelsa
Aloe petricola
Aloe chabaudii
Aloe vryheidensis
Aloe striata subsp. karasbergensis
Aloe hereroensis
Aloe greatheadii var. davyana
Aloe dewinteri
Aloe anivoranoensis
Aloe propagulifera
Aloe perfoliata
Aloe pearsonii
Aloe arenicola
Aloe melanacantha
Aloe tenuior
Aloe ciliaris
Aloe striatula
Aloe plicatilis
Aloe plicatilis
Aloe gracilis
Aloe commixta
Aloe brevifolia
Aloe glauca
Aloe chortolirioides
Aloe albida
Aloe succotrina
Aloe dichotoma subsp. pillansii
Aloe eminens
Aloe dichotoma
Aloe dichotoma subsp. ramosissima
Aloe barberae
Bulbine frutescens
Bulbine semibarbata
Jodrellia fistulosa
Kniphofia uvaria
Kniphofia galpinii
Eremurus spectabilis
Asphodeline lutea
Xanthorrhoea sp.
Xanthorrhoea sp.
Xanthorrhoea resinosa
Zephyra elegans
Tecophilaea cyanocrocus
Anthericum liliago
Section
Leptoaloe
Chortolirion
Single−
stemmed
Aloes
[E]
True Aloes
Section
Macrifoliae
[A]
Tree Aloes
Outgroups
Figure 3.3 One of the most parsimonious trees obtained from the maximum parsimony (MP) analyses
of the ITS1 data set for Alooideae (TL = 346; CI = 0.64; RI = 0.92).
62
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Chapter three: Results
!
!
−
0.77
58
0.98
63
0.83
61
− 0.95
0.57
H
70
0.99
97
1.00
80
1.00
G
92
− 1.00
0.93
88
1.00
−
− 0.74
54 0.99
−
− 0.98
0.55
78 0.55
− 1.00
57 0.78
1.00
−
0.64
67
81 0.88
− 1.00
0.75
82
1.00
92
1.00
−
0.67
100
1.00
−
0.74
F
77
0.95
100
75 1.00
79 0.97
0.99
58
0.68
100
1.00
99
1.00
61
0.93
89
1.00
85
− 1.00
−
95 0.77
0.59 1.00
56
0.97
−
0.60
91
− 1.00
− 0.51
55 0.92
0.99
86
1.00
61
1.00
71
1.00
70
1.00
66
51 0.89
84 0.90
53 0.93
0.99
E
−
1.00
−
53 0.73
0.92
90
1.00
−
0.60
80
1.00
80
0.73
G. polita
G. doreeniae
G. disticha
G. tukhelensis
G. pulchra
G. glomerata
G. glauca
G. rawlinsonii
G. obliqua
G. vlokii
G. excelsa
G. ellaphieae
G. batesiana var. dolomitica
G. nitida var. armstrongii
G. acinacifolia
G. carinata var. retusa
G. carinata var. carinata
G. croucheri
G. pillansii var. ernesti-ruschii
G. pillansii var. pillansii
Haworthia coarctata
Haworthia reinwardtii var. brevicula
Haworthia coarctata var. adelaidensis
Haworthia reinwardtii
Haworthia nigra
Haworthia glauca
Haworthia bruynsii
Haworthia sordida
Haworthia longiana
Haworthia fasciata
Haworthia limifolia var. ubomboensis
Haworthia limifolia
Haworthia limifolia
Haworthia venosa var. granulata
Haworthia venosa
Haworthia attenuata
Haworthia kingiana
Haworthia marginata
Haworthia pumila
Haworthia kingiana
Haworthia marginata
Astroloba corrugata
Astroloba herrei
Astroloba foliolosa
Astroloba rubriflora
Aloe aristata
Aloe aristata
Aloe aristata
H. koelmaniorum var. mcmutryi
H. koelmaniorum var. mcmutryi
Aloe nubigena
Aloe ecklonis
Aloe fouriei
Aloe challisii
Aloe vossii
Aloe verecunda
Aloe saundersiae
Aloe kouebokkeveldensis
Aloe buhrii
Aloe striata subsp. kommagasensis
Aloe striata
Aloe reynoldsii
Aloe lutescens
Chortolirion angolense
Aloe spicata
Aloe ferox
Aloe arborescens
Aloe microstigma
Aloe pictifolia
Aloe lineata var. muirii
Aloe lineata var. muirii
Aloe lineata
Aloe thraskii
Aloe rupestris
Aloe comosa
Aloe alooides
Aloe angelica
Aloe excelsa
Aloe petricola
Aloe munchii
Aloe chabaudii
Aloe vryheidensis
Aloe perfoliata
Aloe arenicola
Aloe pearsonii
Aloe melanacantha
Aloe chortolirioides
Aloe albida
Aloe brevifolia
Aloe glauca
Aloe striata subsp. karasbergensis
Aloe greatheadii subsp. davyana
Aloe propagulifera
Aloe dewinteri
Aloe anivoranoensis
Aloe haworthioides
Aloe hereroensis
Aloe succotrina
Gasteria
Hexangulares I
Robustipendunculares
Astroloba
Aloe aristata
Hexangulares II
Section
Leptoaloe
Single−
stemmed True Aloes
Aloes
[Fig 3.4b]
63
!
Chapter three: Results
!
!
[Fig 3.4a]
53
− 0.65
0.61
−
0.62
−
0.59
61
− 0.96
0.56
−
0.61
63
0.80
58
0.92
−
0.99
−
0.79
−
0.76
−
0.53
−
70 0.82
0.88
−
0.74
D
61
0.82
100
1.00
98
1.00
100
1.00
93
1.00
87
74 0.99
95 0.96
1.00
67
0.94
97
1.00
C
80
1.00
B
A
100
1.00
100
1.00
93
76 1.00
1.00 −
1.00
99
90 1.00
1.00
100
− 1.00
0.51 63
0.90
74
100 0.92
1.00
100
99 1.00
1.00
Haworthia herbacea var. paynei
Haworthia reticulata
Haworthia maculata
Haworthia herbacea var. flaccida
Haworthia magnifica var. dekenahii
Haworthia mirabilis var. paradoxa
Haworthia mirabilis var. calcarea
Haworthia mirabilis var. triebneriana
Haworthia retusa
Haworthia mutica var. nigra
Haworthia mutica
Haworthia marumiana var. archeri
Haworthia semiviva
Haworthia marxii
Haworthia floribunda
Haworthia emelyae
Haworthia wittebergensis
Haworthia vlokii
Haworthia cymbiformis var. setulifera
Haworthia cymbiformis
Haworthia cooperi var. doldii
Haworthia decipiens var. pringlei
Haworthia decipiens var. virella
Haworthia cymbiformis var. ramosa
Haworthia cymbiformis var. setulifera
Haworthia cooperi
Haworthia mucronata var. habdomadis
Haworthia lockwoodii
Haworthia pulchella var. globifera
Haworthia monticola var. asema
Haworthia mucronata var. rycroftiana
Haworthia mucronata var. morrisiae
Haworthia mucronata var. bijliana
Haworthia arachnoidea
Haworthia arachnoidea
Haworthia decipiens
Haworthia outeniquensis
Haworthia chloracantha
Haworthia pygmaea
Haworthia variegata
Haworthia truncata
Haworthia emelyae var. comptoniana
Haworthia angustifolia
Haworthia zantneriana
Haworthia bayeri
Haworthia mirabilis
Haworthia blackburniae var. graminifolia
Haworthia blackburniae var. derustensis
Haworthia blackburniae
Aloe tenuior
Aloe ciliaris
Aloe striatula
Aloe gracilis
Aloe commixta
Aloe plicatilis
Aloe plicatilis
Aloe dichotoma subsp. pillansii
Aloe dichotoma
Aloe dichotoma subsp. ramosissima
Aloe eminens
Aloe barberae
Bulbine frutescens
Bulbine semibarbata
Jodrellia fistulosa
Kniphofia uvaria
Kniphofia galpinii
Eremurus spectabilis
Asphodeline lutea
Xanthorrhoea sp.
Xanthorrhoea sp.
Xanthorrhoea resinosa
Zephyra elegans
Tecophilaea cyanocrocus
Anthericum liliago
Haworthia subg.
Haworthia
Section
Macrifoliae
Aloe plicatilis
Tree Aloes
Outgroups
Figure 3.4 Majority-rule consensus tree obtained from the Bayesian analysis of the combined plastid
and nuclear internal transcribed spacer (ITS1) data set for Alooideae plus outgroup (TL = 1159; CI =
0.63; RI = 0.89). Numbers above the branches are MP bootstrap support and ones below are BI
posterior probabilities.
64
!
Chapter three: Results
!
Per_col
Per_tube
16
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Ped_length
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Per_symmetry
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Tepals
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Fl_arrange
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Perianth
Fl_orient
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Astroloba
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Aloe aristata ●
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5
!
G. polita
G. doreeniae
G. disticha
G. tukhelensis
G. pulchra
G. glomerata
G. glauca
G. rawlinsonii
G. obliqua
G. vlokii
G. excelsa
G. ellaphieae
G. batesiana var dolomitica
G. nitida var armstrongii
G. acinacifolia
G. carinata var. retusa
G. carinata var. carinata
G. croucheri
G. pillansii var. ernesti−ruschii
G. pillansii var. pillansii
H. coarctata
H. reinwardtii var. brevicula
H. coarctata var. adelaidensis
H. reinwardtii
H. nigra
H. glauca
H. bruynsii
H. sordida
H. longiana
H. fasciata
H. limifolia var. ubomboensis
H. limifolia
H. limifolia
H. venosa var. granulata
H. venosa
H. attenuata
H. kingiana
H. marginata
H. pumila
H. kingiana
H. marginata
Astroloba corrugata
Astroloba herrei
Astroloba foliolosa
Astroloba rubriflora
A. aristata
A. aristata
A. aristata
H. koelmaniorum var. mcmurtryi
H. koelmaniorum var. mcmurtryi
A. nubigena
A. ecklonis
A. fouriei
A. challisii
A. vossii
A. verecunda
A. saundersiae
A. kouebokkeveldensis
A. buhrii
A. striata subsp. kommagasensis
A. striata
A. reynoldsii
A. lutescens
Chortolirion angolense
A. spicata
A. ferox
A. arborescens
A. pictifolia
A. microstigma
A. lineata var. muirii
A. lineata var. muirii
A. lineata
A. thraskii
A. rupestris
A. comosa
A. alooides
A. angelica
A. excelsa
A. petricola
A. munchii
A. chabaudii
A. vryheidensis
A. striata subsp. karasbergensis
A. greatheadii var. davyana
A. hereroensis
A. propagulifera
A. dewinteri
A. anivoranoensis
A. haworthioides
A. brevifolia
A. glauca
A. succotrina
A. perfoliata
A. arenicola
A. pearsonii
A. melanacantha
A. eminens
A. chortolirioides
A. albida
A. tenuior
A. ciliaris
A. striatula
A. gracilis
A. commixta
H. herbacea var. paynei
H. reticulata
H. maculata
H. herbacea var. flaccida
H. magnifica var. dekenahii
H. mirabilis var. paradoxa
H. mirabilis var. calcarea
H. mutica var. nigra
H. mutica
H. mirabilis var. triebneriana
H. retusa
H. floribunda
H. emelyae
H. wittebergensis
H. vlokii
H. marumiana var. archeri
H. semiviva
H. marxii
H. cymbiformis var. setulifera
H. cymbiformis
H. cooperi var. doldii
H. decipiens var. pringlei
H. decipiens var. virella
H. cymbiformis var. ramosa
H. cymbiformis var. setulifera
H. cooperi
H. mucronata var. habdomadis
H. lockwoodii
H. pulchella var. globifera
H. mucronata var. rycroftiana
H. mucronata var. morrisiae
H. mucronata var. bijliana
H. arachnoidea
H. arachnoidea
H. decipiens
H. chloracantha
H. pygmaea
H. variegata
H. truncata
H. emelyae var. comptoniana
H. monticola var. asema
H. angustifolia
H. zantneriana
H. bayeri
H. outeniquensis
H. mirabilis
H. blackburniae var. graminifolia
H. blackburniae var. derustensis
H. blackburniae
A. plicatilis
A. plicatilis
A. dichotoma subsp. pillansii
A. dichotoma
A. dichotoma subsp. ramosissima
A. barberae
Bulbine frutescens
Bulbine semibarbata
Jodrellia fistulosa
Kniphofia uvaria
Kniphofia galpinii
Eremurus spectabilis
Asphodeline lutea
Xanthorrhoea sp.
Xanthorrhoea sp.
Xanthorrhoea resinosa
Zephyra elegans
Tecophilaea cyanocrocus
Anthericum liliago
Figure 3.4: Morphological trait distribution along phylogeny of Alooideae. All trait values are centred and scaled, resulting in circles of different colours.
Figure 3.5 Distribution of morphological traits along phylogeny of Alooideae. All trait values are
centred and scaled, resulting in circles of different colours. The colours of the symbols are proportional
to the absolute value of each trait.
65
!
Chapter three: Results
!
!
= stemless
= caulescent
= arborescent
a
b
= polystichous
= distichous
= concolorous
= white
c
C
Figure 3.6 Parsimony-based reconstructions of a) habit character (character 1), b) leaf insertion
(character 2), and c) leaf tubercles (character 5) on the majority-rule consensus tree from the combined
molecular data.
66
!
Chapter three: Results
!
!
= spreading
= pendulous
a
b
= actinomorphic
= weakly bilabiate
= strongly bilabiate
= straight
= curved upwards
Cc
Figure 3.7 Parsimony-based reconstructions of a) flower orientation (character 11), b) periath
symmetry (character 14), and c) perianth tube (character 17) on the majority-rule consensus tree from
the combined molecular data.
67
!
Chapter three: Results
!
!
= cylindrical and tapering into pedicel
= inflated basally
a
b
= exserted
= included
= both whorls connate basally
= both whorls connate 1/2 or more of length
= only outer whorls connate for 1/2 or more of length
c
Figure 3.8 Parsimony-based reconstructions of a) periath shape (character 18), b) stamen inclusion
(character 20), and c) tepal fusion (character 19) on the majority-rule consensus tree from the
combined molecular data.
68
!
Chapter four: Discussion and Conclusion
!
!
CHAPTER(FOUR(
Discussion(and(Conclusion(
!
69
!
Chapter four: Discussion and Conclusion
!
!
4. DISCUSSION AND CONCLUSIONS
4.1 Phylogenetic Relationships within Alooideae
Based on congruence between the plastid and nuclear DNA data sets, the discussion is
restricted to results obtained from the combined data set (Fig. 3.4). Results strongly confirm
the monophyly of Alooideae as previously shown by Linnaeus (1753) and more recently by
Treutlein et al. (2003a, b) and Ramdhani et al. (2011). Furthermore, the genus Aloe is
polyphyletic as currently circumscribed, with all other genera of Alooideae nested in different
parts of Aloe. The analysis identifies seven primary monophyletic lineages in Alooideae,
corresponding to the following currently recognised generic and infrageneric groups:
1. Aloe sect. Dracoaloe + Aloe sect. Aloidendron
2. Aloe sect. Macrifoliae
3. Haworthia subg. Haworthia
4. Haworthia subg. Robustipendunculares + Astroloba + Aloe sect. Aristatae
5. Haworthia subg. Hexangulares
6. Gasteria
7. Remaining Aloe spp.
This topology is congruent with that derived earlier by Treutlein et al. (2003b) with
much smaller taxon sampling and analysis of just two plastid gene regions. Among the smaller
segregate genera, only Gasteria and Astroloba are retrieved as monophyletic (Chortolirion is
monotypic). The analysis indicated that Haworthia is not monophyletic but rather represents
three lineages corresponding more or less to the subgenera proposed by Bayer (1982, 1999).
70
!
Chapter four: Discussion and Conclusion
!
!
4.1.1 Relationships within Aloe
Relationships within Aloe sect. Dracoaloe + sect. Aloidendron ! The analysis did
not retrieved the ‘tree aloes’ sensu Van Wyk & Smith (2003) as a monophyletic group, but
rather as two separate lineages. Lineage 1 includes sections Aloidendron + Dracoaloe which
appears to be the earliest diverging lineage within the subfamily. Aloe section Dracoaloe (A.
dichotoma, A. dichotoma subsp. ramosissima, and subsp. pillansii) known as the “Quiver tree”
(Van Wyk & Smith, 2003), occurs in the Northern Cape region and Namibia. Capitate raceme
is typical of this group (Glen & Hardy, 2000). The section Aloidendron comprise only of Aloe
barberae Dyer, which is the largest African Aloe species (growing up to 10-18 m tall). It is
distributed along the eastern coast of southern Africa (Mpumalanga, Swaziland, KwazuluNatal, Eastern Cape and Mozambique). Aloe plicatilis (lineage 2) (also tree Aloe sensu Van
Wyk & Smith, 2003, endemic to the Cape Floristic region) did not group with the other ‘tree
aloes’, but was retrieved as the early diverging lineage to Haworthia subgenus Haworthia.
Although all three sections share a tree-like growth form, arborescence appears to have
developed convergently in several Aloe lineages. I also recovered a strong support for A.
eminens grouping with A. barberae (A. bainesii sensu Reynolds, 1966). This grouping might
be a confirmation of their similar arborescent habit, although they are geographically isolated
(A. barberae from South Africa and A. eminens from Somalia) plus the slight differences in
some floral details, my analysis places them together as part of the earliest diverging lineage
within Alooideae.
Relationships within Aloe section Macrifoliae ! Aloe sect. Macrifoliae, the
‘rambling Aloes’ (sensu Van Wyk & Smith, 2003), comprising five closely related species (A.
ciliaris, A. commixta, A. gracilis, A. striatula, and A. tenuior), is recovered as a strongly
supported clade sister to the true aloes + all segregate genera. The cane-like stems of these
71
!
Chapter four: Discussion and Conclusion
!
!
species, with their slender, sheathing, unspotted and mesophytic leaves with minute marginal
teeth are unique to members of this section (Glen & Hardy, 2000).
Relationships within the true Aloe species ! The relationships among the remaining
species of Aloe included in the analysis are poorly resolved. Although some of the currently
recognised sections may be monophyletic, other sections are clearly not, and a much more
extensive sampling is required to evaluate taxonomic and evolutionary relationships among
them. The position of Chortolirion is noteworthy, where it is deeply embedded within this
group, as part of a clade including most of the grass aloes. However the precise relationships
are still unclear. Close morphological similarity in vegetative parts, namely the grass-like
leaves with bulb-like swelling, and the small, bilabiate flowers, to species such as A. bowiea
and A. inconspicua suggest a close relationship to sect. Leptoaloe. This resemblance between
Chortolirion and the grass aloes was also pointed out in previous studies (e.g. Mabberley,
2008; Treutlein et al., 2003b).
4.1.2 Relationships within Haworthia
Relationships within Haworthia subg. Haworthia ! Species of H. subgenus Haworthia
comprise a strongly supported monophyletic clade, defined morphologically by the basally
triangular perianth, obclavate flowers and upcurved style (Bayer, 1982, 1999). Additional
support for this comes from Smith et al. (2001), which reported the occurrence of lower (<
than 50 %) sucrose concentrations in H. subg. Haworthia in contrast to higher (> 60 %)
sucrose concentrations in subg. Hexangulares and Robustipendunculares.
Relationships within Haworthia subg. Robustipendunculares + Astroloba + Aloe
sect. Aristatae ! Haworthia subg. Robustipendunculares is a small group of three species
(Haworthia marginata, H. kingiana, and H. pumila) that is well-supported as monophyletic
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Chapter four: Discussion and Conclusion
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!
and sister to Astroloba. Members of subg. Robustipendunculares are often large in size, about
100-300 mm across (with the largest been H. pumila) and restricted to the southern parts of the
Western Cape, from Worcester to Mossel Bay. It is defined morphologically by its more or
less straight perianth abruptly joined to the pedicel (Bayer, 1982, 1999). The flower type
found in H. subg. Robustipendunculares is not dissimilar to that in Astroloba, differing
essentially in its slight zygomorphy. Members of Robustipendunculares are often robust with
attenuate leaves, often scabrid and patterned with white tubercles (Bayer, 1999). Sister to
subg. Robustipendunculares + Astroloba is Aloe aristata, which is unique in the genus Aloe by
having “Haworthia-like” leaves with dry awn-tipped apices and white tubercles (Glen &
Hardy,
2000).
The
sister
relationship
between
Astroloba
and
Haworthia
subg.
Robustipendunculares is further supported by similarities in nectar sucrose proportions (Van
Wyk et al., 1993).
Relationships within Haworthia subg. Hexangulares ! Bayer (1999) recognised 16
species within H. subg. Hexangularis. The current analysis however, indicates that H. subg.
Hexangulares is polyphyletic, with H. koelmanniorum more closely related to H. subg.
Robustipendunculares. Haworthia koelmanniorum occupies an isolated position sister to subg.
Robustipendunculares + Astroloba + Aloe aristata. The geographical distribution of this
species is also disjunct from that of other species of Haworthia. Significantly, species of
Hexangulares display the largest vegetative diversity in Haworthia with some closely
resembling members of Astroloba and Robustipendunculares respectively, in their vegetative
morphology.
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Chapter four: Discussion and Conclusion
!
!
4.1.3 Relationships within smaller segregate genera
Relationships within Astroloba ! Astroloba, consisting of 6 species, are retrieved as a
monophyletic clade sister to Haworthia subg. Robustipendunculares. The genus is
morphologically defined by its caulescent habit with stiff, imbricate leaves, and small,
actinomorphic flowers with included stamens. Results from this study confirm the inclusion of
Poellnitzia rubriflora within Astroloba.
Relationships within Gasteria ! Gasteria forms a strongly supported monophyletic
clade with several morphological synapomorphies, notably the unarmed, verrucose leaves and
inclined secund inflorescences of pendulous, gasteriform flowers with a well-developed floral
tube (Van Jaarsveld, 2007). Bimodal karyotype is uniform in all Gasteria species and consists
of eight large and six chromosomes (Vosa & Bennett, 1990).
4.2 Evolution of selected morphological characters in Alooideae
I analysed 20 morphological characters commonly used to differentiate between Alooideae
species, following extensive literature review (as presented in section 2.5). I then
reconstructed the likely evolutionary history of nine morphological characters that are
important in generic and infrageneric delimitation within the subfamily Alooideae onto the
majority rule consensus tree from the BI. The importance of these selected characters in
Alooideae systematics are not only employed in previous Alooideae systematics (e.g. Bayer,
1999; Smith & van Wyk, 1991; Van Jaarsveld, 1994), they are shown to be unique to certain
clades in this study.
Habit. ! Acaulescence is plesiomorphic within Alooideae, with certain lineages
characterised by secondary caulescence and arborescence (Fig. 3.6a). Caulescence has
74
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Chapter four: Discussion and Conclusion
!
!
developed several times in the subfamily, including Aloe itself (notably section Macrifoliae)
and in some of the segregate genera, notably Astroloba and also some species of Gasteria and
Haworthia subg. Hexangulares. Arborescence is relatively uncommon in the subfamily but is
characteristic of species in Aloe sect. Aloidendron and Dracoaloe, the clade including A.
eminens. It is implicit in previous classifications (e.g. Reynolds, 1966; Holland, 1978) that
arborescence in Aloe is the derived state. Although the cytogenetic analysis by Brandham
(1983) proposed that scandent Aloe species with usually relatively mesophytic leaves, e.g. A.
tenuior Haw. and A. ciliaris Haw. (sect. Macrifoliae), represent the primitive state in Aloe
species, our analysis suggests that caulescence is derived in the genus. Smith & van Wyk
(1991) argued that both small, highly succulent taxa and arborescent forms were derived from
a mesophytic, comparatively acaulescent taxon. The current study supports this hypothesis,
showing that arborescence in Aloe is found not only in early diverging lineages but also in
others deeply embedded within Aloe, and that the grass Aloes (sect. Leptoaloe) are clearly
derived.
Leaf insertion ! Polystichous leaf insertion is common within the subfamily, and
distichous leaf insertion is rare, having evolved independently several times in Aloe, once in
Haworthia and possibly once in Gasteria (Fig. 3.6b). Distichy is evidently the juvenile
condition, present in all Aloe and Gasteria seedlings, and its persistence in adult plants is best
interpreted as neoteny. The ecological advantages are unclear.
Leaf tuberculation ! The presence of white tubercles on the leaves is largely
restricted to Gasteria and Haworthia subg. Hexangulares and Robustipendunculares but
tubercules are also developed in some Aloe species (e.g. Aloe aristata, A. haworthioides and
A. verrecunda) and in the genus Chortolirion (Fig. 3.6c). Tuberculation is certainly a derived
75
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Chapter four: Discussion and Conclusion
!
!
condition, as hypothesised by Smith & van Wyk (1991). In Gasteria, Van Jaarsveld (1994)
has proposed that its evolution was driven by the absence of the bitter constituent typical of
Aloe species, suggesting that the rigid tubercles may make the leaves less palatable. This
remains to be tested.
Perianth colour ! Vivid, yellow, orange or reddish flowers are clearly plesiomorphic
for the subfamily, and are strongly associated with ornithophily. The whitish or greenish
perianth that is characteristic of Haworthia and Astroloba (excluding Poellnitzia) and several
species of Aloe, especially several members of sect. Leptaloae, appear to be adaptations to
entomophily (Botes et al., 2008; Hargreaves et al., 2008) and are derived.
Flower orientation at anthesis ! Representatives of Astroloba, Chortolirion and
Haworthia, bear flowers ascending on vertical peduncles. This is hypothesised by Smith &
Van Wyk (1991) to be the derived condition, which developed in response to a specialised,
insect pollination syndrome. Pendulous flowers at anthesis are found in all Gasteria and some
Aloe species. Flower orientation in Gasteria appears to be an adaptation to bird pollination,
the horizontal peduncles acting as perches for bird visitors such as sunbirds (Nectariidae;
Smith & Van Wyk, 1991), enabling easy access to nectar reward.
Tepal connation ! The perianth in Alooideae comprises six tepals arranged in two
whorls of three each, variously connate into a short or prominent tube. The plesiomorphic
condition is for the tepals to be basally connate only, and this is charateristic even in bird
pollinated Aloe species (Fig. 3.8c). In several of the ornithopilous Aloe species, however, the
outer tepal whorl is connate in the basal half, forming a distinct perianth tube. Tepals with
both whorls connate for half or more of their length are diagnostic for Gasteria, Astroloba
(Poellnitzia) rubriflora, and Aloe kouebokkeveldensis and appear to have evolved
76
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Chapter four: Discussion and Conclusion
!
!
independently in these three lineages from ancestral types with tepals connate at the base only.
Floral syndromes indicate that all of these taxa are bird pollinated and the results from this
study suggest that in Gasteria and Astroloba these are secondary modification of an
entomophilous Haworthia-type flower rather than developments from a more typical Aloetype flower.
Haworthia-type flowers are small, spreading flowers with whitish, sometimes strongly
bilabiate perianths, and included anthers are diagnostic of the genus Haworthia (Bayer, 1999).
Cladistic analysis suggests that this flower type is homoplasious, and that it has evolved
independently at least three and possibly four times (once each in Chortolirion and H. subg.
Haworthia and once or twice in subg. Hexangulares/Robustipendunculares with possible
reversals in A. aristata and Gasteria) (Figs. 3.7a & b). Relatively short-tubed whitish or
sometimes cream flowers in some Aloe species (e.g. Aloe inconspicua) have been revealed as
an adaptation to insect pollination (Botes et al., 2009). Although still recognisably ‘Aloe-like’,
the flowers of bee-pollinated Aloe species (sensu Botes et al., 2009) such as A. linearifolia and
A. minima, display several characteristics of the Haworthia-flower apart from reduced size,
namely their nearly horizontal orientation, whitish and weakly bilabiate perianth, and
sometimes included stamens. In term of perianth symmetry, Smith & van Wyk (1991)
suggested that zygomorphy (as revealed from the flowers of Astroloba, Chortolirion and
Haworthia) represents an advanced state derived from the plesiomorphic actinomorphic
pattern, and the results in this study confirmed this hypothesis.
Gasteria-type flowers are curved and flask-shaped with an ovoid, inflated tube at least
half as long as the perianth, and included or shortly exserted stamens, is characteristic of the
genus Gasteria. The flowers in the genus are also often bicoloured, with greenish tips to the
tepals, and are borne secund on inclined racemes. Flask-shaped flowers with inflated bases
77
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Chapter four: Discussion and Conclusion
!
!
occur widely in Aloe and have evidently evolved several times (Fig. 3.8a), although in this
genus the stamens are often but not always exserted (Fig. 3.8b). Similar flowers are
characteristic also of Poellnitzia but here are uniquely held erect on an inclined raceme. A
well-developed perianth tube formed by the fusion of both tepal whorls has evolved
independently in Gasteria, Astroloba and Aloe kouebokkeveldensis (Fig. 3.8c).
78
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Chapter four: Discussion and Conclusion
!
!
4.3 Conclusions
Results of this study were largely conducted with new data collected for four gene regions
(rbcLa, matK, trnH-psbA and ITS1), with greater sampling than in any previously published
analyses (e.g. Treutlein et al., 2003a, b; Ramdhani et al., 2011) and essentially congruent with
those of earlier studies. Evidence from morphological (Smith & Steyn, 2004), cytological
(Taylor, 1925), cladistic (Smith & Van Wyk, 1991; Klopper et al., 2010) and chemical
analyses (Viljoen et al., 1998; Viljoen, 1999) have been routinely applied to the current
classification of Alooideae, which recognises five genera. These studies, although unable to
resolve the relationships within the subfamily, provided a working hypothesis for this study.
Gasteria is well defined morphologically by its verrucose, edentate leaves and inclined
inflorescences with ± secund, ‘gasteriform’ flowers, and it is not coincidental that it was one
of the earliest groups of species to be recognised. The phylogenetic analysis places Gasteria
sister to Haworthia subg. Hexangulares, which includes species with remarkably similar
leaves, and the unique Gasteria-type flowers, are most parsimoniously interpreted as a
reversion to bird-pollination from the entomophilous Haworthia-type flower with included
stamens. One of the most unexpected results of the analyses is the apparent convergent
evolution of the Haworthia-type flower. It appears to have evolved at least three times: at least
twice and possibly three times in Haworthia itself and independently in Chortolirion. The
close phylogenetic association between Poellnitzia, Astroloba and Haworthia subg.
Robustipendunculares is reflected in the vegetative similarity among these three genera, and
the floral differences between them are in effect very minor, relying on differences in perianth
symmetry, size and colour. Indeed, Manning & Smith (2000) proposed the inclusion of
Poellnitzia in Astroloba and this was confirmed in the current study. It is significant that Aloe
aristata, treated as the monoptypic sect. Aristatae on the basis of its distinctive foliage
79
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Chapter four: Discussion and Conclusion
!
!
(edentate, aristate and strikingly marked with transverse verrucae), is shown to be closely
allied with H. sect. Robustipendunculares + Astroloba. Vegetatively, A. aristata would fit
perfectly in H. sect. Robustipendunculares.
It is thus clear that few of the characters used to define genera within Alooideae at
present are autapomorphies, and Aloe already includes most of the variation evident in the
subfamily. Furthermore, Aloe is shown to be polyphyletic with the other four Alooideae
genera nested in different parts of Aloe. There is an increasing trend to align classifications
with phylogenies at all taxonomic levels, and for taxa to be circumscribed as monophyletic
lineages (Backlund & Bremer, 1998). In Alooideae there are essentially just two options
available. The first ! splitter’s approach ! requires the recognition of at least four and
possibly more additional segregates among Aloe in order to render a rump Aloe monophyletic,
plus the fragmentation of Haworthia into three smaller genera. The second ! lumper’s
approach ! requires combining all of the genera into a single genus Aloe, with appropriate
infrageneric groups.
Finally, the results present a milestone in Alooideae systematics with a phylogenydriven generic classification proposed in which Aloe is expanded to include all members of
Alooideae (Appendices II & III).
!
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References
!
!
CHAPTER(FIVE(
References(
!
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!
References
!
!
REFERENCES
Adams SP, Leitch IJ, Bennett MD, Leitch AR. 2000b. Aloe L. – a second plant family
without (TTTAGGG)n telomeres. Chromosoma 109: 201–205.
Adams SP, Leitch IJ, Bennett MD. 2000a. Ribosomal DNA evolution and phylogeny in
Aloe (Asphodelaceae). American Journal of Botany 87: 1578–1583.
APG II (Angiosperm Phylogeny Group). 2003. An update of the Angiosperm
Phylogeny Group, classification for the orders and families of flowering plants:
APG II. Botanical Journal of the Linnean Society 141: 399–436.
APG III (Angiosperm Phylogeny Group) 2009. An update of the Angiosperm
Phylogeny Group classification for the orders and families of flowering plants:
APG III. Botanical Journal of the Linnean Society 161: 105–121.
Backlund A, Bremer K. 1998. To be or not to be - principles of classification and
monotypic plant families. Taxon 47: 391—400.
Baldwin BG, Sanderson MJ, Porter JM, Wojciechowski MF, Campbell CS,
Donoghue MJ. 1995. The ITS region of the nuclear ribosomal DNA: A valuable
source of evidence on angiosperm phylogeny. Annals of the Missouri Botanical
Garden 82: 247—277.
Bayer MB. 1982. The new Haworthia handbook. National Botanic Gardens, South
Africa.
Bayer MB. 1999. Haworthia revisited. A revision of the genus. Umdaus Press, Hatfield
South Africa.
Bayer MB. 2002. Haworthia update: Essays on Haworthia-Volume 1. Umdaus Press,
Pretoria, South Africa.
Bayer MB. 2009. Haworthia supplement. The Haworthia Society, London.
82
!
References
!
!
Botes C, Johnson SD, Cowling RM. 2008. Coexistence of succulent tree aloes:
partitioning of bird pollinators by floral traits and flowering phenology. Oikos
117: 875—882.
Botes C, Wragg PD, Johnson SD. 2009. New evidence for bee-pollination systems in
Aloe (Asphodelaceae: Aloideae), a predominantly bird-pollinated genus. South
African Journal of Botany 75: 675—681.
Brandham PE. 1977. The inheritance of leaf pigmentation in Gasteria (Liliaceae). Kew
Bulletin 32: 13—17.
Brandham PE. 1983. Evolution in a stable chromosome system. In: Brandham PE,
Bennett MD. (Eds.). Kew Chromosome Conference II London.
CBOL Plant Working Group. 2009. A DNA barcode for land plants. Proceedings of
the National Academy of Sciences, USA 106: 12794—12797.
Chase MW, De Bruijn AY, Cox AV, Reeves G, Rudall PJ, Johnson MAT, Eguiarte
LE. 2000. Phylogenetics of Asphodelaceae (Asparagales): An analysis of plastid
rbcL and trnL-F DNA sequences. Annals of Botany 86: 935–951.
Chase MW, Reveal JL, Fay MF. 2009. A subfamilial classification for the expanded
asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae.
Botanical Journal of the Linnean Society 161: 132–136.
Cronquist A. 1981. An integrated system of classification of flowering plants. Columbia
University Press, New York.
Devey DS, Leitch I, Rudall PJ, Pires JC, Pillon Y, Chase MW. 2006. Systematics of
Xanthorrhoeaceae sensu lato, with emphasis on Bulbine. Aliso 22: 345—351.
Doyle JJ, Doyle JL. 1987. A rapid DNA isolation procedure for small quantities of fresh
leaf tissue. Phytochemical Bulletin 19: 11—15.
83
!
References
!
!
Duval HA. 1809. Plantae succulentae, in Horto Alenconio. Parisiis apud Gabon et
Socios, Paris.
Edgar RC. 2004. MUSCLE: multiple sequence alignment with high accuracy and high
throughput. Nucleic Acids Research 32: 1792-97.
Felsenstein J. 1985. Confidence levels on phylogenies: an approach using the bootstrap.
Evolution 39: 783—791.
Germishuizen G, Meyer NL, Steenkamp Y, Keith M. eds. 2006. A checklist of South
African plants. Southern African Botanical Diversity Network Report No. 41.
SABONET, Pretoria, South Africa.
Gildenhuys SD. 2007. Haworthia marxii (Asphodelaceae/Aloaceae), a new species from
the Little Karoo, South Africa. Aloe 44: 4—8.
Glen HF, Hardy DS. 2000. Aloaceae. Aloe. In: Germishuizen G. Ed. Flora of Southern
Africa. Fascicle 1: Aloaceae (First part): Aloe, vol. 5. National Botanical
Institute, Pretoria, pp. 1-167.
Glen HF, Smith GF. 1995. Notes on the typification of some species of Aloe
(Asphodelaceae/Aloaceae). Bothalia 25: 37—42.
Grace OM, Simmonds MSJ, Smith GF, Wyk AE. 2009. Taxonomic significance of
leaf surface morphology in Aloe section Pictae (Xanthorrhoeaceae). Botanical
Journal of the Linnean Society 160: 418–428.
Grace OM, Simmonds MSJ, Smith GF, Wyk AE. 2010. Chemosystematic evaluation
of Aloe section Pictae (Asphodelaceae). Biochemical Systematics and Ecology
38: 57–62.
Hargreaves AL, Harder LD, Johnson SD. 2008. Aloe inconspicua: The first record of
an exclusively insect-pollinated aloe. South African Journal of Botany 74: 606–
612.
84
!
References
!
!
Hillis DM, Dixon MT. 1991. Ribosomal DNA: Molecular evolution and phylogenetic
inference. The Quarterly Review of Biology 66: 411—453.
Holland PG. 1978. An evolutionary biogeography of the genus Aloe. Journal of
Biogeography 5: 213—226.
Huelsenbeck JP, Ronquist F. 2001. MRBAYES, Bayesian inference of phylogenetic
trees. Bioinformatics 17: 754—755.
IUCN 2001. IUCN Red List Categories and Criteria: version 3.1. IUCN Species Survival
Commision. Gland, Switzerland, and Cambridge, UK.
Jeppe B. 1969. South African Aloes. Purnell, Cape Town, South Africa.
Jombart T, Dray S. 2010. adephylo: exploratory analyses for the phylogenetic
comparative method. Bioinformatics 26: 1907—1909.
Kim JH, Kim DK, Forest F, Fay MF, Chase MW. 2010. Molecular phylogenetics of
Ruscaceae sensu lato and related families (Asparagales) based on plastid and
nuclear DNA sequences. Annals of Botany 106: 775—790.
Klopper RR, Smith GF. 2007. The genus Aloe L. (Apshodelaceae: Alooideae) in
Namqualand, South Africa. Haseltonia 13: 1—13.
Klopper RR, van Wyk AE, Smith GF. 2010. Phylogenetic relationships in the family
Asphodelaceae (Asparagales). Biodiversity and Ecology3: 9-36.
Linnaeus C. 1753. Species plantarum. Editio Secunda, aucta. Impensis Direct, Laurentii
Salvii, Stockholm, Sweden.
Linnaeus C. 1774. Systema Vegetabilium: Secundum Classes, Ordines, Genera, Species.
Cum Characteribus Differentiis Et Synonymis, Volume 7, Part 1.
Mabberley DJ. 2008. The Plant-Book – A portable dictionary of the vascular plants,
3rdedn. Cambridge: Cambridge University Press.
85
!
References
!
!
Maddison WP, Maddison DR. 2011. Mesquite: a modular system for evolutionary
analysis. Version 2.75, http://mesquiteproject.org
Manning JC, Smith GF. 2000. The genus Poellnitzia included in Astroloba. Bothalia
30: 53.
Meyer NL, Smith GF. 1998. Astroloba corrugata: description of a long-known species
in a southern African endemic Alooid genus. Bothalia 28: 60—62.
Meyer NL, Smith GF. 2001. Aloaceae. In Eggli, U. (Ed.), Illustrated Handbook of
succulent plants: Monocotyledons. Springer, Berlin.
Mössmer M, Smith GF, van Wyk BE, Condy G. 1995. Haworthia fasciata fa. fasciata:
Flowering Plants of Africa 54: 88—93.
Obermeyer AA. 1973. Liliaceae: Aloe, Chamaealoe, Haworthia, Astroloba, Poellnitzia
and Chortolirion. Bothalia 11: 119.
Palumbi SR. 1996. Nucleic acids II: the polymerase chain reaction, pp. 241-246. In
Hillis DM, Moritz C, Mable BK, eds. Molecular systematics, 2nd ed. Sinauer,
Inc. Publishers, Sunderland, MA.
Paradis E, Claude J, Strimmer K. 2004. APE: analyses of phylogenetics and evolution
in R language. Bioinformatics 20: 289—290.
Posada D, Crandall KA. 1998. MODELTEST: testing the model of DNA substitution.
Bioinformatics (application note) 14: 817—818.
R Development Core Team 2011. R: A language and environment for
statistical
computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3900051-07-0, URL http://www.R-project.org/.
Ramdhani S, Barker NP, Cowling RM. 2011. Revisiting monophyly in Haworthia
Duval
(Asphodelaceae):
incongruence,
hybridization
and
contemporary
speciation. Taxon 60: 1001-1014.
86
!
References
!
!
Ratnasingham S, Herbert PD. 2007. BOLD: the Barcode of Life Data System
(http://www.barcodinglife.org). Molecular Ecology Notes 7: 355—364.
Reynolds GW. 1966. The aloes of tropical Africa and Madagascar. The Trustees: The
Aloes Book Fund, Mbabane, Swaziland.
Reynolds GW. 1969. The aloes of South Africa. A. A. Balkema, Cape Town, South
Africa.
Roberts R. 1965. The genus Astroloba Uitewaal (Liliaceae). M.Sc. thesis, University of
Cape Town, South Africa.
Ronquist F, Huelsenbeck JP. 2003. MRBAYES 3: Bayesian phylogenetic inference
under mixed models. Bioinformatics 19: 1572—1574.
Sang T, Crawford DJ, Steussy TF. 1997. Chloroplast DNA phylogeny, reticulate
evolution and biogeography of Paeonia (Paeoniaceae). American Journal of
Botany 84: 1120—1136.
Scott CL. 1985. The genus Haworthia (Liliaceae): a taxonomic revision. Aloe Books,
Johannesburg, South Africa.
Seelanan T, Schnabel A, Wendel JF. 1997. Congruence and consensus in the cotton
tribe (Malvaceae). Systematic Botany 22: 259—290.
Small RL, Cronn RC, Wendel JF. 2004. Use of nuclear genes for phylogeny
reconstruction in plants. Australian Systematic Botany 17: 145—170.
Smith GF, Steyn EMA, Steyn C, van Wyk AE. 1996. Notes on the leaf anatomy of
Chortolirion and Haworthia (Aloaceae). South African Journal of Botany 62:
217—219.
Smith GF, Steyn EMA. 2004. Taxonomy of Aloaceae. In: Reynolds, T. (Ed.), Aloes:
The genus Aloe. CRC Press, London, UK, pp. 15—36.
87
!
References
!
!
Smith GF, Steyn EMA. 2005. Notes on the phenology, natural geographical distribution
range and taxonomy of Aloe dichotoma (Aloaceae). Bradleya 23: 17—22.
Smith GF, van Wyk AE. 1993. The generic status of Chortolirion (Aloaceae): Evidence
from leaf anatomy. Kew Bulletin 48: 105—113.
Smith GF, van Wyk BE, Condy G. 1995. Poellnitzia rubriflora. Flowering Plants of
Africa 54: 94—98.
Smith GF, van Wyk BE, Steyn EMA, Breuer I. 2001. Infrageneric classification of
Haworthia (Aloaceae): perspectives from nectar sugar analysis. Systematics and
Geography of Plants 71: 391—397.
Smith GF, van Wyk BE. 1991. Generic relationships in the Alooideae (Asphodelaceae).
Taxon 40: 557—581.
Smith GF, van Wyk BE. 1998. Asphodelaceae. In: Kubitzki K, ed. The Families and
genera of vascular plants, III. Flowering Plants. Monocotyledons, Lilianae
(except Orchidaceae), Springer-Verlag, New York, pp. 130-140.
Smith GF. 1995a. FSA contributions 2: Asphodelaceae/Aloaceae, 1029010 Chortolirion.
Bothalia 25: 31—33.
Smith GF. 1995b. FSA contributions 3: Asphodelaceae/Aloaceae, 1028010 Poellnitzia.
Bothalia 25: 35—36.
Swofford DL. 2002. PAUP*: phylogenetic analysis using parsimony (* and other
methods), version 4.0b10. Massachusetts: Sinauer Associates.
Taylor WR. 1925. Cytological studies on Gasteria II. A comparison of the chromosomes
of Gasteria, Aloe and Haworthia. American Journal of Botany 12: 219—223.
The Plant List 2010. Version 1. Published on the Internet; http://www.theplantlist.org/
(accessed 1st January).
Treutlein J, Smith GF, van Wyk BE, Wink M. 2003a. Evidence for the polyphyly of
88
!
References
!
!
Haworthia (Asphodelaceae subfamily Alooideae; Asparagales) inferred from
nucleotide sequences of rbcL, matK, ITS1 and genome fingerprinting with
ISSR-PCR. Plant Biology 5: 513—521.
Treutlein J, Smith GF, van Wyk BE, Wink M. 2003b. Phylogenetic relationships in
Asphodelaceae (subfamily Alooideae) inferred from chloroplast DNA sequences
(rbcL, matK) and from genomic finger- printing (ISSR). Taxon 52: 193—207.
Van Jaarsveld EJ, Low B, van Wyk AE. 2004. Aloe kouebokkeveldensis: a new species
from the Western Cape. Aloe 41: 36—37.
Van Jaarsveld EJ, van Wyk AE. 2004. Gasteria doreeniae, a new species from the
Eastern Cape. Aloe 41: 81—83.
Van Jaarsveld EJ, van Wyk AE. 2005. Gasteria tukhelensis, a new species from
Kwazulu-natal, South Africa. Bothalia 35: 164—166.
Van Jaarsveld EJ, van Wyk AE. 2006. Aloe challisii, a new cliff-dwelling aloe from
Mpumalanga, and a checklist of the obligate cliff-dwelling aloes in South Africa
and Namibia. Aloe 43: 36—41.
Van Jaarsveld EJ. 1992. The genus Gasteria: A Synoptic Review. Aloe 29: 5—33.
Van Jaarsveld EJ. 1994. Gasterias of South Africa. A new revision of a major succulent
group. Cape Town: Fernwood Press in association with the National Botanical
Institute, South Africa.
Van Jaarsveld EJ. 1998. A new taxon and new combinations in the Gasteria carinata
complex. Cactus and Succulent Journal (US) 70: 65—71.
Van Jaarsveld EJ. 2001. Gasteria polita van Jaarsv., a new species from the Western
Cape. Cactus and Succulent Journal (US) 73: 127—130.
Van Jaarsveld EJ. 2007. The genus Gasteria: A Synoptic Review (new taxa and
combinations). Aloe 44: 84—104.
89
!
References
!
!
Van Wyk BE, Smith GF. 2003. Guide to Aloes of South Africa. Briza Publications,
Pretoria, South Africa.
Van Wyk BE, Whitehead CS, Glen HF, Hardy DS, Van Jaarsveld EJ, Smith GF.
1993. Nectar Sugar Composition in the Subfamily Alooideae (Asphodelaceae).
Biochemical Systematics and Ecology 21: 249—253.
Victor JE, Dold AP. 2003. Threatened plants of the Albany centre of Floristic
Endemism, South Africa. South African Journal of Science 99: 437—446.
Viljoen AM, Van Wyk BE, Van Heerden FR. 1998. Distribution and chemotaxonomic
significance of flavonoids in Aloe (Asphodelaceae). Plant systematics and
evolution 211:31—42.
Viljoen AM. 1999. A chemotaxonomic study of phenolic leaf compounds in the genus
Aloe. Ph.D. thesis, Rand Afrikaans University, Johannesburg.
Vosa CG, Bennett ST. 1990. Chromosome studies in the southern African flora. 58-94.
Chromosome evolution in the genus Gasteria Duval. Caryologia 43: 235—247.
Vosa CG. 2004. On the classification of some species of the genus Haworthia, subgenus
Haworthia (Asphodelaceae). Caryologia 57: 395—399.
Wiens JJ. 1998. Testing phylogenetic methods with tree-congruence: Phylogenetic
analysis of polymorphic morphological characters in phrynosomatid lizards.
Systematic Biology 47:427—444.
Zonneveld BJM, van Jaarsveld EJ. 2005. Taxonomic implications of genome size for
all species of the genus Gasteria Duval (Aloaceae). Plant Systematics and
Evolution 251: 217—227.
90
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Appendices
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!
APPENDIX I
This appendix includes all abstracts of papers presented at conferences.
PAPER PRESENTATIONS
Molecular phylogeny of Haworthia (Asphodelaceae): Evidence from plastid and
nuclear DNA sequences
Barnabas H. Daru1, Olivier Maurin1, Norman Maclean2, Bruce M. Bayer3, and
Michelle van der Bank1
1
African Centre for DNA Barcoding, Faculty of Science,
University of Johannesburg, P. O. Box 524 Auckland Park, 2006 Johannesburg, South
Africa.
2
School of Biological Sciences, University of Southampton, Highfield, Southampton,
Hants,
SO16 7PX, United Kingdom.
3
PO Box 960, 7579 Kuilsriver, Western Cape, South Africa.
Haworthia Duval (Asparagales: Asphodelaceae: Alooideae) is endemic to southern
Africa with modifications to withstand relatively waterless or desert environments. There
is considerable variation in form between members of the genus and also, to a lesser
extent, between populations of the same species. It is likely that cryptic species are yet to
be described. Despite their importance in commercial horticulture, little is known about
their evolutionary history and their genetic diversity, and the taxonomy remains
unresolved. In the current study, generic limits of Haworthia were addressed using
molecular sequenced data from three plastid (rbcLa, matK and trnH-psbA) and the
nuclear ribosomal internal transcribed spacer (ITS1). Representatives of other genera
within Alooideae were also included to infer the placement of the three subgenera of
Haworthia within Alooideae. Preliminary findings were discussed.
Oral Presentation: 2011 Congress of the Southern African Society for Systematics
Biologists (SASSB), Rhodes University, Grahamstown, South Africa.
Awarded the Best MSc Oral presentation SASSB Congress 2011.
91
!
Appendices
!
!
!
Phylogeny of the subfamily Alooideae (Asphodelaceae): Paraphyly of Aloe and
Haworthia and consequences for classification
Barnabas H. Daru1,3,4, Olivier Maurin1, John C. Manning2, James S. Boatwright2,3,
and Michelle van der Bank1
1
African Centre for DNA Barcoding, University of Johannesburg, P. O. Box 524 Auckland
Park, 2006 Johannesburg, South Africa.
2
Compton Herbarium, South African National Biodiversity Institute, Private Bag X7,
Claremont 7735, South Africa.
3
Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box
524, Auckland Park 2006, Johannesburg, South Africa.
4
Author for correspondence (darunabas@gmail.com)
Abstract
This study addresses generic delimitation of subfamily Alooideae (Asphodelaceae) using
DNA sequences from plastid (rbcLa, matK and trnH-psbA) and nuclear (ITS1) regions
plus morphological trait data. The plastid and nuclear DNA were analysed using maximum
parsimony and bayesian statistics. The morphological data was mapped unto the molecular
phylogeny. Parsimony and bayesian analyses of cpDNA and ITS1 combined together
yielded two clades in Alooideae. The findings were discussed.
Keywords—Asphodelaceae, ITS1, matK, molecular phylogeny, morphology,
nomenclature, rbcLa, trnH-psbA, taxonomy
Oral Presentation: South African Academy for Science and Arts Conference, University
of Johannesburg, South Africa (2011)
92
!
Appendices
!
!
Phylogeny of the subfamily Alooideae (Asphodelaceae): Paraphyly of Aloe and
Haworthia and consequences for classification
Barnabas H. Daru1,3,4, John C. Manning2, James S. Boatwright2,3, Olivier Maurin1,
and Michelle van der Bank1
1
African Centre for DNA Barcoding, University of Johannesburg, P. O. Box 524 Auckland
Park, 2006 Johannesburg, South Africa.
2
Compton Herbarium, South African National Biodiversity Institute, Private Bag X7,
Claremont 7735, South Africa.
3
Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box
524, Auckland Park 2006, Johannesburg, South Africa.
4
Author for correspondence (darunabas@gmail.com)
Abstract
Alooideae (Asparagales–Asphodelaceae) currently comprises five genera, four of which
are endemic to southern Africa. Despite their importance in commercial horticulture,
evolutionary relationships among the genera are still incompletely understood, and the
generic taxonomy remains unresolved. This study addresses generic delimitation in
subfamily Alooideae using an expanded molecular sequenced datasets from three plastid
regions (rbcLa, matK and trnH-psbA) and the first subunit of the nuclear ribosomal
internal transcribed spacer (ITS1). Sequence data were analysed using maximum
parsimony and bayesian statistics, and selected morphological traits were mapped onto the
molecular phylogeny. Among the polytypic genera, only Astroloba and Gasteria were
retrieved as monophyletic. Haworthia is shown to be polyphyletic and to comprise three
main clades largely correllated with current subgeneric circumscriptions. The taxonomic
implications of these findings are examined and a revised, phylogeny-driven generic
classification is proposed in which the genus Aloe is expanded to include all members of
Alooideae.
Keywords—Asphodelaceae, ITS1, matK, molecular phylogeny, morphology,
nomenclature, rbcLa, trnH-psbA, taxonomy
Oral Presentation: South African Association of Botanists (SAAB) 38th Annual
Conference, University of Pretoria, South Africa (2012).
93
!
Appendices
!
!
APPENDIX II
Paper submitted to Taxon – 15 December 2011 (under review).
Phylogeny of the subfamily Alooideae (Asphodelaceae): Paraphyly of Aloe and
Haworthia and consequences for classification
Barnabas H. Daru1,7, John C. Manning2,3, James S. Boatwright2,4, Olivier Maurin1,
Bruce M. Bayer4, Norman Maclean5, Maria Kuzmina6 and Michelle van der Bank1
1
African Centre for DNA Barcoding, University of Johannesburg, P. O. Box 524 Auckland Park, 2006,
Johannesburg, South Africa.
2
Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont, 7735,
South Africa.
3
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.
4
Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box 524, Auckland
Park 2006, Johannesburg, South Africa.
5
Sheilam Nursery, PO Box 960, Kuilsriver, 7579, Western Cape, South Africa.
5
School of Biological Sciences, University of Southampton, Highfield, Southampton, Hants, SO16 7PX,
United Kingdom.
6
International Barcode of Life Project, Biodiversity Institute of Ontario, University of Guelph, Guelph, ON
N1G 2W1, Canada.
7
Author for correspondence (darunabas@gmail.com)
Abstract
Alooideae (Asparagales–Asphodelaceae) currently comprises five genera, four of which
are endemic to southern Africa. Despite their importance in commercial horticulture,
evolutionary relationships among the genera are still incompletely understood, and the
generic taxonomy remains unresolved. This study addresses generic delimitation in
subfamily Alooideae using an expanded molecular sequenced datasets from three plastid
regions (rbcLa, matK and trnH-psbA) and the first subunit of the nuclear ribosomal
internal transcribed spacer (ITS1). Sequence data were analysed using maximum
parsimony and bayesian statistics, and selected morphological traits were mapped onto
the molecular phylogeny. Among the polytypic genera, only Astroloba and Gasteria were
retrieved as monophyletic. Haworthia is shown to be polyphyletic and to comprise three
main clades largely correllated with current subgeneric circumscriptions. The taxonomic
implications of these findings are examined and a revised, phylogeny-driven generic
classification is proposed in which the genus Aloe is expanded to include all members of
Alooideae.
Keywords Asphodelaceae, ITS1, matK, molecular phylogeny, morphology,
nomenclature, rbcLa, trnH-psbA, taxonomy.
!
94
!
Appendices
!
!
APPENDIX III
The taxonomic changes in Alooideae resulting from the current study.
TAXONOMY
Aloe L., Sp. Pl.: 319. 1753. Type species: A. perfoliata L.
I. Section Aloidendron A.Berger in Bot. Jahrb. 36: 48. 1905. Type: Aloe barberae T.Dyer
II. Section Dracoaloe A.Berger in Bot. Jahrb. 36: 48. 1905. Type: Aloe dichotoma
Masson
III. Section Kumara (Medik.) Baker in J. Linn. Soc. Bot. 18: 155. 1880. Type: Aloe
plicatilis (L.) Mill.
IV.
Section
Reticulatae
Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 10. 1836–1863. Lectotype species, here designated: Aloe
reticulata Haw.
=Haworthia Duval, Pl. Succ. Horto Alencon.: 7. 1809. nom. cons., syn. nov. Type:
Haworthia arachnoidea (L.) Duval.
95
!
Appendices
!
!
=Apicra Willd., Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde
Berlin 5: 167. 1811. nom. illegit. pro Haworthia Duval – Type: not designated.
= Aloe section Retusae Salm-Dyck in Monographia generum Aloes et Mesembryanthemi
1: sect. 9. 1836–1863., syn. nov. – Lectotype species, here designated: Aloe retusa L.
=
Aloe
section
Limpidae
Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 11. 1836–1863., syn. nov. – Lectotype species, here
designated: Aloe cymbifolia Schad. [=Aloe cymbiformis Haw.]. [note: as there is no
nominate species in this section, the first listed species is chosen as lectotype].
= Aloe section Setatae Salm-Dyck in Monographia generum Aloes et Mesembryanthemi
1: sect. 12. 1836–1863., syn. nov. – Lectotype species, here designated: Aloe setosa
Schult. & Schult.f. [=Aloe arachnoidea (L.) Thunb.].
= Aloe section Loratae Salm-Dyck in Monographia generum Aloes et Mesembryanthemi
1: sect. 13. 1836–1863., syn. nov. – Lectotype species, here designated: Aloe
chloracantha (Haw.) Schult. & Schult.f.
1. Aloe arachnoidea [as arachnoides] (L.) Thunb., Fl. Cap.: 311. 1823. ≡ Aloe pumila
var. arachnoidea L., Sp.Pl.: 322. 1753. ≡ Haworthia arachnoidea (L.) Duval., Pl.Succ.
Horto Alencon.: 7. 1809. – Type: Illustration in Commelin Praeludea Bot.: t. 27. 1703.
(lecto, designated by Scott, 1977); South Africa, [Western Cape], Buitenstekloof,
Langvlei, Bayer 153 (NBG, epi., designated by Breuer & Metzing, 1997).
2. Aloe bayeri (J.D.Venter & S.A.Hammer) Boatwr. & J.C.Manning, comb. nov. ≡
Haworthia bayeri J.D.Venter & S.A.Hammer in Cact. Succ. J. (US) 69: 75. 1997. –Type:
South Africa, [Eastern Cape], hills S of Uniondale, Steyner s.n. (NBG, holo.).
96
!
Appendices
!
!
3. Aloe blackburniae (W.F.Barker) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
blackburniae W.F.Barker, J. S. Afr. Bot 3: 93. 1937. – Type: South Africa, [Western
Cape], Calitzdorp, Reynolds 1842 (NBG, holo.; BOL, PRE, iso.).
4. Aloe bolusii (Baker) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia bolusii Baker,
J. Linn. Soc. 18: 215. 1880. – Type: South Africa, [Eastern Cape], Graaff-Reinet, Bolus
158 (K, holo.; BOL, iso.).
5. Aloe chloracantha (Haw.) Schult. & Schult. f., Syst. Veg. 7(1): 641. 1829. ≡
Haworthia chloracantha Haw., Saxifrag. Enum. 2: 57. 1821. – Type: South Africa,
[Western Cape], N of Herbertsdale, Bayer s.n. (NBG, neo., designated by Breuer &
Metzing, 1997).
6. Aloe cymbiformis Haw., Trans. Linn. Soc. 7: 8. 1804. ≡ Haworthia cymbiformis
(Haw.) Duval, Pl. Succ. Horto Alencon.: 7. 1809. – Type: South Africa, [Eastern Cape],
Walmer, Smith 2844 (NBG, neo., designated by Breuer & Metzing, 1997).
7. Aloe denticulata (Haw.) Schult. & Schult. f., Syst. Veg. 7(1): 639. 1829. ≡ Haworthia
denticulata Haw., Rev. Pl. Succ.: 58. 1821. – Type: Illustration in K (lecto., designated
by Bayer, 1999).
=Haworthia aristata Haw., Suppl. Pl. Succ.: 51. 1819., non Aloe aristata Haw. 1825. –
Type: Illustration in K (lecto., designated by Bayer, 1999); South Africa, [Eastern Cape],
Dead Man’s Gulch, Smith 3550 (NBG, epi., designated by Bayer, 1999).
8. Aloe emelyae (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia emelyae
Poell., Repert. Spec. Nov. Regni Veg. 42: 271. 1937. – Type: Unpublished photograph
“Haworthia Emelyae v. P.” (B, lecto., designated by Breuer & Metzing, 1997).
97
!
Appendices
!
!
9. Aloe floribunda (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
floribunda Poelln., Repert. Spec. Nov. Regni Veg. 40:149. 1936. – Type: Unpublished
photograph “Haworthia floribunda v. P.” (B, neo., designated by Bayer, 1982); South
Africa, [Western Cape], Black Down, 3.2 km N of Heidelberg, Bayer 158 (NBG, epi.,
designated by Breuer & Metzing, 1997).
10. Aloe heidelbergensis (G.G.Sm.) Boatwr. & J.C.Manning, comb. nov. ≡ Hawortia
heidelbergensis G.G.Sm. in J. S. Afr. Bot. 14: 42. 1948. – Type: South Africa, [Western
Cape], W of Heidelberg, Smith 6566 (NBG, holo.; NBG, iso.).
11. Aloe herbacea Mill., Gard. Dict. ed. 8: Aloe No. 18. 1768. ≡ Haworthia herbacea
(Mill.) Stearn in Cact. J. (Croydon) 7: 40. 1938. – Type: Illustration in Boerhaave, Ind.
Alter Hort. Lugd.-Bat. 2: ad p. 131. 1720 (lecto, designated by Bayer, 1972b); South
Africa, [Western Cape], N of Ribbokkop, SW of Worcester, Bayer 161 (NBG, epi.,
designated by Breuer & Metzing, 1997).
12. Aloe lockwoodii (Archibald) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
lockwoodii Archibald in Fl. Pl. S. Afr.: ad t. 792. 1940. – Type: South Africa, [Western
Cape], Laingsburg, Lockwood Hill 215 (GRA, holo.).
13. Aloe magnifica (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
magnifica Poelln., Repert. Spec. Nov. Regni Veg. 33:240. 1933. – Type: South Africa,
[Western Cape], Riversdale, Ferguson s.n. (BOL, lecto., designated by Breuer &
Metzing, 1997).
14. Aloe maraisii (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia maraisii
Poelln, Repert. Spec. Nov. Regni Veg 38: 194. (1935). – Type: Unpublished photograph
“Haworthia Maraisii v. P.” (B, lecto., designated by Bayer, 1999).
98
!
Appendices
!
!
15. Aloe marumiana (Uitewaal) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
marumiana Uitewaal in Cact. & Vetpl. 6:33. 1940. – Type: South Africa, [Western
Cape], Ladismith ex hort. Stellenbosch sub No. 6610 (AMD, holo.).
16. Aloe mirabilis Haw. in Trans. Linn. Soc. London 7: 9. 1804. ≡ Haworthia mirabilis
(Haw.) Haw., Syn. Pl. Succ.: 95. 1812. – Type: Curtis’s Bot. Mag.: t. 1354. 1811. (epi.,
designated by Bayer, 1977); South Africa, [Western Cape], Skuitsberg, between Caledon
and Greyton, Bayer 2453 (NBG, epi., designated by Breuer & Metzing, 1997).
17. Aloe monticola (Fourc.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia monticola
Fourc., Trans. Roy. Soc. S. Afr. 21: 78. 1937. – Type: South Africa, George and
Uniondale Districts, Fourcade 2498 (K, holo.).
18. Aloe mucronata (Haw.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
mucronata Haw., Suppl. Pl. Succ.: 50. 1819. – Type: Unpublished illustration
“Haworthia mucronata Haw.” (k, lecto., designated by Bayer 1999).
19. Aloe mutica (Haw.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia mutica Haw.,
Saxifrag. Enum 2: 55. 1821. – type: Illustration in Excelsa 8: 50. 1978. (lecto., designated
by Bayer 1978); South Africa, [Western Cape], Bredasdorp to Swellendam, NE of
Soesrivier bridge, Bayer s.n. (NBG, epi., designated by Breuer & Mitzing, 1997).
20. Aloe nortieri (G.G.Sm.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia nortieri
G.G.Sm. in J. S. Afr. Bot. 12: 13. 1946. – Type: South Africa, [Western Cape], Doorn
River between Clanwilliam and Vanrhynsdorp, Smith 6115 (NBG, holo.; NBG, iso.).
21. Aloe outeniquensis (M.B.Bayer) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
outeniquensis M.B.Bayer, Haworthia Revisited: 130. 1999. – Type: South Africa,
[Western Cape], Moerasrivier, Venter, Marx & Kent 94/61 (NBG, holo.).
99
!
Appendices
!
!
22. Aloe parksiana (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
parksiana Poelln., Repert. Spec. Nov. Regni Veg. 41: 205. 1937. – Type: Illustration
published in Desert Pl. Life 10: 48. 1938 “Haw. Parksiana v. P.” (lecto., designated by
Breuer & Metzing, 1997).
23. Aloe pilifera (Baker) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia pilifera
Baker, Saund. Ref. Bot. 4: t. 234. 1871. – Type: Illustration in Saund. Ref. Bot. 4: t. 234.
1871. (lecto., designated by Bayer, 1999).
=Haworthia cooperi Baker, Saund. Ref. Bot. 4: t. 233. 1871., non Aloe cooperi Baker
(1874) – Type: South Africa, [Eastern Cape], without precise locality, Cooper s.n. (K,
holo.).
24. Aloe pringlei (C.L.Scott) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia pringlei
C.L.Scott in Bradleya 12: 103. 1994. – Type: South Africa, [Eastern Cape], Adelaide
Distr., Scott s.n. (PRE, holo.).
=Haworthia decipiens Poelln, Repert. Spec. Nov. Regni Veg. 28: 103. 1930., non Aloe
decipiens Schult. & Schult. f. (1829) [=Aloe nitida Salm-Dyck] – Type: South Africa,
[Eastern Cape], Zuurberg at Georgida, Fourcade 4367 (BOL, neo., designated by Breuer
& Metzing, 1997).
25. Aloe pubescens (M.B.Bayer) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
pubescens M.B.Bayer in J.S. Afr. Bot. 38: 125. 1972. – Type: South Africa, [Western
Cape], Sandberg Hills, 12 km SSE of Worcester, Bayer 161 (NBG, holo.).
26. Aloe pulchella (M.B.Bayer) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
pulchella M.B.Bayer in J. S. Afr. Bot. 39: 232. 1973. – Type: South Africa, [Western
Cape], Avondrust, SE of Touwsrivier, Bayer 162 (NBG, holo.).
100
!
Appendices
!
!
27. Aloe pygmaea (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia pygmaea
Poelln., Repert. Spec. Nov. Regni Veg. 27: 132. 1929. – Type: South Africa, [Western
Cape], hills E of Great Brak River, Fourcade 4759 (BOL, neo., designated by Breuer &
Metzing, 1997).
28. Aloe reticulata Haw. in Trans. Linn. Soc. London 7: 9. 1804. ≡ Haworthia reticulata
(Haw.) Haw., Syn. Pl. Succ.: 94. 1812. – Type: Illustration in Curtis’s Bot. Mag.: t. 1314.
1810. (neo., designated by Bayer, 1972a); South Africa, [Western Cape], Ribbokkop,
19.2 km SW of Worcester, Bayer 160 (NBG, holo.).
29. Aloe retusa L., Sp. Pl.: 322. 1753. ≡ Haworthia retusa (L.) Duval., Pl. Succ.
Alencon.: 7. 1809. – Type: Illustration in Commelin, Horti Med. Amstelad. 2: t. 6. 1701.
(lecto., designated by Scott, 1985); South Africa, [Western Cape], Blikbonnie, 2 km E of
Riversdale, Dekenah s.n. (NBG, epi., designated by Breuer & Metzing, 1997).
30. Aloe semiviva (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia bolusii
var. semiviva Poelln. in Repert Spec. Nov. Regni Veg. 44: 40. 1928. ≡ Haworthia
semiviva (Poelln.) M.B.Bayer, Haworthia Handbook: 153. 1976. – Type: Illustration in
Succulenta (Netherlands) 22: 25. 1940. “Haw. Bolusii var. semiviva v. P. Typ.” (lecto.,
designated by Breuer & Metzing, 1997).
31. Aloe serrata (M.B.Bayer) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia serrata
M.B. Bayer in J. S. Afr. Bot. 39: 249. 1973. – Type: South Africa, [Western Cape],
Oudekraalkop, 40 km SW of Heidelbereg, Bayer 166 (NBG, holo.).
32. Aloe springbokvlaktensis (C.L.Scott) Boatwr. & J.C.Manning, comb. nov. ≡
Haworthia springbokvlaktensis [as springbokvlakensis] C.L.Scott in J. S. Afr. Bot. 36:
101
!
Appendices
!
!
287. 1970. – Type: South Africa, [Eastern Cape], Steytlerville, Farm Springbokvlakte,
Scott 245 (PRE, holo.).
33. Aloe stenophylla Schult. & Schult. f., Syst. Veg. 7(1): 641. 1829. – Type: Illustration
in Salm-Dyck Monographia generum Aloes et Mesembryanthemi 1 sect. 13: fig. 2. 1836–
1863. (neo, designated by Bayer, 1999); Epitype: South Africa, [Eastern Cape],
Highlands, Cooke s.n. (NBG, epi, designated by Bayer, 1999).
=Haworthia angustifolia Haw. in Philos. Mag. J. 66: 283. 1825. non Aloe angustifolia
Haw. 1819. [= Aloe africana Mill.] – Type: South Africa, [Eastern Cape], 34 km from
Grahamstown to Alicedale, Bruyns 1653 (NBG, neo, designated by Breuer & Metzing,
1997).
34. Aloe translucens Haw. in Trans. Linn. Soc. London: 10. 1804. ≡ Haworthia
translucens (Haw.) Haw., Suppl. Pl. Succ.: 52. 1819. – Type: South Africa, [Eastern
Cape], 2 km E of Hankey, Bayer 4476 (NBG, neo, designated by Breuer & Metzing,
1997).
=Haworthia gracilis Poelln., Feddes. Repert. Spec. Nov. 27: 133. 1929. – Type:
Illustration in B.; South Africa, [Eastern Cape], Helspoort, Britten s.n. (PRE, epi.,
designated by Bayer, 1999).
35. Aloe truncata (Schönland) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
truncata Schönland in Trans. Roy. Soc. S. Afr. 1: 391. 1910. – Type: South Africa,
[Western Cape], a farm 11.2 km from Oudtshoorn, Britten s.n. (K., holo.).
36. Aloe turgida (Haw.) Schult. & Schult. f., Syst. Veg. 7(1): 635. 1829. ≡ Haworthia
turgida Haw., Suppl. Pl. Succ.: 52. 1819. – Type: South Africa, [Western Cape],
102
!
Appendices
!
!
Swellendam, Breede River Bridge, Bayer 2420 (NBG, neo., designated by Breuer &
Metzing, 1997).
37. Aloe punctulata Boatwr. & J.C.Manning, nom. nov., pro Haworthia variegata
L.Bolus in J. Bot. Soc. S. Afr.: 137. 1929., non Aloe variegata L. (1753) – Type: South
Africa, [Western Cape], Still Bay, Botterkloof, Ferguson s.n. (BOL, holo.).
38. Aloe venteri Boatwr. & J.C.Manning nom. nov. pro. Haworthia maculata (Poelln.)
M.B.Bayer in Haworthia Handbook: 130. 1976. ≡ Haworthia schuldtiana var. maculata
Poelln., in Repert. Spec. Bot. Regni Veg. 49: 25. 1940., non Aloe maculata All. 1773. –
Type: South Africa, [Western Cape], Worcester, Venter 6 (BOL, lecto., designated by
Breuer & Metzing, 1997)
= Haworthia intermedia Poelln., Kakteenkunde 9: 133. 1937., non. Aloe intermedia Haw.
1804. [=Aloe carinata Mill.]. – Type: Unpublished photograph, “Haworthia intermedia
v. P.” (B, lecto., designated by Bayer, 1999); South Africa, [Western Cape],
Buitenstekloof, Bayer 4461 (NBG, epi., designated by Bayer, 1999).
39. Aloe vlokii (M.B.Bayer) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia vlokii
M.B.Bayer in Haworthia Revisited: 160. 1999. – Type: South Africa, [Western Cape],
Swartberg Mountains, Vlok sub Venter 91/2 (NBG, holo.).
40. Aloe wittebergensis (W.F.Barker) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
wittebergensis W.F.Barker in J. S. Afr. Bot. 8: 245. 1942. – Type: South Africa, [Western
Cape], Witteberg, Pieterse s.n. (NBG, holo.).
41. Aloe zantneriana (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia
zantneriana Poelln. in Repert. Spec. Nov. Regni Veg. 41: 217. 1937. – Type: Illustration
103
!
Appendices
!
!
in Desert Pl. Life 9: 90. 1937. “Haworthia Zantneriana v. P.” (lecto., designated by
Breuer & Metzing, 1997).
V. Section Macrifoliae (Haw.) Glen & D.S.Hardy, FSA 5 (1, 1): 92. 2000. Type: Aloe
ciliaris Haw.
VI. Section Aristatae (A.Berger) Glen & D.S.Hardy, FSA 5 (1, 1): 31. 2000. Type: Aloe
aristata Haw.
VII. Section Poellnitzia (Uitewaal) Boatwr. & J.C.Manning, comb. et stat. nov. ≡
Poellnitzia Uitewaal in Succulenta 22: 61. 1940. – Type: Apicra rubriflora L.Bolus
[=Poellnitzia rubriflora (L.Bolus) Uitewaal].
1. Aloe rubriflora (L.Bolus) G.D.Rowley in Cactus & Succulent Journal of Great Britain
43: 2. 1981. ≡ Apicra rubriflora L. Bolus in Ann. Bolus Herb. 3: 13. 1920. – Type: South
Africa [Western Cape], Bonnie Vale [Bonnievale], Smith s.n. BOL45213 (BOL, holo.).
VIII.
Section
Imbricatae
Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 1. 1836–1863. Lectotype species, designated here: Aloe
imbricata Haw. [=Aloe spiralis L.].
=Section Foliolosae Salm-Dyck in Monographia generum Aloes et Mesembryanthemi 1:
sect. 2. 1836–1863, syn. nov. Lectotype species, here designated: Aloe foliolosa Haw.
104
!
Appendices
!
!
=Astroloba Uitewaal in Succulenta 5: 53. 1947, syn. nov. Type: Astroloba pentagona
(Ait.) Uitewaal [=Aloe spiralis L.].
1. Aloe bullulata Jacq., Fragm. Bot. 72. t. 109. ≡ Astroloba bullulata (Jacq.) Uitewaal in
Succulenta 5: 53. 1947. – Type: Illustration in Fragm. Bot. 72. t. 109.
2. Aloe congesta Salm-Dyck in Monographia generum Aloes et Mesembryanthemi 1:
sect. 2, fig. 1. 1836–1863. ≡ Astroloba congesta (Salm-Dyck) Uitewaal in Succulenta 5:
54. 1947. – Type: Illustration in Monographia generum Aloes et Mesembryanthemi 1:
sect. 2, fig. 1. 1836–1863.
3. Aloe corrugata (N.L.Mey. & G.F.Sm.) Boatwr. & J.C.Manning, comb. nov. ≡
Astroloba corrugata N.L.Mey. & G.F.Sm. in Bothalia 28: 61. 1998. – Type: South
Africa, Western Cape, 7.5 km W of Warmwaterberg turnoff on Montagu-Ladismith road,
Van Jaarsveld 13913 (PRE, holo.).
4. Aloe foliolosa Haw. Trans. Linn. Soc. London 7: 7. 1804. ≡ Astroloba foliolosa (Haw.)
Uitewaal in Succulenta 5: 54. 1947. – Type: Illustration in Curtis’s Bot. Mag. t. 1352.
1811. [This illustration was drawn from Masson’s original introduction cultivated by
Haworth and is accepted as the type in the absence of a specimen.].
5. Aloe herrei (Uitewaal) Boatwr. & J.C.Manning, comb. nov. ≡ Astroloba herrei
Uitewaal in Desert Pl. Life 20. 37. 1948. – Type: Uniondale, Herre s.n. “Stellenbosch
5703” (?STEU, holo.).
6. Aloe spiralis L., Sp. Pl. 1: 322. 1753. ≡ Astroloba spiralis (L.) Uitewaal in Succulenta
5: 53. 1947. Type: “Aloe Afr. erecta rotundata etc. Comm.” in Dillenius, Hort. Eltham. 1:
16, t. 13, f. 14. 1732. (lecto., designated by Wijnands, 1983).
105
!
Appendices
!
!
IX.
Section
Margaritiferae
Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 6. 1836–1863. ≡ Haworthia section Margaritiferae (SalmDyck) Haw. in Handbuch der Succulenten Pflanzen: 710. 1954. Lectotype, here
designated: Aloe pumila L. [=Haworthia margaritifera (L.) Haw.]
≡Haworthia
subgenus
Robustipedunculatae
(Uitewaal)
M.B.Bayer
[as
Robustipedunculares Uitewaal ex M.B.Bayer] in Haworthia Handbook: 14. 1976.
Haworthia [unranked] Robustipedunculatae Uitewaal in Succulenta 51. 1947. Lectotype,
designated by Bayer in Haworthia Handbook: 14. 1976: Haworthia margaritifera (L.)
Haw. [= Aloe pumila L.].
=
Aloe
section
Albicantes
Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 5. 1836–1863., syn. nov. – Type: Aloe albicans Haw. [=Aloe
marginata Lam.] [Note: this section is monotypic].
1. Aloe granata (Willd.) Boatwr. & J.C.Manning, comb. nov. ≡ Apicra granata Willd. in
Berl. Mag.: 269. 1811. – Type: not seen.
=Haworthia minima (Ait.) Haw., Syn. Pl. Succ.: 92. 1812. ≡ Aloe margaritefera var.
minima Ait., Hort. Kew 1:468. 1789. – Type: Illustration in Dillenius, Hort. Eltham.: t.
16, f. 18. 1732 (lecto., designated by Scott, 1985).
2. Aloe kingiana (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia kingiana
Poelln. In Feddes Repert. 41: 203. 1937. – Type: South Africa, [Western Cape], Great
Brak, Dekenah 201 (NBG, neo., designated by Breuer & Metzing, 1997.).
106
!
Appendices
!
!
3. Aloe marginata Lam., encycl. 1: 89. 1783. ≡ Haworthia marginata (Lam.) Stearn in
Cact. J. (Croydon) 7: 34. 1938. – Type: Illustration in Commelin, Praeludia Bot.: t. 30.
1703 (lecto., designated by Scott, 1985).
4. Aloe pumila L., Sp. Pl.: 322. 1753. ≡ Aloe pumila var. margaritifera L., Sp. Pl.: 322.
1753. ≡ Aloe margaritifera (L.) Burm. f., Prodr. Fl. Cap.: 10. 1768. ≡ Haworthia
margaritifera (L.) Haw., Suppl. Pl. Succ.: 55. 1819. ≡ Haworthia pumila (L.) M.B.Bayer,
Haworthia revisited: 214. 1999.– Type: Illustration in Commelin, Hort Med. Amstelad.
Pl. Rar. 2: 19, t. 10. 1701 (lecto., designated by Scott, 1978). [Although Breuer and
Metzing (1997) treated the species under the name H. margaritifera, Scott’s (1978)
lectotypification renders this name homotypic with Aloe pumila L., which as the autonym
has statutory priority (Jarvis, 2007).].
X. Section Parviflorae Haw. in Trans. Linn. Soc. London 7: 6. 1804. ≡ Aloe subgenus
Parviflorae (Haw.) Salm-Dyck in Monographia generum Aloes et Mesembryanthemi 1.
1836–1863. Lectotype species, here designated: Aloe viscosa L. [the first species listed
by Haworth in his A. section Parviflorae is here chosen as lectotype]
≡Haworthia section Hexangulares Uitewaal in Succulenta 5: 51. 1947, syn. nov. ≡
Haworthia subgenus Hexangulares (Uitewaal) M.B.Bayer [as Uitewaal ex M.B.Bayer] in
Haworthia Handbook: 14. 1976. – Lectotype species, designated by Bayer (1976):
Haworthia coarctata Haw.
=
Aloe
section
Triquetrae
Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 3. 1836–1863., syn. nov. –Lectotype species, here designated:
107
!
Appendices
!
!
Aloe cordifolia (Haw.) Schult. & Schult.f. [=Aloe viscosa L.] [note: as there is no
nominate species in this section, the first listed species is chosen as lectotype].
=
Aloe
section
Tortuosae
Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 4. 1836–1863., syn. nov. – Lectotype species, here designated:
Aloe tortuosa Haw. [= Aloe viscosa L.]
= Aloe section Luridae Salm-Dyck in Monographia generum Aloes et Mesembryanthemi
1: sect. 7. 1836–1863., syn. nov. – Lectotype species, here designated: Aloe scabra
(Haw.) Schult. & Schult.f. [Note: As there is no nominate species in this section, the first
listed species is chosen as lectotype].
=
Aloe section Tessellatae Salm-Dyck
in
Monographia
generum
Aloes
et
Mesembryanthemi 1: sect. 8. 1836–1863., syn. nov. – Lectotype species, here designated:
Aloe tessellata (Haw.) Schult. & Schult.f. [=Aloe venosa Lam.].
1. Aloe attenuata Haw., Trans. Linn. Soc. 7: 11. 1804. ≡ Haworthia attenuata (Haw.)
Haw., Syn. Pl. Succ.: 92. 1812. – South Africa, [Eastern Cape], Sandland, 20 km E of
Paternsie, Perry 660 (NBG, neo., designated by Breuer & Metzing, 1997).
2. Aloe brucei Boatwr. & J.C.Manning, nom. nov. pro Haworthia pungens M.B.Bayer in
Haworthia Revisited: 188. 1999. non Aloe pungens A.Berger (1908). – Type: South
Africa, [Eastern Cape], Joubertina, Bruyns 7090 (BOL, holo.).
3. Aloe bruynsii (M.B.Bayer) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia bruynsii
M.B.Bayer in J.S. Afr. Bot. 47: 789. 1981. – Type: South Africa, [Eastern Cape], SE of
Steytlerville, Rossouw 456 (NBG, holo.).
108
!
Appendices
!
!
4. Aloe coarctata (Haw.) Schult. & Schult. f., Syst. Veg. ed 15, v. 7(1): 647. 1829. ≡
Haworthia coarctata Haw., Phil. Mag. 44: 301. 1824. – Type: South Africa, [Eastern
Cape], 16 km from Grahamstown to Bathurst, Smith 7092 (NBG, neo., designated by
Breuer & Metzing, 1997).
5. Aloe fasciata (Willd.) Salm-Dyck in Schult. & Schult. f., Syst. Veg. ed 15, v. 7(2):
1713. 1830. ≡ Apicra fasciata Willd. In Ges. Naturf. Freunde Berlin Mag. Neuesten
Entdeck. Gesammten Naturk. 5: 270. 1811. ≡ Haworthia fasciata (Willd.) Haw.,
Saxifrag. Enum. 2: 54. 1821. – Type: South Africa, [Eastern Cape], Hankey, Stayner s.n.
(NBG, neo., designated by Breuer & Metzing, 1997).
6. Aloe glabrata Salm-Dyck, Hort. Dyck.: 325. 1834. ≡ Haworthia glabrata (Salm-Dyck)
Baker, J. Linn. Soc., Bot. 18: 206. 1880. – Type: Illustration in Salm-Dyck, Monographia
generum Aloes et Mesembryanthemi 1 sect. 3: fig. 7 [sect. 6, fig. 13] (neo., designated by
Smith & Greyling, 1990).
7. Aloe glauca (Baker) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia glauca Baker,
J. Linn. Soc. Bot. 18: 203. 1880. –Type: South Africa, [Eastern Cape], Cape, without
precise locality, Cooper s.n. (K., holo.).
8. Aloe koelmaniorum (Oberm. & Hardy) Boatwr. & J.C.Manning, comb. nov. ≡
Haworthia koelmaniorum [as koelmaniora] Oberm. & Hardy, Fl. Pl. Afr.: t1502. 1967. –
Type: South Africa, [Mpumalanga], Groblersdal, Hardy & Mauve 2267 (PRE, holo.).
9. Aloe limifolia (Marloth) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia limifolia
Marloth, Trans. Roy. Soc. S. Afr. 1: 409. 1910. – Type: Mozambique, W of Delagoa Bay,
Marloth 4678 (PRE, holo.).
109
!
Appendices
!
!
10. Aloe longiana (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia longiana
Poelln. in Repert. Spec. Nov. Regni Veg. 41: 203. 1937. – Type: unpublished
photographic icon, “Haworthia Longiana v. P.” (B, lecto. designated by Breuer &
Metzing, 1997).
11. Aloe nigra (Haw.) Schult & Schult. f., Syst. Veg. ed 15, v. 7(1): 647. 1829. ≡ Apicra
nigra Haw. in Philos. Mag. J 64: 302. 1824. ≡ Haworthia nigra (Haw.) Baker in J. Linn.
Soc., Bot 18: 203. 1880. –Type: South Africa, [Eastern Cape], Campherspoort
[Kamferspoort], Barker 5099 (NBG, neo., designated by Breuer & Metzing, 1997).
12. Aloe reinwardtii Salm-Dyck in Observ. Bot. Horto Dyck: 37. 1821. ≡ Haworthia
reinwardtii (Salm-Dyck) Haw., Saxifrag. Enum. 2: 53. 1821. – Type: Illustration in
Salm-Dyck, Monographia generum Aloes et Mesembryanthemi 1 sect. 6: fig. 12. 1836–
1863. (neo, designated by Scott, 1981a); South Africa, [Eastern Cape], near top of hill
above Ncera River Bridge, Smith 3563 (NBG, epi., designated by Breuer & Metzing,
1997).
13. Aloe scabra (Haw.) Schult. & Schult. f., Syst. Veg. ed 15, v. 7(1): 644. 1829. ≡
Haworthia scabra Haw., Suppl. Pl. Succ.: 58. 1819. – Type: South Africa, without
precise locality, illustration in Cact. Succ. J. (Los Angeles) 52: 274. 1980. (lecto.,
designated by Scott, 1980).
14. Aloe sordida (Haw.) Schult. & Schult. f., Syst. Veg. ed 15, v. 7(1): 644. 1829. ≡
Haworthia sordida Haw. in Saxifrag. Enum. 2: 51. 1821. – Type: South Africa, without
locality, illustration in Salm-Dyck, Monographia generum Aloes et Mesembryanthemi 1:
sect. 7: fig. 1. 1836–1863. (neo, designated by Scott, 1985).
110
!
Appendices
!
!
15. Aloe venosa Lam., Encycl. 1: 89. 1873. ≡ Haworthia venosa (Lam.) Haw., Saxifrag.
Enum.: 51. 1821. – Type: South Africa, without locality, illustration in Commelin,
Praeludia Bot.: t. 29. 1703. (lecto., designated by Scott, 1978); South Africa, [Western
Cape], Swellendam, W of Breede River Bridge, Bayer 168 (NBG, epi., designated by
Breuer & Metzing, 1997).
16. Aloe viscosa L., Sp. Pl.: 322. 1753. ≡ Haworthia viscosa (L.) Haw., Syn. Pl. Succ.:
90. 1812. – Type: Illustration in Commelin, Praeludia Bot.: t. 31. 1703. (lecto.,
designated by Scott, 1981b); South Africa, [Western Cape], Calitzdorp, Blackburn
Valley, Barker 5073 (NBG, epi., designated by Breuer & Metzing, 1997).
XI. Section Curviflorae Haw. in Trans. Linn. Soc. Lond. 7: 12. 1804. – Lectotype
species, here designated: Aloe verrucosa Mill. [=Aloe carinata Mill.] [Note: the species is
selected as lectotype being the first listed in Haworth’s section].
=Gasteria Duval, Plantae succulentae in Horto Alenconio: 6. 1809. ≡ Aloe section
Gasteria [as Gasteriae] (Duval) Salm-Dyck, Monographia generum Aloes et
Mesembryanthemi 1: sect. 29. 1836–1863., syn. nov. – Lectotype, designated by Maire
in Flore de l’Afrique du nord 5: 71. 1958.: Gasteria angustifolia (Ait.) Duval. [= Aloe
disticha L. non Thunb.].
=Gasteria section Breviflorae Haw., Rev. Pl. Succ.: 47. 1821. nom. nud.
=Gasteria section Longiflorae Haw., Rev. Pl. Succ.: 47. 1821. nom. nud.
=Gasteria section Mediocres Haw., Rev. Pl. Succ.: 47. 1821. nom. nud.
111
!
Appendices
!
!
1. Series Bifariae Haw. in Trans. Linn. Soc. London 7: 12. 1804. Lectotype species, here
designated: Aloe verrucosa Mill. [=Aloe carinata Mill.] [note: the species is selected as
lectotype being the first listed in Haworth’s series and displaying the characteristics of
the series.].
1.1 Aloe acinacifolia J.Jacq., Ecloge pl. rar.: 49. 1811–1816. ≡ Gasteria acinacifolia
(J.Jacq.) Haw., Suppl. Pl. succ.: 49. 1819. – Type: South Africa, without precise locality,
illustration in J.Jacq., Ecloge pl. rar. t. 31. 1811–1816.
1.2. Aloe armstrongii (Schönland) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
armstrongii Schönland in Records of the Albany Museum 2: 258. 1912. – Type: South
Africa, without precise locality, Drège 566 (GRA, holo.).
1.3. Aloe batesiana (G.D.Rowley) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
batesiana G.D.Rowley in National Cactus & Succulent Journal 10: 32. 1955. – Type:
South Africa, without locality, Rowley s.n. (RNG, neo., designated by Van Jaarsveld,
1992).
1.4. Aloe carinata Mill., Gard. Dict. ed. 8. 1768. ≡ Gasteria carinata (Mill.) Duval in
Plantae succulentae in Horto Alenconio: 6. 1809. – Type: South Africa, without precise
locality, illustration in Commelin, Hort. Med. Amst. t. 9. 1701., (lecto. designated by
Wijnands, 1983).
1.5. Aloe croucheri Hook. f. in Curtis’s Bot. Mag. 95: t. 5812. 1869. ≡ Gasteria
croucheri (Hook. f.) Baker in J. Linn. Soc., Bot. 18: 196. 1880. – Type: South Africa,
without precise locality, Cooper s.n. (K, holo.).
112
!
Appendices
!
!
1.6. Aloe ellaphieae (Van Jaarsv.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
ellapieae Van Jaarsv. in Journal of the Cactus & Succulent Society (US) 63: 3. 1991. –
Type: South Africa [Eastern Cape], 20 km W of Patensie, Van Jaarsveld et al. 9904
(NBG, holo.).
1.7. Aloe fuscopunctata (Baker) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
fuscopunctata Baker in J. Linn. Soc. Bot. 18: 195. 1880. – Type: South Africa, “C. B
Spei”, Cooper s.n. (?K). [Note: selection of this name is based on the synonymy given in
Van Jaarsveld (2007). In an earlier treatment (Van Jaarsveld, 1994) this species was
treated under the synonymy of G. acinacifolia].
=Gasteria excelsa Baker in J. Linn. Soc. Bot. 18: 195. 1880. non Aloe excelsa A.Berger
(1906) – Type: South Africa [Eastern Cape], Chalumna River, Cooper s.n. (K, holo.).
1.8. Aloe nitida Salm-Dyck, Catalogue raisonne des especes d’Aloes: 13. 1817. ≡
Gasteria nitida (Salm-Dyck) Haw. in Philosophical Mag. 2: 359. 1827. –Type: South
Africa, without locality, illustration in Salm-Dyck, Monographia generum Aloes et
Mesembryanthemi 1 sect. 29: fig. 17. 1836–1863 (neo., designated by Van Jaarsveld,
1992).
1.9. Aloe polita (Van Jaarsv.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria polita Van
Jaarsv. in Cactus & Succulent Journal (US) 73: 127–130. 2001. – Type: South Africa
[Western Cape], Whisky Creek, Plettenberg Bay, Van Jaarsveld & Kok 13742 (NBG,
holo.).
1.10. Aloe pulchra (Ait.) Jacq., Pl. Rar. Hort. Schon.: t. 419. 1805 “1804”. ≡ Aloe
maculata var. pulchra Ait. in Hort. Kew.: 469. 1789. ≡ Gasteria pulchra (Ait.) Haw.,
Synopsis Plantarum Succulentarum: 86. 1812. – Type: South Africa, without precise
113
!
Appendices
!
!
locality, illustration in Miller, Fig. Pl.: t. 292. 1759 (lectotype designated by Van
Jaarsveld, 1992).
1.11. Aloe thunbergii (N.E.Br.) Boatwr. & J.C.Manning, comb.nov. ≡ Gasteria
thunbergii N.E.Br. in Bothalia 1: 140. 1923. – Type: South Africa, without precise
locality, Thunberg s.n. UPS-THUNB 8595 (UPS-THUNB, holo.).
1.12. Aloe tukhelensis (Van Jaarsv.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
tukhelensis Van Jaarsv. in Bothalia 35: 164. 2005. –Type: South Africa, KwaZulu-Natal,
Ngubevu, Van Jaarsveld et al. 17996 (NBG, holo.).
1.13. Aloe vanjaarsveldii Boatwr. & J.C.Manning, nom. nov. pro Gasteria glauca Van
Jaarsv. in Cactus & Succulent Journal (US) 70: 65. 1998, non Aloe glauca Mill. (1768) –
Type: South Africa [Eastern Cape], Kouga River, E of Guernakop, Van Jaarsveld &
Welsh 14670 (PRE, holo.).
1.14. Aloe vlokii (Van Jaarsv.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria vlokii
Van Jaarsv. in Cactus and Succulent Journal (US) 59: 170. 1987. –Type: South Africa
[Western Cape], Waboomsberg, Vlok 880 (NBG, holo.).
2. Series Pictae Haw. in Trans Linn. Soc. London 7: 14. 1804. Type: Aloe obliqua Haw.,
nom. illegit. [=Gasteria bicolor Haw.]
2.1. Aloe baylissiana (Rauh.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
baulissiana Rauh. in J. S.Afr. Bot. 43: 187. 1977. – Type: South Africa [Eastern Cape],
Farm Oudekraal, S of Somerset-East, Bayliss 2796 (HEID, holo.; PRE, iso.).
114
!
Appendices
!
!
2.2. Aloe bicolor (Haw.) Schult. & Schult. f., Syst. Veg. ed 15, v. 7: 682. 1829. ≡
Gasteria bicolor Haw. in Philosophical Mag. 57: 275. 1826. – Type: South Africa,
without locality, illustration in Salm-Dyck, Monographia generum Aloes &
Mesembryanthemi 1 sect. 29: fig. 5. 1836–1863 (neo., designated by Van Jaarsveld,
1992).
2.3. Aloe brachyphylla Salm-Dyck in Monographia generum Aloes & Mesembryanthemi
1 sect. 29: fig. 8. 1836–1863. ≡ Gasteria brachyphylla (Saml-Dyck) Van Jaarsv. in Aloe
29:19. 1992. – Type: South Africa, without locality, illustration in Saml-Dyck,
Monographia generum Aloes & Mesembryanthemi: t. 8. 1836–1863 (lecto., designated
by Van Jaarsveld, 1992).
2.4. Aloe disticha L. Sp. Pl.: 321. 1753. ≡ Gasteria disticha (L.) Haw. in Philosophical
Mag. 2: 352. 1927. –Type: South Africa, without precise locality, illustration in
Commelin, Hort. Med. Amst. t. 8. 1701 (lecto., designated by Wijnands, 1983).
2.5. Aloe doreeniae (Van Jaarsv. & A.E.van Wyk) Boatwr. & J.C.Manning, comb. nov.
≡ Gasteria doreeniae Van Jaarsv. & A.E.van Wyk in Aloe 41: 81. 2004. – Type: South
Africa, Eastern Cape, Swartwaterspoort, W of Riebeeck East, Quart 448 (NBG, holo.).
2.6. Aloe glomerata (Van Jaarsv.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
glomerata Van Jaarsv. in Bradleya 9: 100. 1991. – Type: South Africa, [Eastern Cape],
Kouga Dam, 25 km W of Patensie, Van Jaarsveld & Sardien 11054 (NBG, holo.).
2.7. Aloe neliana (Poelln.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria neliana
Poelln. in Feddes Rep. 27: 139. 1930. non Aloe pillansii L.Guthrie (1928)–Type: South
Africa, Clanwilliam, Poellnitz 559 (holo., supposedly in STE, specimen not found).
115
!
Appendices
!
!
=Gasteria pillansii Kensit in Trans. Roy. Soc. S. A. 1: 163. 1909. – Type: South Africa
[Western Cape], Clanwilliam, Pillans 833 (BOL, holo.; PRE, iso.).
2.8. Aloe rawlinsonii (Oberm.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria
rawlinsonii Oberm., Flow. Pl. Afr. 43: t. 1701. 1976. – Type: South Africa [Eastern
Cape], Willowmore Distr., Baviaanskloof, Rawlinson s.n. PRE34421 (PRE, holo.).
2.9. Aloe retusa (Van Jaarsv.) Boatwr. & J.C.Manning, comb. nov. ≡ Gasteria carinata
var. retusa Van Jaarsv. in Aloe 29: 15. 1992. ≡ Gasteria retusa (Van Jaarsv) Van Jaarsv.
in Aloe 44: 100. 2007. – Type: South Africa, [Western Cape], Orange Grove near De
Wet, Van Jaarsveld & Steyner 4656 (NBG, holo.).
XII. et seq. Remaining sections of Aloe s.s. (including Lomatophyllum, Chortolirion),
following Glen & Hardy (2000) and Reynolds (1966, 1969).
Section Chortolirion (Berger) Boatwr. & J.C.Manning, stat. et sect. nov. ≡ Chortolirion
Berger in Pflanxenr. 4, 38, 3, 2: 723. 1908., syn. nov. – Type: Chortolirion angolense
(Bak.) Berger [=Aloe tenuifolia (Engl.) Boatwr. & J.C.Manning]
1. Aloe tenuifolia (Engl.) Boatwr. & J.C.Manning, comb. nov. ≡ Haworthia tenuifolium
Engl. In Bot. Jahrb. 10: 2, t. 1. 1889. – Type: South Africa, [Northern Cape], near
Kuruman, Marloth 1049 (B, holo.; PRE, iso.).
=Haworthia angolensis Bak. in Trans. Linn. Soc. Bot. 1: 263. 1878. ≡ Chortolirion
angolense (Bak.) Berger Berger in Pflanxenr. 4, 38, 3, 2: 723. 1908., non Aloe angolensis
Bak. – Type: Angola, Huilla, Welwitsch 3756 (BM, holo.).
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