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Streptocarpus redefined to include all Afro-Malagasy Gesneriaceae:
Molecular phylogenies prove congruent with geographical
distribution and basic chromosome numbers and uncover remarkable
morphological homoplasies
Kanae Nishii,1,4 Mark Hughes,1 Marie Briggs,2 Elspeth Haston,1 Frieda Christie,1 Margaret J. DeVilliers,3
Thea Hanekom,3 Wiets G. Roos,3 Dirk U. Bellstedt3 & Michael Möller1
1 Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland, U.K.
2 Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, England, U.K.
3 Department of Biochemistry, University of Stellenbosch, Private Bag X1, Matieland 7602, South Africa
4 Tokyo Gakugei University, 4-1-1 Nukuikitamachi, Kogagei, Tokyo 184-8501, Japan
Author for correspondence: Michael Möller, m.moeller@rbge.ac.uk
ORCID: KN, http://orcid.org/0000-0001-8141-8908; MH, http://orcid.org/0000-0002-2168-0514; MB, http://orcid.org/0000-0003-29880032; EH, http://orcid.org/0000-0001-9144-2848; MM, http://orcid.org/0000-0002-2819-0323
DOI http://dx.doi.org/10.12705/646.8
Abstract The taxonomy of the African, Madagascan and Comoro Island (Afro-Malagasy) Gesneriaceae attracts a large amount
of interest given the horticultural importance of Cape Primroses (Streptocarpus) and African Violets (Saintpaulia). Earlier
studies indicated that the Afro-Malagasy genera form a strongly supported clade, and recent classifications have included some
of the genera within an expanded Streptocarpus. Given the global importance of this group, we carried out a comprehensive
molecular phylogenetic analysis of all Afro-Malagasy genera in subfamily Didymocarpoideae, tribe Trichosporeae, subtribe
Streptocarpinae, to investigate species relationships in these genera as the basis for a new classification. Phylogenetic analyses of the nuclear ribosomal spacer (ITS, 5S NTS) and chloroplast intron and spacer regions (rpl20-rps12 spacer, trnL intron,
trnLF spacer) of 226 samples were performed, including all Streptocarpinae genera, except the monotypic Nodonema. The
molecular phylogenies demonstrate that the genera with non-twisted fruits are nested within Streptocarpus which has twisted
fruits. Two main clades were found, one comprising herbaceous caulescent Streptocarpus that also included Saintpaulia, the
caulescents Hovanella and Schizoboea, and the unifoliates Acanthonema and Trachystigma. The second clade comprises the
woody caulescents and acaulescent Streptocarpus, Colpogyne and Linnaeopsis. Altogether, twelve well-supported subclades
can be recognized, each with a combination of distinct morphological characteristics. A new classification of tribe Streptocarpinae, de facto Streptocarpus, is presented, retaining the two subgenera, Streptocarpus and Streptocarpella, and dividing
them into five and seven sections respectively. Nodonema is attributed to subg. Streptocarpus for morphological reasons. The
former genus Saintpaulia is classified as Streptocarpus subg. Streptocarpella sect. Saintpaulia with ten species recognized.
Keywords Acanthonema; Colpogyne; Hovanella; infrageneric classification; Linnaeopsis; Nodonema; Saintpaulia;
Schizoboea; Trachystigma.
Supplementary Material The Electronic Supplement (Tables S1–S2; Figs. S1–S18) is available in the Supplementary Data
section of the online version of this article at http://www.ingentaconnect.com/content/iapt/tax; matrices and trees have been
submitted to TreeBase (http://purl.org/phylo/treebase/phylows/study/TB2:S17292).
INTRODUCTION
Cape Primroses (Streptocarpus Lindl.) and African Violets
(Saintpaulia H.Wendl.) have a significant economic importance with an estimated annual world trade value of over
$30 million (Buta & al., 2010). The taxonomy of such important horticultural plants is of interest for horticulturists and
botanists alike, for the global trade and the communication
of research findings. A recent classification has synonymised
Saintpaulia with Streptocarpus but did not consider the many
other African and Madagascan and Comoro Island (Afro-Malagasy) Gesneriaceae genera (Christenhusz, 2012). A new family classification for the Gesneriaceae was published recently,
following molecular phylogenetic relationships (Weber & al.,
2013). Here, we follow this approach for the Afro-Malagasy
Gesneriaceae. Saintpaulia and Streptocarpus are among the
10 Gesneriaceae genera in the subfamily Didymocarpoideae
occurring in Africa, Madagascar and the Comores (Möller
& Clark, 2013; Weber & al., 2013). The other genera are Epithema Blume, Acanthonema Hook.f., Colpogyne B.L.Burtt,
Received: 14 Mar 2015 | returned for (first) revision: 25 Apr 2015 | (last) revision received: 25 Jun 2015 | accepted: 26 Jun 2015 || publication date(s):
online fast track, n/a; in print and online issues, 31 Dec 2015 || © International Association for Plant Taxonomy (IAPT) 2015
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Nishii & al. • Streptocarpus redefined
Hovanella A.Weber & B.L.Burtt, Linnaeopsis Engl., Nodonema
B.L.Burtt, Schizoboea (Fritsch) B.L.Burtt and Trachystigma
C.B.Clarke (Burtt & Wiehler, 1995; Weber, 2004). Within
the subfamily Didymocarpoideae, Epithema belongs to tribe
Epithemateae, subtribe Epithematinae, a South and Southeast
Asian group, with only E. tenue C.B.Clarke occurring in Africa
(Mayer & al., 2003). The other nine genera belong to subtribe
Streptocarpinae in tribe Trichosporeae (Weber & al., 2013),
and are the subject of the study presented here. One of them,
Streptocarpus, is characterized by twisted capsules, while the
remainder have straight (non-twisted) capsules (Weber, 2004;
Weber & al., 2013).
Most genera in the Streptocarpinae are monotypic (Colpogyne, Nodonema, Schizoboea, Trachystigma), or small (Acanthonema 2 spp., Hovanella 2–3 spp., Linnaeopsis 3 spp.), with
only Streptocarpus being large (154 spp.; Hilliard & Burtt,
1971; Skog & Boggan, 2007; Möller & Clark, 2013). Acanthonema, Linnaeopsis, Nodonema, Saintpaulia, Schizoboea,
and Trachystigma are endemic to Africa and Colpogyne and
Hovanella endemic to Madagascar. Streptocarpus is the most
widespread genus, occurring in Africa (111 spp.), Madagascar
and the Comores (43 spp.) and Asia (4 spp.; Hilliard & Burtt,
1971), although the Asian species are not allied to the AfroMalagasy species (Hellens & al., 1999; Möller & al., 2009)
and fall outside the new classification proposed in this paper.
Earlier molecular phylogenetic analyses have shown that
subtribe Streptocarpinae forms a strongly supported clade
(Möller & Cronk, 1997a, b; Möller & al., 2009, 2011a; Weber
& al., 2011a). Phylogenetic studies using the internal transcribed
spacer (ITS) sequences of nuclear ribosomal DNA (nrDNA)
indicated that Saintpaulia, Linnaeopsis and Schizoboea are
nested within Streptocarpus (Möller & Cronk, 1997a, 2001a, b;
Möller & al., 1999). The nested position of Saintpaulia was
supported by chloroplast DNA (cpDNA) ndhF sequence data
(Smith & al., 1998). Later analyses using additional cpDNA
sequences (trnLF, atpB-rbcL: Möller, 2003; Möller & al., 2009)
showed that Acanthonema, Colpogyne and Hovanella are also
nested within Streptocarpus. Thus far, all Streptocarpinae analyzed are phylogenetically nested within Streptocarpus. Only
Nodonema and Trachystigma have not been placed using DNA
sequence data to date.
On the basis of molecular phylogenies, several genera have
already been synonymized with Streptocarpus, namely Linnaeopsis (Darbyshire, 2006), Saintpaulia, Schizoboea, Colpogyne
and Hovanella (Christenhusz, 2012). With respect to Saintpaulia, these studies also considered a number of the 21 species
accepted by Burtt (1958a, 1964) to be synonyms, reducing the
number of accepted species to 6 (Darbyshire, 2006) and a further 2 species being considered as synonyms by Christenhusz
(2012). However, molecular phylogenetic relationships among
species (e.g., Möller & Cronk, 1997b) appear not to have been
taken into consideration fully.
Once the first molecular phylogenies became available,
it became apparent that the generic and infrageneric circumscription within subtribe Streptocarpinae was in need of
revision (e.g., Möller & Cronk, 1997a, b; Smith & al., 1998;
Möller & Cronk, 1999). The division of Streptocarpus into two
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subgenera was based on morphological and cytological grounds
(Hilliard & Burtt, 1971). Streptocarpus species, like most Old
World Gesneriaceae (Burtt, 1970), have unequally developing
seedling cotyledons (anisocotyly) due to the extended activity of
a basal meristem in one cotyledon (macrocotyledon) (e.g., Jong,
1970; Jong & Burtt, 1975). The macrocotyledon of caulescent
species grows for a few days or weeks only and then becomes
petiolated (Jong, 1970). In acaulescent species, to which unifoliates and rosulates belong, the macrocotyledon continues
to grow for months, and in extreme cases, for years in some
unifoliates, until inflorescences are produced (Hilliard & Burtt,
1971). Because of the extended period of growth and the bearing
of inflorescences, Jong (1970) termed the macrocotyledonary
leaf a phyllomorph. In unifoliates only the seedling cotyledonary phyllomorph is produced, while in rosulates further phylomorphs are produced from a meristem at the base of the lamina
of a preceding phyllomorph. Based on chromosome data then
known, Hilliard & Burtt (1971) placed the acaulescent species
into subg. Streptocarpus, and the caulescent ones into subg.
Streptocarpella, with the former having x = 16, and the latter
x = 15. The subgeneric division based on chromosome number
showed inconsistencies when it was found that the basic chromosome number for the caulescent Streptocarpus schliebenii
Mansf. was x = 16 (Milne, 1975). The molecular phylogenetic
position of this species in the genus was, however, consistent
with chromosome counts, since this species fell among acaulescent species of subg. Streptocarpus (Möller & Cronk, 2001a, b).
In their monograph, Hilliard & Burtt (1971) pointed out that
many Madagascan species were insufficiently known, and that
groups of species exist which cannot be easily assigned to either
of the two subgenera: (i) herbaceous species with leaves in a
basal rosette with long petioles, sharply marked off from the
orbicular blades, unbranched veins arising from the midrib,
filaments fused to about a third of the corolla tube; (ii) herbaceous species with leaves in a basal rosette (rarely solitary),
often attenuate at the base (petiole ill-defined), veins ascending
from the base and branched, filaments fused to about a third
of the corolla tube; and (iii) species of shrubby-woody habit,
unbranched veins arising from the midrib, characterized also by
short filaments arising at the base of the corolla. The presence
of sclereids and non-coherent anthers can be added as further
synapomorphies for the last group of shrubby Streptocarpus
species (Hilliard & Burtt, 1971; Jong & al., 2012). The shrubby
species share the basic chromosome number of x = 16 (Jong
& Möller, 2000; Möller & Kiehn, 2004) and form a clade with
acaulescent species (Möller & Cronk, 2001a, b; Möller & al.,
2009, 2011a). Members of group (iii) were placed in subg. Streptocarpella by Hilliard & Burtt (1971), although there are other
compelling reasons not to place them there. Members of group
(i) were also placed in subg. Streptocarpella and those of group
(ii) in subg. Streptocarpus (Hilliard & Burtt, 1971). However,
since increased morphological, cytological and molecular phylogenetic information for the Malagasy Streptocarpinae species
has become available, it is time to propose a new classification.
Molecular phylogenetic studies that include samples of
subtribe Streptocarpinae are somewhat scattered and did not
include large combined datasets of both nuclear and chloroplast
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genes. Here we present data of two nuclear and three chloroplast
intron-spacer regions (ITS at genus level; 5S non-transcribed
spacer of nrDNA [5S NTS] at sectional level for Saintpaulia;
rpl20-rps12 spacer, trnL intron, trnLF spacer) for 260 samples covering ~80% of species of subtribe Streptocarpinae,
including all genera except the monotypic genus Nodonema for
which no suitable material for DNA extraction was available.
Furthermore, several species morphologically linked to Str.
schliebenii, that had not been included in previous molecular
studies, i.e., Str. montanus Oliv. and Str. parensis B.L.Burtt
from Tanzania have also been included with multiple samples
here, as well as early diverging lineages of Saintpaulia. These
samples cover important gaps in the morphological diversity
of the Afro-Malagasy Gesneriaceae.
The aim of our study is to provide a state of the art classification for Afro-Malagasy Gesneriaceae, based on our current
knowledge of phylogeny, morphology and cytology. We present phylogenies augmented with a considerable amount of new
data, both in terms of taxon and gene sampling. The phylogenies
shed light on the geographic relationships among the species
and the distribution of key morphological and basic chromosome numbers of the species. Based on this evidence, a new
infrageneric division of Streptocarpus with two subgenera and
12 sections is proposed. We also aim to re-evaluate and adjust
the species delimitations in the erstwhile genus Saintpaulia.
MATERIALS AND METHODS
Plant materials. — For simplicity, we have adopted the
generic concept of Möller & Clark (2013) and Weber & al.
(2013), who recognized nine genera among the Afro-Malagasy
Streptocarpinae. We gathered 260 samples including 189 samples of Streptocarpus (116 out of 154 described species, 75%),
58 samples of Saintpaulia (including all species and infraspecific taxa described, except for the doubtful Sa. amaniensis
F.Roberts; see Burtt, 1958a), 4 samples of Linnaeopsis (1 out
of 3 species), two of Hovanella (2 out of 2 species), 2 of Acanthonema (1 out of 2 species), 2 each of the monotypic Schizoboea (Sch. kamerunensis (Engl.) B.L.Burtt) and Colpogyne
(C. betsiliensis (Humbert) B.L.Burtt), and 1 of the monotypic
Trachystigma (T. mannii C.B.Clarke) (Appendices 1, 2).
Outgroup taxa for the Streptocarpus analyses were chosen on the basis of previous work (Möller & Cronk, 1997a, b,
2001a, b; Möller & al., 2009, 2011a; Weber & al., 2011a). All
came from tribe Trichosporeae and include the European
Haberlea rhodopensis Friv. of subtribe Ramondinae, Paraboea rufescens (Franch.) B.L.Burtt of subtribe Loxocarpinae,
and two samples of subtribe Didymocarpinae, Didymocarpus
citrinus Ridl. and Primulina spadiciformis (W.T.Wang) Mich.
Möller & A.Weber.
DNA extraction, PCR, and direct sequencing. — Genomic
DNA was extracted following Möller & al. (2009) and De
Villiers & al. (2013). DNA extractions from herbarium material were carried out using the CTAB method (Doyle & Doyle,
1987, 1990), but with a longer precipitation time that increased
the amount of recovered DNA.
PCR and sequencing approaches followed that of Möller
& Cronk (1997a), Mayer & al. (2003) and De Villiers & al. (2013).
For the ITS region, containing the internal transcribed spacers
1 and 2 and the intervening 5.8S gene, the primers used were
ITS5P and ITS8P (Möller & Cronk, 1997a), for trnL intron and
trnLF spacer (trnLF), primers c and f (Taberlet & al., 1991), and
for rpl20-rps12, primers rpl20 and 5′-rps12 (Hamilton, 1999).
For partially degraded DNA from herbarium specimens,
internal primers ITS2G and ITS3P (Möller & Cronk, 1997a)
were used for ITS, primers d and e (Taberlet & al., 1991) for
trnLF, and rpl20-3r and rpl20-2f for rpl20-rps12, which were
designed in this study (rpl20-3r: 5′-CGT TGT GAA AGA CCT
ATT ACT GGA T-3′; rpl20-2f: 5′-TCT TCT CTA TAC ACC
GGA GCC-3′) to PCR amplify the regions in smaller fragments. For low yield DNA extracts, booster PCR was carried
out for the ITS region, by re-amplifying PCR products that
were initially amplified with ITS5P and ITS8P with primers
ITS1 and ITS4 (White & al., 1990).
PCR conditions for amplifying ITS, trnLF, and rpl20-rps12
were according to De Villiers & al. (2013). The short fragments obtained using internal primers were amplified under
the following conditions: 94°C for 5 min, then 35 cycles of
94°C for 30 s, 55°C for 30 s, 72°C for 30 s, with a final extension at 72°C for 10 min. The amplified PCR fragments were
purified with ExoZAP-IT (Affymetrix U.K., Voyager, U.K.)
and cycle-sequenced in both forward and reverse directions
(Terminator Ready Reaction Mix, Applied Biosystems, Foster City, California, U.S.A.), using the PCR primers following the manufacturer”s protocol and analyzed on a capillary
sequencer ABI3730 at the GenePool facilities of the University
of Edinburgh, U.K. The electropherograms were assembled
and sequences edited in Sequencher v.4.7 (Gene Codes, Ann
Arbor, Michigan, U.S.A.). Base polymorphisms detected in
both forward and reverse direction electropherograms were
IUB coded. The GenBank accession numbers of these newly
acquired sequences are indicated in Appendices 1, 2.
Phylogenetic analyses. — The newly acquired sequences
and sequences downloaded from GenBank were added to existing sequence matrices (ITS: Möller & Cronk, 2001a; trnLF,
rpl20-rps12: De Villiers & al., 2013) and manually aligned by
eye. We assembled matrices with data for ITS, the trnL intron,
trnLF spacer and rpl20-rps12 spacer sequences that included all
available Afro-Malagasy samples assembled, 21 out of the 66
Saintpaulia samples representing 19 taxa, plus the 4 outgroup
samples, and rooted on Haberlea (Möller & al., 2009, 2011a).
This matrix contained a total of 226 samples (“Str226”), of which
117 (rpl20-rps12 spacer), 130 (trnL intron), 130 (trnLF spacer) and
75 (ITS) were newly acquired (Appendix 1). Additionally, to test
for incongruences in phylogenetic signal between the nuclear
and chloroplast markers, the ITS and plastid matrices (combined
trnL intron, trnLF spacer, rpl20-rps12 spacer sequences) for
both sampling levels were analysed individually. Due to missing samples for cpDNA data in the large analysis, this matrix
contained only 209 samples (“Str209-cpDNA”).
To investigate the phylogenetic relationships and taxon
coherence among the Saintpaulia samples in detail, a matrix
containing ITS and 5S NTS nrDNA sequences was assembled
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that included 57 Saintpaulia samples with multiple samples
per taxon and Streptocarpus caulescens Vatke as outgroup and
root (“Sa58”), based on the large analysis and previously published work (Möller & Cronk, 2001,a, b). Forty ITS sequences
were newly acquired, while all 5S NTS sequences were downloaded from GenBank. These were essentially those generated
by Lindqvist & Albert (1999, 2001) and had been generated for
the same accessions as the ITS data (Appendix 2). The ITS
region resides in the tandemly arranged 18S, 5.8S and 26S ribosomal genes that are clustered in the nucleolar organizer region
(NOR; Hillis & Dixon, 1991). The 5S gene, between which the
5S NTS resides, is also tandemly repeated but not linked to the
other three genes in plants except bryophytes and can exist on
separate chromosomes (Murata & al., 1997; Sone & al., 1999).
Thus, to test for phylogenetic congruences, separate analyses
were carried out for the ITS region and 5S NTS sequences on
matrices of both sampling levels. Due to missing sequences of
the larger set, the ITS matrix contained 47 (“Sa47-ITS”), and
the 5S NTS matrix 49 sequences (“Sa49-5S”).
All matrices and trees have been submitted to TreeBase,
study no. TB2: S17292 (http://purl.org/phylo/treebase/phylows/
study/TB2:S17292).
It is well known that homoploid hybridization is widespread in Streptocarpus, particularly among members of the
South African Cape Primrose clade (Hilliard & Burtt, 1971; De
Villiers & al., 2013). To assess the impact of this on the phylogenetic analysis we compared topology differences among
the individual marker analyses with reference to major clades
(that we use as the basis for the classification here) for the
Streptocarpus analyses and “species” monophyly for the Saintpaulia analyses. Incongruences supported by higher than 75%
bootstrap values and/or 0.95 posterior probabilities would be
regarded as significant.
A limited proportion of within-electropherogram single
nucleotide polymorphisms were found in ITS and 5S NTS
(Electr. Suppl.: Table S1), amounting to 0.06% for the Streptocarpus matrices and 0.02%–0.39% for the Saintpaulia matrices.
These polymorphisms may have their origin in past hybridisation and incomplete lineage sorting events (De Villiers & al.,
2013). Alternatively, they may result from incomplete concerted
evolution across the two NOR sites known to be present in
Streptocarpus (Möller & al., 2008). Mutations arising here may
sometimes be transiently present in evolutionary terms (others
may become homogenised in parallel with speciation events;
Hughes & al., 2005), reducing their significance in phylogenetic analyses. Polymorphisms may also arise as artefacts of
PCR and/or sequencing protocols (cf. Möller, 2000). Because
of the complex nature of polymorphisms in the multicopy
ribosomal DNA sequences, their low levels of occurrences in
the matrices and the fact that the matrices consisted of a mix
of sequences downloaded fromGenBank and newly acquired
sequences, the scoring of polymorphisms was not necessarily uniformly performed, and so these polymorphisms were
treated as uncertainties in the phylogenetic analyses.
Maximum parsimony (MP) and Bayesian inference (BI)
analysis were run in PAUP* v.4.0b10 (Swofford, 2002), and
MrBayes v.3.2.2 (Ronquist & al., 2012), respectively. The MP
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analysis followed Möller & al. (2009). Best-fitting models for
the BI analyses were obtained independently for each gene
region using MrModeltest v.2.3 (Nylander, 2004). Under the
Akaike information criterion (Akaike, 1974), GTR + I + G was
given for rpl20-rps12, GTR + G for trnL intron and trnLF spacer,
SYM + I + G for both the 5.8S gene region and ITS spacers for
matrix Str226. For matrix Sa58, K80 + I was given for the 5.8S
gene, SYM + G for the ITS spacers, and K80 + G for the 5S NTS
sequences. For matrix Str226 10 million generations and for
Sa58 5 million generations were run in two independent parallel runs and sampled every 1000th generation. The first 500
(Str226) or 250 (Sa58) trees (5%) were discarded as burn-in prior
to calculating the BI consensus trees and posterior probabilities
(PP). The statistics for each analysis are shown in Figs. S1, S4,
S7, S10, S13, S16 (Electr. Suppl.), as run online under AWTY
(Nylander & al., 2008). RAxML analysis was carried out using
the raxmlGUI (Silvestro & Michalak, 2012), with GTRGAMMA
setting and 1000 bootstrap replicates. Partitions were set for
rpl20-rps12, trnL intron and trnLF spacers, ITS1, 5.8S, ITS2
and 5S NTS sequences (Marx & al., 2010).
Geography, morphology and cytology. — Geographical
and morphological data were taken from living plants cultivated
in the research collection at the Royal Botanic Garden Edinburgh
(RBGE), herbarium specimens at E, and consultation of the relevant literature, primarily Hilliard & Burtt (1971) for Streptocarpus, and for other genera as cited hereafter in the relevant
sections. Living material was either fixed in Farmer’s Fluid to
aid clearing (Jong & al., 2012), or directly observed under a dissecting microscope. Some material used came from the RBGE
spirit collection and was preserved in Copenhagen mix.
Streptocarpus species display an array of morphological
diversity (Jong, 1970; Hilliard & Burtt, 1971; Jong & Burtt,
1975). We categorized the species into the main forms irrespective of herbaceous or woody habit, including caulescents
with stems and decussate leaf arrangement (Fig. 1A), plants
with creeping rhizomatous stem and alternate leaf arrangement (Fig. 1B), Saintpaulia-like rosettes (Fig. 1C), acaulescents
lacking a stem and shoot apical meristem and either producing
leaves in an irregular rosette (rosulates; Fig. 1D), or only retaining a single enlarged cotyledon as the sole photosynthetic organ
(unifoliates; Fig. 1E).
We used the categorization into floral types of Harrison
& al. (1999) for Streptocarpus flowers, these included the small
pouch (Fig. 1F), open tube (Fig. 1G), personate (having the
lower lip pushed upward so as to close the gap between the
lips, as in the snapdragon; Fig. 1H), keyhole (where the opening of the corolla is laterally compressed to a narrow vertical
slit; Fig. 1I), Saintpaulia-type (Fig. 1J), and bird-pollination
(possessing a red tubular flower; Fig. 1K). We distinguished
a further two types, the Acanth-type (Fig. 1L) for the recently
described Str. lilliputana Bellstedt & T.J.Edwards that exhibits
an extremely long proximate alignment channel (viz. Westerkamp & Claßen-Bockhoff, 2007), and the labellanthus-type
with a forward directing lip and reduced upper lip (Fig. 1M).
Seeds were either categorised as reticulate, where the lateral
testa cell walls form a raised reticulum on the seed surface, and
verruculose where the outer testa cell walls produce a papilla.
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Data for basic chromosome numbers were taken predominantly from the RBGE WebCyte2 (Möller & Pullan, 2015–),
from Briggs (2004), or obtained from mitotic chromosome
counts according to Jong & Möller (2000).
RESULTS
Phylogenetic congruence between the individual marker
matrices. — A comparison of the topologies of the ITS with
the cpDNA trees for the Streptocarpus analyses revealed no
strong conflict in relationships between the main clades (Electr.
Suppl.: Figs. S1–S6; summarised in Table S2). Some “clades”,
such as “h” and “i” were not resolved as monophyletic in some
analyses, especially those involving ITS matrices, but no
strongly supported topological conflict existed. A similar situation was revealed for clades “k” and “l”. All clades received
predominantly high support in the combined ITS & cpDNA
analyses. For Saintpaulia, the MP, ML and BI analyses of neither ITS nor 5S NTS separately indicated strongly supported
incongruences in topologies (Electr. Suppl.: Figs. S7–S12). A
few instances occurred of changing relationships between ITS
and 5S NTS trees. None received consistently strong support.
As the Streptocarpus and Saintpaulia analyses revealed no
strong topological incongruences, our report is based on the
results of the combined analyses only.
Fig. 1. Major growth forms (A–E)
and floral types (F–M) found
among the Afro-Malagasy
Gesneriaceae. A & C, modified
from Humbert (1971: figs. XIII4,
X8, © Publications Scientifiques
du Muséum national d’Histoire
naturelle, Paris); B, from Weigend
(2000: fig. 2:5, © Elsevier Ltd.
Oxford); D–E, from Hilliard &
Burtt (1971: figs. 2, 5, © University of Natal Press, Pietermaritzburg); F–K, from Harrison & al.
(1999: fig. 1A, © Oxford University Press); L, from Bellstedt &
Edwards (2004: fig. 1D–E,
© Cambridge University Press);
M, modified from Hilliard &
Burtt (1975: figs. 1Aa–1Ab,
© Royal Botanic Garden Edinburgh).
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Characteristics and phylogenetic relationships revealed
by the Str226 analysis. — The combined rpl20-rps12 spacer,
trnL intron, trnLF spacer and ITS matrix included 2589 characters, of which 1526 (58.9%) were constant, 298 (11.5%) were parsimony uninformative and 765 (29.6%) parsimony informative.
The MP analysis resulted in 48,847 most parsimonious
trees of a length of 3021 steps (consistency index [CI] = 0.5144,
retention index [RI] = 0.8610). The BI tree of the same matrix
had a topology very similar to the RAxML tree and MP consensus tree (Electr. Suppl.: Figs. S13, S14, S15 respectively). The
subclades relevant for the subgenus and sectional division were
identical in all analyses, with minor differences in unsupported
topologies within subclades. The Bayesian inference phylogenies are presented in the main text, with branch support values
from the BI, ML and MP analyses shown for main branches
(Fig. 2A–B). The individual analyses with all support values
are given in Figs. S13–S15 (Electr. Suppl.).
Two strongly supported large clades were identified, one
(Clade I, PP = 1.00; ML = 99%; BS = 84%; Fig. 2A) being
composed mainly of subg. Streptocarpella and Saintpaulia, and
the other (Clade II, PP = 1.00; ML = 100%; BS = 92%; Fig. 2B)
including mostly subg. Streptocarpus, but also some caulescent species (see below). Within Clade I, several subclades
were strongly supported; subclade “a” contains Str. nobilis
C.B.Clarke, Trachystigma and the Acanthonema samples (PP
= 1.00; ML = 97; BS = 75%), subclade “b” (PP = 1.00; ML =
100%; BS = 99%) contains the Madagascan species of group (i)
of Hilliard & Burtt (1971). Subclade “c” (PP = 1.00 ML = 100%;
BS = 100%) contains the Madagascan herbaceous species and
the two Hovanella species. Subclade “d” (PP = 1.00; ML =
99%; BS = 92%) was composed of Str. thysanotus Hilliard
& B.L.Burtt, Str. kimbozanus B.L.Burtt, Str. elongatus Engl.
and the Schizoboea samples. Subclade “e” (PP = 1.00; ML =
100%; BS = 100%) included three species, Str. saxorum Engl.,
A
subclades
1/97/75
0.56
68
*
1
100
100
1/100/99
1
100
100
1/99/84
1/99/92
Clade I
1
97
81
1/100/100
1
99
95
1
100
100
1/100/96
0.99
75
81
1
100
99
Clade II
Hab.rhodopensis
Par.rufescens
Did.citrinus
Prim.spadiciformis
Str.nobilis -1
Str.nobilis -2
Str.nobilis -3
Tra.mannii
Aca.strigosum -1
Aca.strigosum -2
Str.beamp.ssp.beamp -1
Str.beamp.ssp.beamp -2
6x Str.andohahelensis -1
6x Str.andohahelensis -2
2x/4x Str.thompsonii -1
2x/4x Str.thompsonii -2
Hov.madagascarica
Str.venosus
Str.muscosus
Str.levis -2
Str.hilsenbergii -1
Str.hilsenbergii -2
Hov.vestita
Str.tanala -1
Str.tanala -2
Str.oliganthus -1
Str.oliganthus -2
Str.thysanotus
Str.kimbozanus -1
Str.kimbozanus -2
Schiz.kamerunensis -1
Schiz.kamerunensis -2
Str.elongatus -1
Str.elongatus -2
Str.saxorum
Str.kirkii
Str.stomandrus -1
Str.stomandrus -2
Str.inflatus
Str.glandulosissimus
Str.buchananii
Str.holstii
Str.pallidiflorus
Str.caulescens
Sa.ulugurensis
Sa.goetzeana -2
Sa.pusilla -2
Sa.inconspicua -1
Sa.inconspicua -2
Sa.watkinsii-1
Sa.watkinsii-2
Sa.teitensis -1
Sa.rupicola -1
Sa.nitida -1
Sa.brevipilosa -1
Sa.pendula
Sa.shumensis -1
Sa.diplotricha
Sa.grandifolia -1
Sa.difficilis-1
Sa.cf.ionantha
Sa.tongwensis -1
Sa.velutina
Sa.magungensis -1
Sa.grotei -1
‘a’
‘b’
‘c’
‘d’
‘e’
‘f’
‘g’
subg. Streptocarpus
Fig. 2A–B. Bayesian inference tree for matrix Str226 of combined ITS, trnL intron, trnLF spacer and rpl20-rps12 spacer sequences, with branch
support (BI / ML / MP) along the branches, indicated for branches supporting sections and higher-level taxa. For intrasectional branch support
values see Electr. Suppl.: Figs. S1A, S2, S3. * indicates branches receiving values less than 0.5 (BI) or 50% (ML, MP). Characters for subg.
Streptocarpella (A) and subg. Streptocarpus (B) are indicated. First column, country of origin (geography): ■ Africa, ▲ Madagascar and
Comores. Second column, seed ornamentation (seed): ▲ verruculose, ■ reticulate. Third column, growth form (habit): ● caulescent, creeping
rhizomatous stem, ■ rosulate, ▬ Saintpaulia-like rosette, ▲ unifoliate, ç shrubby. Fourth column, floral types: ▲ small pouch, ◊ labellanthus,
□ keyhole, ● open tube, ▬ Saintpaulia, personate, ç bird-pollination, Acanth. Fifth column, basic chromosome number (chromosomes): ▲
x = 15, ■ x = 16; ploidy level: given as numbers to the right. Gaps indicate missing data.
1248
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B
subclades
Clade I
Hab.rhodopensis
Par.rufescens
Did.citrinus
Prim.spadiciformis
subg. Streptocarpella
Str.papangae-1
Str.papangae-2
Str.tsaratananensis
Str.suffruticosus
Str.macropodus
Str.lanatus-1
Str.lanatus-2
Str.lanatus-3
Str.itremensis-1
Str.itremensis-2
Str.itremensis-3
Str.itremensis-4
Col.betsiliensis-1
Col.betsiliensis-2
Str.ibityensis-1
Str.ibityensis-2
Str.ibityensis-3
Str.exsertus-1
Str.exsertus-2
Str.capuronii-1
Str.capuronii-2
Str.brevistamineus
8x Str.hildebrandii
Str.sambiranensis
6x Str.variabilis-3
6x Str.variabilis-1
6x Str.variabilis-2
2x/4xStr.perrieri
Str.cf.mangindranensis
Str.lokohensis
Str.parensis-1
Str.parensis-2
Str.schliebenii-1
Str.schliebenii-2
Str.schliebenii-3
Str.montanus-1
Str.montanus-2
Str.montanus-3
Str.bullatus-2
Str.bullatus-1
Str.bullatus-3
Lin.alba
Lin.sp.-3
Lin.sp.-1
Lin.sp.-2
Str.huamboensis
Str.hirticapsa-3
Str.hirticapsa-1
Str.hirticapsa-2
Str.nimbicola
Str.hirtinervis
Str.dolichanthus
Str.milanjianus
Str.decipiens
Str.galpinii
Str.micranthus-1
Str.micranthus-2
Str.umtaliensis
Str.cf.michelmorei-1
Str.cf.michelmorei-2
Str.davyi
Str.pusillus-1
Str.pusillus-2
Str.occultus
Str.fanniniae-1
Str.fanniniae-2
Str.daviesii
Str.wilmsii
Str.makabengensis
Str.rimicola-1
Str.rimicola-2
Str.erubescens
Str.cyanandrus
Str.pumilus
Str.pole-evansii
Str.dunnii-3
Str.dunnii-1
Str.dunnii-2
Str.denticulatus-1
Str.denticulatus-2
Str.grandis-1
Str.grandis-2
Str.molweniensis
Str.wittei
Str.cooperi
Str.cooksonii
Str.solenanthus
Str.bindseilii
Str.kungwensis
Str.monophyllus-1
Str.monophyllus-2
Str.compressus
Str.vandeleurii-1
Str.vandeleurii-2
Str.eylesii
Str.goetzei
Str.bolusii-1
Str.bolusii-2
Str.porphyrostachys-1
Str.porphyrostachys-2
Str.confusus
Str.haygarthii
Str.trabeculatus
Str.saundersii-2
Str.prolixus
Str.pentherianus
Str.polyanthus-1
Str.silvaticus
Str.polyanthus-2
Str.sp.nov.-1
Str.sp.nov.-2
Str.saundersii-1
Str.wendlandii-1
Str.wendlandii-2
Str.caeruleus-1
Str.caeruleus-2
Str.candidus
Str.gardenii-1
Str.gardenii-2
Str.longiflorus
Str.kentaniensis-1
Str.kentaniensis-2
Str.aylae
Str.kunhardtii
Str.floribundus
Str.johannis-2
Str.lilliputana-1
Str.lilliputana-2
Str.parviflorus.subsp.soutpansbergensis
Str.cyaneus.subsp.nigridens
Str.montigena-1
Str.montigena-2
Str.meyeri-6
Str.meyeri-3
Str.meyeri-4
Str.meyeri-5
Str.primulifolius-1
Str.baudertii-2
Str.johannis-1
Str.formosus-1
Str.formosus-2
Str.primulifolius-2
Str.primulifolius-4
Str.fasciatus
Str.cyaneus-1
Str.fenestradei
Str.cyaneus.subsp.longitommii
Str.cyaneus-2
Str.roseoalbus
Str.parviflorus-1
Str.parviflorus-2
Str.primulifolius-6
Str.modestus-1
Str.modestus-2
Str.primulifolius-3
Str.primulifolius-5
Str.meyeri-1
Str.meyeri-2
Str.baudertii-1
Str.rexii-1
Str.rexii-2
‘h’
‘i’
‘j’
Clade II
‘k’
‘l’
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Nishii & al. • Streptocarpus redefined
Str. kirkii Hook.f., and Str. stomandrus B.L.Burtt. The species
with personate flowers were in subclade “f” (PP = 1.00; ML =
100%; BS = 96%). The final subclade in Clade I, subclade “g”
(PP = 1.00; ML = 100%; BS = 99%), contained all Saintpaulia
species included in this analysis. Relationships between the
subclades were found to be identical in the three phylogenetic
analyses of the Str226 dataset and well supported in the BI
(PP = 0.98–1.00), ML (75%–100%) and MP (BS = 81%–100%)
analyses, except for the position of subclade “a”, which was
sister to subclade “b” and “c” in the BI and ML analyses (Electr.
Suppl.: Figs. S13–S14), but collapsed into a basal polytomy in
Clade I in the MP analysis (Electr. Suppl.: Fig. S15).
Clade II showed similar resolution but overall less branch
support in the MP, but high branch support in the BI and ML
analyses (Figs. 2–3; Electr. Suppl.: Figs. S13–S15). Two subclades, “h” (PP = 1.00; ML = 98; BS = 86%) containing the
shrubby Madagascan species of group (iii) of Hilliard & Burtt
(1971), and subclade “i” (PP = 1.00; ML = 98%; BS = 88%)
including Madagascan Streptocarpus species with a dense
woolly indumentum, plus Colpogyne, formed sister clades
(PP = 1.00; ML = 97%; BS = 86%) on the earliest diverging
lineage in Clade II. The two samples of Str. exsertus Hilliard
& B.L.Burtt and Str. capuronii Humbert were resolved in different positions in the different analyses, either in a polytomy
with the remaining subclades in the BI analysis (Electr. Suppl.:
Fig. S13), in a basal polytomy of Clade II in the MP analysis
(Electr. Suppl.: Fig. S15), or resolved in grades between the two
subclades “h” and “i” in the ML analysis (Electr. Suppl.: Fig.
S14), although never with significant branch support. Subclade
“j” (PP = 1.00; ML = 81%; BS = 85%) included species of group
(ii) of Hilliard & Burtt (1971) with ascending veins, and subclade “k” (PP = 0.98, ML = 75%; BS = 59%) contained the African species Str. bullatus Mansf., Str. montanus, Str. parensis,
Str. schliebenii, and the Linnaeopsis samples. The remaining
107 samples were unifoliate and rosulate African species and
formed subclade “l” (PP = 1.00; ML = 80%; BS < 50%). Within
this subclade there were several larger supported clades, such
as the ITS2 deletion clade (PP = 1.00; ML = 98%; BS = 90%)
and an extended Cape Primrose clade (sensu Möller & Cronk
2001a, b) (PP = 1.00; ML 91%; BS < 78%) (see Figs. S1A, S2,
S3 in Electr. Suppl.), that includes exclusively rosulate species
geographically restricted to the northern and eastern provinces
of South Africa and Swaziland.
Characteristics and phylogenetic relationships revealed
by the Sa58 analysis. — The combined ITS, 5S NTS nrDNA
matrix had 991 characters of which 690 (69.6%) were constant,
97 (9.8%) uninformative and 204 (20.6%) parsimony-informative in matrix Sa58.
In the MP analysis 13,925 most parsimonious trees were
found of 469 steps (CI = 0.8081, RI = 0,9112). In the MP majority consensus tree, BI and ML trees, sequences of samples
from the Uluguru Mts. (PP = 1.00; ML = 97%; BS = 94%),
Teita Hills (PP = 1.00; ML = 100%; BS = 100%), and Nguru
Mts. (PP = 1.00; ML = 98%; MP = 98%) were retrieved in
respective clades, with identical topologies between these
clades in all three analyses and high branch support (Electr.
Suppl.: Figs. S16–S18; summarized in Fig. 4). Within each of
1250
these clades, the samples fell into species-specific clades in
the case of Sa. watkinsii Haston (PP = 1.00; ML = 100%; BS =
98%), Sa. inconspicua B.L.Burtt (PP < 0.50; ML < 50%; BS
< 50%), Sa. pusilla Engl. (PP = 1.00; ML = 97%; BS = 85%),
Sa. goetzeana Engl. (PP = 1.00; ML = 100%; BS = 100%), Sa.
teitensis B.L.Burtt (PP = 1.00; ML = 100%; BS = 100%), Sa.
brevipilosa B.L.Burtt (PP = 1.00; ML = 100%; BS = 91%), and
Sa. nitida B.L.Burtt (PP = 1.00; ML = 98%; BS = 86). Only
for Sa. inconspicua were different topologies of the individual
samples found between the analyses, as sister pair in the MP,
in grades in the ML and in separate positions in the BI analysis
(Electr. Suppl.: Figs. S16–S18). This might be linked to the fact
that of Sa. cf. inconspicua-1, only ITS1 sequences were available and no 5S NTS data.
The clade of the West Usambara Mts. species Sa. shumensis B.L.Burtt (PP = 1.00; ML = 100%; MP = 91%) fell between
the Nguru Mts. clade and the remainder of the Saintpaulia
samples. These were the Usambara Mts. samples and the
Kenyan samples of Sa. rupicola B.L.Burtt that formed a clade
without support, and the latter in a separate clade (PP = 1.00;
ML = 96%; BS = 92%) within the former. Where multiple
taxon samples existed within the Sa. ionantha complex (sensu
Möller & Cronk, 1997b) none, except Sa. rupicola, formed a
taxon-specific clade.
Character distribution. — As the relationships at the base
of Clade II were unresolved, and the Saintpaulia analyses
showed a polytomy for the Sa. ionantha complex (= Str. ionanthus complex), the characters for geographic origin, morphology
and cytology were presented next to the taxa in the phylogenetic trees (Fig. 2, summarized for the Streptocarpus analysis in
Fig. 3, Fig. 4), rather than carrying out a character optimization,
and their utilization for the classification discussed.
DISCUSSION
In this study we have assembled the most comprehensive
molecular dataset to date for the Afro-Malagasy didymocarpoid Gesneriaceae, that include the well-known African Violets
and Cape Primroses, covering about 83% out of the 211 taxa in
176 species (equivalent to 76% sampling at the species level).
On the basis of the phylogenetic trees reconstructed, we assess
morphological character states and their use in a new classification. Such an approach has been widely applied before
(e.g., Marx & al., 2010; Möller & al., 2011b; Weber & al., 2011b;
Iles & al., 2012). Our results are in stark contrast to previous
classifications of the Afro-Malagasy Gesneriaceae that viewed
the traditional genera as independent evolutionary lineages
(Hilliard & Burtt, 1971; Weber, 2004; Weber & al., 2013). The
results are particularly significant for the horticulturally important genus Saintpaulia, which can be distinguished morphologically from an archetypical Streptocarpus such as the rosulate
Str. rexii (Bowie ex Hook.) Lindl. or the caulescent Str. caulescens (Hilliard & Burtt, 1971). However, with the increased
knowledge of the Madagascan species these morphological
boundaries became weakened (Briggs, 2004), and its inclusion
in Streptocarpus seems a viable option. The ramifications of
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our findings with view to a new classification of the AfroMalagasy Gesneriaceae are discussed in detail here.
Molecular phylogenetic structure. — Overall, we found
that the reconstructed phylogenies showed few incongruences
between the plastid and nuclear datasets at the clade level, and
none were strongly supported. The largest matrix with combined
ITS, trnL intron, trnLF spacer and rpl20-rps12 spacer sequences
resulted in a well-resolved and generally well-supported tree
topology. The high resolution is perhaps the effect of the combination of fast evolving ITS with slower evolving plastid intron
and spacer sequences, a well-known observation (e.g., Sullivan,
1996; Gontcharov & al., 2004). This is despite the fact that this
matrix was incomplete and had data gaps in the plastid sequence
partitions. The effect of missing data on phylogenies has been
investigated previously and their effect on retrieving accurate
topologies found to be minimal (e.g., Wiens, 2003, 2006; also
in Gesneriaceae, Möller & al., 2009).
Based on our phylogenetic analyses using three different
approaches, BI, ML, and MP, we retrieved very strong support
for the finding that all Afro-Malagasy Gesneriaceae genera with
non-twisted fruits included in this analysis, i.e., Acanthonema,
Colpogyne, Hovanella, Linnaeopsis, Saintpaulia, Schizoboea
and Trachystigma, have evolved from within the genus Streptocarpus (Figs. 2–3). We found only small differences in branch
support between the BI, ML, and MP analyses, increasing the
confidence for inferences regarding sectional coherence and
Hab.rhodopensis
Par.rufescens
Did.citrinus
Prim.spadiciformis
Clade I
x = 15
Clade II
x = 16
‘a’ Trachystigma
Af
ver
cau/uni open t. tw
‘b’ Parasaintpaulia
Mg
ver
sp
pouch tw
‘c’ Hova
Mg
ver
cau
pouch tw
‘d’ Schizoboea
Af
ver
cau
lab.
tw
‘e’ Carnosifolii
Af
ver
cau
mix
tw
‘f’ Caulescentes
Af
ver
cau
pers.
str
‘g’ Saintpaulia
Af
ver
sp/cau
sp
tw
‘h’ Lignostreptocarpus
Mg
ret
shr
pouch tw
‘i’ Colpogyne
Mg
ver
ros/uni
pouch tw
Str.exsertus 1
Af
ret
uni
pouch
tw
Str.exsertus 2
Af
ret
uni
pouch
tw
Str.capuronii 1
Mg
ret
uni
pouch
tw
Str.capuronii 2
Mg
ret
uni
pouch
tw
‘j’ Plantaginei
Mg
ret
ros/uni
mix
tw
‘k’ Protostreptocarpus
Af
ver
cau/rhz pouch tw
‘l’ Streptocarpus
Af
ret
ros/uni
mix
tw
Fig. 3. Summary of phylogenetic relationships among sections of Streptocarpus based on BI, ML, and MP analyses with branch support
(BI / ML / MP) indicated for branches supporting sections and higher-level taxa, for the Str226 analysis. * indicates branches receiving values less
than 0.5 (BI) or 50% (ML, MP); °, with minor exceptions. Lowercase letters a–l identify sections (names in bold italics). First column, geographic
origin (geography): Af., Africa; Mg; Madagascar and Comores. Second column, seed ornamentation (seed): ret, reticulate; ver, verruculose.
Third column, growth form (habit): cau, caulescent; rhz, creeping rhizomatous; ros, rosulate; shr, shrubby; sp, Saintpaulia-like rosette; uni,
unifoliate. Fourth column, floral type: pouch, small pouch; lab., labellanthus-type; mix, mixed subclades; open t., open tube; pers., personate;
sp, Saintpaulia-type. Fifth column, fruit development: str, not twisted; tw, twisted.
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Str.caulescens
Sa.ulugurensis
Sa.watkinsii-1
Sa.watkinsii-2
Sa.cf.inconspicua-1
Sa.inconspicua-2
Sa.pusilla-1
Sa.pusilla-2
Sa.goetzeana-3
Sa.goetzeana-1
Sa.goetzeana-2
Sa.teitensis-1
Sa.teitensis-2
Sa.teitensis-3
Sa.brevipilosa-1
Sa.brevipilosa-2
Sa.nitida-1
Sa.nitida-2
Sa.shumensis-1
Sa.shumensis-2
Sa.diplotricha
Sa.ion.SigiFalls
Sa.grotei-1
Sa.grandifolia-2
Sa.grandifolia-3
Sa.grandifolia-1
Sa.mag.var.min.-2
Sa.magungensis-1
Sa.cf.ionantha
Sa.tongwensis-2
Sa.tongwensis-3
Sa.orb.var.pur.
Sa.velutina
Sa.tongwensis-1
Sa.confusa-2
Sa.difficilis-2
Sa.magungensis-2
Sa.grotei-2
Sa.orbicularis-2
Sa.difficilis-1
Sa.mag.var.min.-1
Sa.orbicularis-3
Sa.orbicularis-1
Sa.difficilis-3
Sa.confusa-1
Sa.pen.var.pen.
Sa.mag.var.occ.
Sa.grandifolia-4
Sa.intermedia
Sa.pen.var.Kiz.
Sa.ionantha
Sa.rupicola-1
Sa.rupicola-2
Sa.rupicola-3
Sa.rup.Kachor.-2
Sa.rup.Mwarak.
Sa.rup.Kachor.-1
Sa.rup.Mwache.
Ulu.
Ulu.
Ulu.
Ulu.
Ulu.
Ulu.
Ulu.
Ulu.
Ulu.
Ulu.
Teit.
Teit.
Teit.
Ngu.
Ngu.
Ngu.
Ngu.
W.Us.
W.Us.
E.Us.
E.Us.
E.Us.
W.Us.
W.Us.
W.Us.
E.Us.
E.Us.
E.Us.
E.Us.
E.Us.
W.Us.
E.Us.
E.Us.
E.Us.
E.Us.
E.Us.
E.Us.
W.Us.
E.Us.
E.Us.
W.Us.
W.Us.
E.Us.
E.Us.
E.Us.
W.Us.
W.Us.
E.Us.
E.Us.
E.Us.
K.Ka.
K.Ka.
K.Ka.
K.Ka.
K.Ka.
K.Ka.
K.Ka.
I
I
I
I
I
R
R
I
I
I
I
I
I
R
R
R
R
R
R
R
R
I
R
R
R
I
I
R
R
R
R
R
R
R
R
I
I
R
R
I
R
R
R
R
I
I
R
I
I
R
R
R
R
R
R
R
R
?
?
?
?
glab.
LE
LE
LE
LE
LE
LE
LE
LE
SE
SE
SA
SA
LE
LE
LESE
LE
LASA
LE
LE
LE
LASA
LASA
LE
LE
LE
LASA
LESE
LE
LASA
LASA
LASA
LASA
LASA
LASA
LASA
LASA
LASA
LASA
LASA
LE
LASA
LE
LA
LE
LE
SE
SE
SE
SE
SE
SE
SE
Str.ulugurensis
Str.watkinsii
Str.inconspicuus
Str.afroviola
Str.goetzeanus
Str.teitensis
Str.brevipilosus
Str.nitidus
Str.shumensis
Str.ionanthus
Fig. 4. Majority-rule consensus tree of the maximum parsimony analysis of the Saintpaulia Sa58 matrix comprised of combined ITS and 5S NTS
nrDNA data, with branch support values indicated along branches (BI / ML / MP). * indicates branches receiving values less than 0.5 (BI) or 50%
(ML, MP). First column, old names: Sa, Saintpaulia; Str, Streptocarpus. Second column, geographic origin (geography): E.Us., East Usambara
Mts.; K.Ka., Kenya, Kaloleni; Ngu., Nguru Mts.; Teit., Teita Hills; Ulu., Uluguru Mts.; W.Us., West Usambara Mts. Third column, growth
form (habit): I, opposite leaves and conspicuous internodes; R, alternate leaves in rosette. Fourth column, leaf indumentum (indumentum): A,
appressed hairs; E, erect hairs; glab., glabrous; L, long hairs; S, short hairs. Fifth column, new names at species rank.
1252
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relationships where they were strongly supported. In our phylogenetic analyses, we found two strongly supported clades.
Clade I is comprised of the herbaceous caulescent Streptocarpus species of subg. Streptocarpella including samples of five
non-twisted genera, i.e., Acanthonema, Hovanella, Saintpaulia,
Schizoboea, and Trachystigma. Clade II primarily included
species of subg. Streptocarpus and samples of Colpogyne and
Linnaeopsis possessing non-twisted fruits. Since no molecular
data could be generated for Nodonema, its taxonomic placement was decided on the basis of morphological features shared
with members of subg. Streptocarpus (see below).
Character evolution. — Despite the low resolution at the
base of Clade II that precludes a full character optimization
analysis, several conclusions can be drawn from the molecular
phylogenies and consequently for the new infrageneric classification presented here. We follow a character order of decreasing
congruence with infrageneric clade topology.
Cytology. – The basic chromosome number is the only
character that is, to our present knowledge, fully congruent
with the tree topology, albeit only to support the split between
Clade I and Clade II. An interesting aspect is the occurrence of
polyploidy. This has only been detected in the four species on
Madagascar and the Comoro Islands (Str. variabilis Humbert
on Anjouan), and in both main evolutionary lineages (Fig. 2).
This recalls the idea of island colonization for which polyploidy
has been argued to be particularly advantageous (Stebbins,
1950; Barrier & al., 1999).
Geography. – Our phylogeny of the Streptocarpinae shows
a strong geographic pattern (Fig. 2). It is interesting to note that
the species resolved into clades of either African, or Madagascan
and Comores distributions in both Clade I and, to a lesser extent,
in Clade II. This confirms that interchanges between the two
geographic regions were rare events (as previously suggested
by Möller & Cronk, 2001b). Because of the polytomy at the base
of Clade II, it is not possible to state with certainty whether the
Streptocarpinae originated in Africa or Madagascar, or whether
there were one or two geographic exchanges in Clade II.
Seed ornamentation. – The seed testa cell surface ornamentation of the Streptocarpinae is either reticulate (Fig. 5K)
or verruculose (Fig. 5L). The correlation between caulescence
and verruculose seeds, and acaulescence and reticulate seeds in
Streptocarpus on the African continent and the lack of this correlation in Madagascan and Comores species has been observed
before (Hilliard & Burtt, 1971). We can define this pattern more
precisely here. Most members of Clade I have verruculose seeds
irrespective of their geographic origin (Fig. 2A), while those
in Clade II exhibit a more complex pattern (Fig. 2B). The few
exceptions in Clade I are likely to be incorrect reports and
are dealt with in detail below. The absence of reticulate seeds
in Clade I, the mixed reticulate/verruculose subclades in the
basal lineages and the predominantly reticulate seeds of the
derived subclade “l”, may suggest that verruculose seed testa
ornamentation is the ancestral state from which reticulate seeds
have evolved. However, a conclusive answer to the evolutionary scenario here is hampered by the low basal resolution in
Clade II. However, the presence of verruculose seeds in some
members in subclade “l” (Str. dunnii Hook.f., Str. decipiens
Hilliard & B.L.Burtt, Str. hirtinervis C.B.Clarke, Str. nimbicola
Hilliard & B.L.Burtt, and some populations of Str. galpinii
Hook.f.) are clearly homoplastic.
Fruit twist. – This study shows that this character, traditionally used to define the genus Streptocarpus, is perhaps the
most labile of all the morphological characters used to define
it. The repeated loss of this character is not unprecedented
in Gesneriaceae, and has also been observed in Paraboea
(C.B.Clarke) Ridl. (Puglisi & al., 2011). In Streptocarpus, the
twisted fruit was lost on at least seven occasions (Fig. 2). Given
that the fruit twist has perhaps evolved only once or twice in
Gesneriaceae (Möller & al., 2009), the results here and perhaps
for Paraboea (Puglisi & al., 2011) are an interesting demonstration of uni-directionality in character evolution.
Floral types. – Similar to other genera of Didymocarpoideae, such as Oreocharis Benth. (Möller & al., 2011b), or
Petrocodon Hance (Weber & al., 2011b), Streptocarpus exhibits
a high diversity of floral sizes and forms which can be ascribed
to several main types (Harrison & al., 1999; Hughes & al.,
2006). However, this rough grouping does not fully reflect
floral diversity in Streptocarpus. Among the small-flowered
type, there is a group of species in subclade “d” with a forward directing lip and reduced upper lip that cannot be adequately described as a small pouch type and is described here
as labellanthus-type (Fig. 6D). The open flower type exhibits
perhaps the most diversity, and may contain several subtypes,
without or with distinct pollination chambers (viz. Westerkamp
& Claßen-Bockhoff, 2007), such as the Acanth-type present
in Str. lilliputana (Bellstedt & Edwards, 2004; De Villiers
& al., 2013). However, differences are often only slight and
some flowers are difficult to classify. Cases of parallelisms in
the evolution of main floral types occur, such as for the keyhole flower for example (Fig. 6E), occurring in Str. saxorum
in Clade I, and several species in Clade II (Fig. 2). However,
several subclades have more conserved flower characteristics
(Fig. 2A), such as subclade “d” with the labellanthus-type,
subclade “f” with personate flowers (Fig. 6F) or the flat-faced
corollas of Saintpaulia (Figs. 6G–H). Such cases have been
utilized for our classification here.
Growth form. – After the Madagascan species of Humbert
(1967, 1971) became known, the subdivision of Streptocarpus
into the unranked groups Caulescentes, Rosulati and Unifoliati
of Fritsch (1894) was considered not to capture the breadth of
morphological diversity in this genus. With the phylogenetic
inclusion of other genera of subtribe Streptocarpinae into Streptocarpus, there are at least two more fundamental growth patterns that can be recognized, (1) Saintpaulia-like rosette plants
(group (i) of Hilliard & Burtt, 1971; Fig. 7B) and (2) shrubby
caulescent plants (group (iii) of Hilliard & Burtt, 1971; Fig. 7I).
Species with a creeping rhizomatous stem (Fig. 8C–E) could be
regarded as a subform of caulescents, while those with leaves
with branched veins ascending from the base (Fig. 7J; group
(ii) of Hilliard & Burtt, 1971) are best described as a subform of
acaulescents, since they include rosulates and unifoliates (Fig. 2).
The evolution of growth form has been studied previously
in Streptocarpus and the habit found to be rather plastic (Möller
& Cronk, 2001a). In the expanded phylogenies presented here,
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Fig. 5. Leaves, seeds, and chromosome numbers of selected Streptocarpus species. A, Str. beampingaratrensis subsp. antambolorum; B, Str.
ionanthus subsp. grotei; C, Str. levis; D, Str. glandulosissimus; E, Str. stomandrus; F, Str. schliebenii; G, Str. parensis; H, Str. papangae; I, Str.
rexii; J, Str. wendlandii; K, Str. rexii; L, Str. vestitus; M, Str. vestitus, late prometaphase; N, Str. vestitus, prometaphase; O, Streptocarpus sp.
nov., early metaphase; P, Streptocarpus sp. nov., prometaphase. — Scale bars: A–J = 1 cm; K–L = 100 µm; M–P = 10 µm. Photos: A–J, M–P by
M. Möller, RBGE; K, L by F. Christie, RBGE.
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Fig. 6. Floral form (front and side view) of representative species of sections of Streptocarpus. A, sect. Trachystigma: Str. nobilis (open tube); B,
sect. Parasaintpaulia: Str. beampingaratrensis (small pouch); C, sect. Hova: Str. muscosus (small pouch); D, sect. Schizoboea: Str. thysanotus
(labellanthus); E, sect. Carnosifolii: Str. saxorum (keyhole); F, sect. Caulescentes: Str. caulescens (personate); G, sect. Saintpaulia: Str. ionanthus
(Saintpaulia-type); H, sect. Saintpaulia: Str. goetzeanus comb. nov. (Saintpaulia-type); I, sect. Lignostreptocarpus: Str. papangae (small pouch);
J, sect. Plantaginei: Str. hildebrandtii (small pouch); K, sect. Colpogyne: Str. ibityensis (small pouch); L–M, sect. Protostreptocarpus (small
pouch): L, Str. montanus; M, left to right: Str. bullatus, Str. montanus, Str. parensis, Str. schliebenii; N–O, sect. Streptocarpus: N, Str. rexii (open
tube); O, Str. dunnii (bird pollination). — Scale bars: 5 mm. All photos by M. Möller, RBGE.
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most morphs form separate subclades and have been found
useful in the classification of the Streptocarpinae, with the
exception of the unifoliates and rosulates, but sometimes with
parallelisms such as the Saintpaulia-rosette, that has evolved
once in Africa and once in Madagascar. The unifoliate form
shows a high level of homoplasy, and has most likely evolved
several times, at least twice in Clade II within the African and
the Madagascan acaulescents and once among the caulescents
of Clade I.
There is an enormous wealth of publications devoted to
the study of growth and development of Streptocarpus plants.
However, a detailed discussion of the historic (e.g., Crocker,
Fig. 7. Habit and growth form of representative species of sections of Streptocarpus subg. Streptocarpella and subg. Streptocarpus. A, sect.
Trachystigma: Str. nobilis (facultative caulesent); B, sect. Parasaintpaulia: Str. beampingaratrensis (rosette); C, sect. Hova: Str. muscosus (caulesent); D, sect. Schizoboea: Str. elongatus (caulesent); E, sect. Carnosifolii: Str. saxorum (caulesent); F, sect. Caulescentes: Str. caulescens (caulesent); G, sect. Saintpaulia: Str. ionanthus (rosette); H, sect. Saintpaulia: Str. goetzeanus comb. nov. (caulescent); I, sect. Lignostreptocarpus:
Str. papangae (caulesent); J, sect. Plantaginei: Str. hildebrandtii (acaulesent); K, sect. Colpogyne: Str. lanatus (rosulate); L, sect. Colpogyne: Str.
betsiliensis (rosulate). — Photos: A–C, E–H, J–L by M. Möller, RBGE; D by Matt Walters; I by L.Wilson, RBGE.
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1860; Jong, 1970) and evolutionary developmental work (e.g.,
Jong & Burtt, 1975; Harrison & al., 2005; Nishii & al., 2010)
would be beyond the scope of this work.
Inter- or infrageneric division. — In our phylogenetic analyses, we retrieved strong support for a subdivision within the
main clades and we found additional support from vegetative,
floral, seed morphological characters and basic chromosome
number or combinations thereof. These are used here to propose a new classification for the Streptocarpinae. The question
arises as to the rank at which to divide them. There are different
options to classify the Afro-Malagasy Streptocarpinae. These
options include:
Fig. 8. Habit and growth form of representative species of sections of Streptocarpus subg. Streptocarpus. A–F, sect. Protostreptocarpus: A–B, Str.
parensis (caulescent); C, Str. bullatus (creeping rhizomatous); D–E, Str. montanus (creeping rhizomatous; F, seedling of Str. schliebenii (caulescent); G, sect. Carnosifolii: seedling of Str. kirkii (caulescent); H–J, sect. Streptocarpus: H–I, Str. rexii (rosulate); J, Str. dunnii (unifoliate). — All
photos by M. Möller, RBGE.
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1.
Retaining all nine genera in subtribe Streptocarpinae
with no nomenclatural changes, but leaving a paraphyletic Streptocarpus.
2. Assigning Streptocarpus species to other genera with
which they form subclades in order to obtain monophyletic
genera. This would involve extreme splitting, with new
genera needing to be established from within paraphyletic
Streptocarpus subclades, resulting in over 20 (often small)
genera, sometimes even requiring the splitting of genera
such as Hovanella whose two species were not sister to
each other in any of our analyses. The morphological definition and differentiation between these “genera” would
also be weak. Raising subclades “a–l” to monophyletic
genera would still result in around a dozen genera, with the
recurring issue of difficulty in their delineation sometimes
on the basis of a single character, e.g., flower form (e.g.,
subclades “f ” and “g”), geography (e.g., subclades “c”, and
“k”), or basic chromosome number (subclades “d” and “k”)
and much nomenclatural change. For all other characters
used in the traditional classification of the Streptocarpinae
exceptions exist (e.g., Str. thysanotus does not have verruculose seeds, despite its placement in Clade I, and similar
exceptions exist in subclade “j” and “l”). For growth form
and floral form, exceptions exist in 9 out of the 12 subclades. Furthermore, the existence of morphologically
intermediate species, such as Str. semijunctus B.L.Burtt,
Str. stenosepalus B.L.Burtt or Str. tsimihetorum Humbert,
would, without additional data, only allow an uncertain or
no generic placement, further destabilizing the classification. The 12-genus approach would thus raise the level of
uncertainty in the generic placement of new species, that
has caused the misplacement of species in genera among
the Chinese Gesneriaceae, a situation that was resolved
by enlarging genera such as Oreocharis or Petrocodon
along phylogenetic boundaries and their redefinition using
synapomorphies (Möller & al., 2011b; Weber & al., 2011b).
Finally, the uncertainty of a phylogenetic placement of
species such as Str. exsertus and Str. capuronii, and the
insufficient knowledge of the morphology and affinities
of another seven species would result in unplaced species
at the genus level. Their existence in a taxonomic limbo
is not desirable.
3. Establishing two genera divided along Clade I and Clade II
species. The only clear synapomorphy for such a division
would be the basic chromosome number.
4. Sinking the genera with non-twisted fruits into Streptocarpus. This would result in a single genus, Streptocarpus, encompassing all Afro-Malagasy species with two
subgenera.
Nomenclatural considerations. — The alternatives of
subdividing the Streptocarpinae have different nomenclatural
consequences. In option 3, the two genera would comprise
Clade I, hitherto subg. Streptocarpella and Saintpaulia, and
Clade II, species of the former subg. Streptocarpus including
the generic type Str. rexii. The oldest genus name in Clade I
is Acanthonema (Hooker, 1862), and this name would have to
be used as a genus name for this clade under the International
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Code of Nomenclature (ICN, McNeill & al., 2012). However,
this is a small genus with two species, and would require 63
new combinations to accommodate the remaining species in
this genus. For the horticulturally important name Saintpaulia
to be used, a proposal to conserve this name over Acanthonema
(and any other older genus names) would be required. An argument for conserving Saintpaulia could be the fact that some
species placed in subg. Streptocarpella by Hilliard & Burtt
(1971), group (i), do resemble Saintpaulia very closely in vegetative habit, and that some Saintpaulia species of basal lineages
have prominent internodes and erect habit, and decussate leaf
arrangement (e.g., Fig. 7H, see also below). The globose-ovoid
non-twisted capsules of derived Saintpaulia species occur also
in Acanthonema, further blurring the morphological boundaries between Saintpaulia s.str. and other taxa in Clade 1.
The single-genus solution, Streptocarpus, in option 4,
could contain two subgenera, Streptocarpus and Streptocarpella, with expanded circumscriptions, and several sections,
one to accommodate Saintpaulia. Saintpaulia has a single synapomorphy in the tubeless flat corolla to support this section.
Of all the above options, this would keep the morphological
concept of Saintpaulia most intact, though at the rank of section. The plants are of significant horticultural importance,
and the name well established in the trade and the public. The
retention of the name at section rank would enable it to be used
in colloquial terms. The option of a single genus Streptocarpus
requires the fewest name changes, and provides a phylogenetictaxonomic framework across the genus with sufficient support
at the infrageneric level and we propose this option in our
classification.
Subdivision in Streptocarpus. — Fritsch (1894: 151) recognized three unranked groups in Streptocarpus, which Engler
(1921) later established as sections, i.e., sect. Unifoliati, sect.
Rosulati and sect. Caulescentes, the last of which Fritsch (1904:
158) raised to subg. Streptocarpella and combined the former
two into subg. Eustreptocarpus (now correctly Streptocarpus,
Art. 22 of the ICN, McNeill & al., 2012). Fritsch established
subg. Streptocarpella to comprise herbaceous caulescent species, while in subg. Streptocarpus he included the unifoliate
and rosulate species known at the time. Hilliard & Burtt (1971)
expanded the concept to accommodate some Madagascan
groups of species that did not fit well into either subgenus. Not
having seen nor studied these species in the field, they included
the shrubby species (group iii) and species with leaves in a basal
rosette with long petioles (group i) in subg. Streptocarpella,
and those with leaves in basal rosettes with veins ascending
from the leaf base (group ii) in subg. Streptocarpus although
they admitted to place these taxa for convenience rather than
conviction until more data became available. With the additional
information now available, we retain a subdivision into two
subgenera and establish a separate section for each of the three
Madagascan groups, but with the shrubby species falling into
subg. Streptocarpus. This is supported by cytology and perhaps
leaf base morphology, which is attenuate in subg. Streptocarpus
and cordate in subg. Streptocarpella (Fig. 5A–J, M–P). The
only further subdivision within subgenera was attempted by
Engler (1921) for subg. Streptocarpella, who recognized seven
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“Gruppen” which Hilliard & Burtt (1971) interpreted as series.
None of Engler’s Gruppen form a natural group and we refrain
from using these as section names. New sections have been
established here to accommodate the diversity now known in
the genus. A key to the subgenera and sections is provided.
In the new circumscription, the genus Streptocarpus
includes all species of Acanthonema, Colpogyne, Hovanella,
Linnaeopsis, Nodonema, Saintpaulia, Schizoboea and Trachystigma. These names are cited as synonyms under the section into which their types fall. Some of the former genus names
were adopted as sectional names for nomenclatural stability
(e.g., Colpogyne, Saintpaulia), some had to be used for priority reasons (e.g., Hova, Schizoboea, Trachystigma). Out of the
176 species in the newly defined Streptocarpus only 9 could
not be placed to section because of uncertainties in their phylogenetic placement or insufficient knowledge of their morphology. A table is provided listing the species in their respective
sections (Appendix 3).
TAXONOMY AND NOMENCLATURE
Key to subgenera and sections in Streptocarpus
1.
1.
x = 15 chromosomes; herbaceous caulescents with short
phyllomorphic phase of petiolated macrocotyledon (Fig.
6G), or unifoliates with non-twisted fruits .................
.................................... subg. Streptocarpella — 2
x = 16 chromosomes; woody caulescents, or herbaceous
caulescents with extended phyllomorphic phase of apetiolated macrocotyledon (Fig. 6F), or unifoliates and rosulates with twisted fruits ........ subg. Streptocarpus — 8
Streptocarpus subg. Streptocarpella (includes Acanthonema,
Hovanella, Saintpaulia, Schizoboea, Trachystigma)
2. Unifoliates with fruits not twisted, or facultative unifoliates with twisted fruits ......... sect. Trachystigma (“a”)
2. Rosette or caulescent herbs, fruits twisted or not twisted 3
3. Rosette herbs, or herbs with distinct internodes, corollas
flat ............................................................... 4
3. Caulescent herbs with tubular corollas .................... 5
4. Rosette or herbs with distinct internodes, corollas flat,
fruits not twisted .................... sect. Saintpaulia (“g”)
4. Rosette herbs, corollas tubular, fruits twisted ..............
.................................. sect. Parasaintpaulia (“b”)
5. Endemic to Madagascar and the Comoro Islands ..........
................................................. sect. Hova (“c”)
5. Endemic to Africa ............................................ 6
6. Small-flowered (≤ 10 mm corolla tube length), often with
forward directed lip and reduced upper lip .................
......................................... sect. Schizoboea (“d”)
6. Larger-flowered (≥ 15 mm corolla tube length) ........... 7
7. Succulent leaves with thick subepidermis, flowers not
± personate ...........................sect. Carnosifolii (“e”)
7. Leaves thinner, flowers ± personate ..........................
....................................... sect. Caulescentes (“f”)
Streptocarpus subg. Streptocarpus (includes the former Colpogyne, Linnaeopsis, Nodonema)
8.
8.
9.
9.
10.
10.
11.
11.
Unifoliates and rosulates, lateral leaf veins ascending from
the base of the lamina .............. sect. Plantaginei (“j”)
Shrubs, unifoliates, rosulates and caulescent herbs, lateral
leaf veins spreading at right angle to midrib ............. 9
Shrubs, anthers not coherent, some species with sclereids .......................... sect. Lignostreptocarpus (“h”)
Unifoliates, rosulates and caulescent herbs, anthers coherent, sclereids absent .......................................... 10
Unfoliates and rosulates, leaves with very dense and long
indumentum............................ sect. Colpogyne (“i”)
Unifoliates, rosulates and caulescent herbs, leaves glabrous
or slightly hairy................................................ 11
Caulescent herbs, with erect or rhizomatous stems, leaves
alternate or opposite, leaves without abscission zones,
seeds verruculose ......... sect. Protostreptocarpus (“k”)
Unifoliates or rosulates, leaves with abscission zones, seeds
mostly reticulate .................. sect. Streptocarpus (“l”)
Streptocarpus Lindl. in Bot. Reg. 14: t. 1173. 1828 – Type: Streptocarpus rexii (Bowie ex Hook.) Lindl. in Bot. Reg. 14:
t. 1173. 1828 ≡ Didymocarpus rexii Bowie ex Hook., Exot.
Fl. 3: 227. 1827.
Streptocarpus subg. Streptocarpella Fritsch, Keimpfl. Gesneriaceen: 158. 1904 ≡ Streptocarpus [unranked] Caulescentes
Fritsch in Engler & Prantl, Nat. Pflanzenfam. IV(3b): 151.
1894 (“§ 1.”) – Type (designated by Burtt in Notes Roy. Bot.
Gard. Edinburgh 21: 208. 1954): Streptocarpus caulescens
Vatke in Linnaea 43: 323. 1882.
See Figs. 5A–E, 6A–H, 7A–H.
Description. – In its broader circumscription, this subgenus comprises plants with x = 15 chromosomes and verruculose
seed ornamentation, and includes forms with the following
vegetative character combinations, (a) herbaceous caulescents
with decussate leaves at least in the seedling stage (for Saintpaulia see below) with a distinctly short phyllomorphic phase
of the macrocotyledon, and strongly petiolated macrocotyledon that is seldom larger than the subsequent cauline leaves
(Hilliard & Burtt, 1971: 118) (Fig. 8G), and (b) unifoliates with
non-twisted fruits.
Notes. – This subgenus contains ca. 65 taxa in 49 species and includes the former genera Acanthonema, Hovanella,
Saintpaulia, Schizoboea and Trachystigma (Clade I in Fig. 2A).
The fruit type in this subgenus is diverse, due to the inclusion of the five genera with non-twisted fruits. Acanthonema
and Saintpaulia have globose or ovoid fruits, while Hovanella,
Schizoboea and Trachystigma have slender cylindrical fruits.
The spiral leaf arrangement of leaves in mature Saintpaulia
species in the Sa. ionantha complex is the result of a shift in
leaf position and disjunct leaf development in a pair, superficially resembling anisophylly. Fritsch (1904: fig. 1) observed
that seedlings of Sa. ionantha H.Wendl. develop their first few
pairs of leaves in decussate arrangement and only later, due to
initial bending of the long petioles and shifts of the insertion
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points of leaves of a pair, the phyllotaxis becomes spiral. Hilliard & Burtt (1971) found a clear separation between their acaulescent Streptocarpus (2n = 32) and caulescent Streptocarpella
(2n = 30) species, but at that time chromosome counts for the
herbaceous caulescent Str. schliebenii (2n = 32; Milne, 1975)
and woody caulescent species (i.e., Str. papangae Humbert
and Str. suffruticosus Humbert both 2n = 32; Jong & Möller,
2000) were unavailable. So far, basic chromosome numbers for
species of this clade representing subg. Streptocarpella are consistent with x = 15 (24 species out of 49 now included). The seed
ornamentation is known for 38 taxa out of 65 in this subgenus.
All but three have verruculose seeds. Reticulate seed testa cell
surfaces were reported for Acanthonema strigosum Hook.f.
and Sa. inconspicua. The report for A. strigosum (BeaufortMurphy, 1983: 287, based on Mann 1948, P) is, on closer inspection of the published SEM image, perhaps based on the analysis
of an immature seed, since the specimen appears shrivelled
and not fully formed. Specimens observed in the MNHN herbarium in Paris (R. Letouzey 12925) clearly showed verruculose seed ornamentation for A. strigosum. The report for Sa.
inconspicua (Beaufort-Murphy, 1983: pl. 20F) came from a
herbarium specimen which we have now seen (Pócs T. & al.
6296, E); the testa sculpturing in Beaufort-Murphy (1983) is
more like that found in seeds of other genera, such as certain
Cyrtandra J.R.Forst. & G.Forst. (Beaufort-Murphy, 1983: pls.
6–9; Mühlbauer & Kiehn, 1997) or Didissandra C.B.Clarke
(e.g., D. elongata (Jack) C.B.Clarke or D. sprengelii C.B.Clarke;
Beaufort-Murphy, 1983: pl. 18I; Sontag & Weber, 1998). However, on reexamining seeds of Pócs T. & al. 6296, we find
that the seeds, while more strongly ribbed than other Saintpaulia seeds clearly possess verruculose seed testa cells. The
only species possessing seeds with a confirmed absence of a
bladder-like protrusion or papilla is Str. thysanotus. The phyllomorphic phase of seedlings of herbaceous caulescent Streptocarpus subg. Streptocarpella species is decidedly short and
their macrocotyledon cordate and strongly petiolated (Fig. 8G;
Jong, 1970; Hilliard & Burtt, 1971).
Streptocarpus sect. Trachystigma (C.B.Clarke) Mich.Möller
& Nishii, comb. nov. ≡ Trachystigma C.B.Clarke in
Candolle & Candolle, Monogr. Phan. 5: 131. 1883 ≡ Roettlera sect. Trachystigma (C.B.Clarke) Fritsch in Engler
& Prantl, Nat. Planzenfam. IV(3b): 148. 1894 – Type: Trachystigma mannii C.B.Clarke in Candolle & Candolle,
Monogr. Phan. 5: 131. 1883 ≡ Roettlera mannii (C.B.Clarke)
Fritsch in Engler & Prantl, Nat. Planzenfam. IV(3b): 148.
1894 ≡ Streptocarpus mannii (C.B.Clarke) Mich.Möller
& Nishii (see below). — Figs. 6A, 7A.
= Acanthonema Hook.f. in Bot. Mag. 88: t. 5339. 1862 – Type:
Acanthonema strigosum Hook.f. in Bot. Mag. 88: t. 5339.
1862 ≡ Streptocarpus strigosus (Hook.f.) Nishii & Mich.
Möller (see below).
= Streptocarpus [unranked] Brevilobati Engl. in Bot. Jahrb.
Syst. 57: 210. 1921 (“§”) – Type (designated by Hilliard
& Burtt, Streptocarpus: 120. 1971): Streptocarpus princeps
Mildbr. & Engl. in Bot. Jahrb. Syst. 57: 211. 1921 (= S. nobilis C.B.Clarke).
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Description. – Facultative unifoliate caulescents with
decussate leaf arrangement and twisted fruits, and unifoliates
with non-twisted fruits. Seeds verruculose. Distributed in western tropical Africa.
Notes. – Subclade “a” (Fig. 2A). Contains five species,
including Str. nobilis, which become unifoliate under unfavorable conditions such as short day length (Lawrence, 1943). Other
members of the section are Acanthonema strigosum, A. diandrum (Engl.) B.L.Burtt, and Trachystigma mannii, three obligate
unifoliate species. It appears that in these plants the transition
to the unifoliate state is irreversible, as opposed to Str. nobilis,
and the plants only produce one macrocotyledon as the sole
photosynthetic organ. Streptocarpus nobilis shares a relatively
large infundibuliform dark purple corolla with Acanthonema
strigosum. There are characters that differentiate Acanthonema
and Trachystigma from the former delimitation of Streptocarpus, not least the non-twisted fruits of both genera, but also the
ovoid-globose shape of the capsule, flattened toothed filaments,
unilateral disc and bilocular ovary for Acanthonema, and the
cylindrical capsule opening along the median line of the upper
carpel for Trachystigma (Hilliard & Burtt, 1971: 115; Weber
& Skog, 2007–). However, they share characters with Streptocarpus such as being unifoliate, having an acropetal succession
of inflorescences on the leaves, two coherent fertile stamens in
anterior position, and unilocular ovaries (Trachystigma only)
that place them firmly in the genus. Interestingly, Acanthonema
and Trachystigma share toothed filaments, and the phylogenetic work retrieves them as strongly supported sister taxa. This
suggests that the toothed filament has evolved prior to a split
between the two taxa, and that characters pertaining to the ovary
and fruit morphology have evolved more rapidly. This section
is essentially of western tropical African origin (Cameroon,
Fernando Po Island off the west coast of Cameroon), with Str.
nobilis the most widespread (from Gambia, through Cameroon
to Central African Republic).
Streptocarpus diandra (Engl.) Nishii & Mich.Möller, comb.
nov. ≡ Carolofritschia diandra Engl. in Bot. Jahrb. Syst.
26: 362. 1899 ≡ Acanthonema diandrum (Engl.) B.L.Burtt
in Bull. Mus. Natl. Hist. Nat., B, Adansonia, ser. 4,
3(4): 416. 1982 (“1981”) – Syntypes: CAMEROUN. Um
Lolodorf, März 1895, Staudt 118 (B, destr.; BM barcode
BM001122850!); zwischen Lolodorf und Carantschiamasdorf, Mai 1907, Zenker 1381 (B, destr.; BM barcode
BM000984380!, E barcode E00632283!, P barcode
P00057466!).
Streptocarpus mannii (C.B.Clarke) Nishii & Mich.Möller,
comb. nov. ≡ Trachystigma mannii C.B.Clarke in Candolle & Candolle, Monogr. Phan. 5: 131, pl. 14. 1883 ≡
Roettlera mannii (C.B.Clarke) Fritsch in Engler & Prantl,
Nat. Pflanzenfam. IV(3b): 148. 1894 ≡ Didymocarpus mannii (C.B.Clarke) Wonisch in Sitzungsber. Kaiserl. Akad.
Wiss., Wien. Math.-Naturwiss. Cl., Abt. 1, 118: 483. 1909
– Holotype: Africa occidentalis tropica. Sinus Corisco,
in montibus Sierra del Crystal, Mann 1682 (K barcodes
K000378484! & K000378485!).
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Streptocarpus strigosus (Hook.f.) Nishii & Mich.Möller, comb.
nov. ≡ Acanthonema strigosum Hook.f. in Bot. Mag. 88: pl.
5339. 1862 – Neotype (designated here): CAMEROUN,
Fernando Po, epiphyte on rocks and trees, corolla dark
purple, calyx brown[red], 1–2000 feet, Nov 1860, Mann
569 (K barcode K000378483!).
Note. – In the protologue, Hooker (1862) refers only to
“living plants were sent home by M. Gustav Mann from an
elevation of 4000–5000 feet in the plains of Fernando Po, where
it grows on rocks and epiphytically on trees.”
Streptocarpus sect. Parasaintpaulia Mich.Möller & Nishii,
sect. nov. – Type: Streptocarpus beampingaratrensis
Humbert in Adansonia, sér. 2, 7: 287. 1967. — Figs. 6B, 7B.
Description. – Leaves in basal rosette, long petioles sharply
marked off from the (sub-)orbicular blades, axillary inflorescence, small short wide corollas and relatively short spirally
twisted fruits. Seeds verruculose. Distributed in Madagascar.
Notes. – Subclade “b” (Fig. 2A). Six taxa in three species.
The section is named for their similarity in vegetative morphology with African Saintpaulia. This section corresponds to
group (i) of Hilliard & Burtt (1971), for which they recognized
a close affinity to the African Saintpaulia, but which can be
distinguished easily by their tubular corolla and twisted fruits.
A section that includes polyploids, all with a basic number of
x = 15 (Fig. 2).
Streptocarpus sect. Hova (C.B.Clarke) Mich.Möller & Nishii,
comb. nov. ≡ Didymocarpus sect. Hova C.B.Clarke in Candolle & Candolle, Monogr. Phan. 5: 108. 1883 ≡ Hovanella
A.Weber & B.L.Burtt in Beitr. Biol. Pflanzen 70: 333.
1998 – Type: Didymocarpus madagascaricus C.B.Clarke
in Candolle & Candolle, Monogr. Phan. 5: 108. 1883 ≡
Streptocarpus madagascaricus (C.B.Clarke) Christenh.
in Phytotaxa 46: 8. 2012. — Figs. 6C, 7C.
= Streptocarpus [unranked] Longiflori Engl. in Bot. Jahrb.
Syst. 57: 216. 1921 (“§”) – Type (designated by Hilliard
& Burtt, Streptocarpus: 120. 1971): Streptocarpus hilsenbergii R.Br. in Bennett, Pl. Jav. Rar.: 119. 1840.
Description. – Herbaceous caulescent plants with decussate leaves, axillary inflorescences, mostly with small,
short-tubed pouch flowers. Seeds verruculose. Distributed
in Madagascar.
Notes. – Subclade “c” (Fig. 2A). Seventeen taxa in 13 species plus 4 varieties. Species in this section have the strong
tendency to possess small corollas (mostly 5–10 mm) with
short tubes (4–8 mm), which are the ancestral type in the
genus (Hughes & al., 2006). The exception is Str. hilsenbergii R.Br. with the largest flower of all Madagascan and
Comores Gesneriaceae, although within this species flower
size varies greatly (25–45 mm corollas with 20–30 mm tubes),
indicating an inherent flexibility in these characteristics. Its
phylogenetic position in the middle of this section indicates
an isolated co-speciation event with a long-tongued pollinator (Potgieter & Edwards, 2005). Section Hova includes the
erstwhile genus Hovanella. The main character distinguishing it from Streptocarpus s.str. is the non-twisted fruit. The
two species included here have distinctive fruit orientations
and dehiscence. In H. madagascarica (C.B.Clarke) A.Weber
& B.L.Burtt they are plagiocarpic, held horizontally at a ±
right angle to the pedicel and dehisce follicle-like, only along
the dorsal suture, whereas in H. vestita (Baker) A.Weber
& B.L.Burtt they are in line with the pedicel and dehisce
along the dorsal and ventral suture. It is no surprise to find
that they were not sister species in the phylogenetic trees, and
that the losses of fruit twist in Hovanella represent parallel,
independent events. Seven chromosome counts are available
for species in this section, most with 2n = 30 (Möller & Pullan,
2015–). The one of 2n = 28 by Kiehn for Hovanella madagascarica should be treated with caution since the count is based
on field-fixed flower bud material. Preliminary counts exist
for two further taxa, H. vestiata and a putative new species,
with 2n = 30 (Fig. 5M–P).
Streptocarpus sect. Schizoboea (Fritsch) Mich.Möller &
Nishii, comb. nov. ≡ Roettlera sect. Schizoboea Fritsch
in Engler & Prantl, Nat. Planzenfam. Nachtr. 1: 300. 1897
≡ Schizoboea (Fritsch) B.L.Burtt in Notes Roy. Bot. Gard.
Edinburgh 33: 266. 1974 – Type: Didymocarpus kamerunensis Engl. in Bot. Jahrb. Syst. 18: 79. 1894 ≡ Schizoboea
kamerunensis (Engl.) B.L.Burtt in Notes Roy. Bot. Gard.
Edinburgh 33: 266. 1974 ≡ Streptocarpus kamerunensis
(Engl.) Christenh. in Phytotaxa 46: 7. 2012. — Figs. 6D, 7D.
= Streptocarpus [unranked] Ventricosi Engl. in Bot. Jahrb.
Syst. 57: 210. 1921 (“§”) – Type: Streptocarpus gonjaensis
Engl. in Bot. Jahrb. Syst. 57: 210. 1921.
= Streptocarpus [unranked] Elongati Engl. in Bot. Jahrb. Syst.
57: 213. 1921 (“§”) – Type (Art. 22.6): Streptocarpus elongatus Engl. in Bot. Jahrb. Syst. 18: 76. 1894.
Description. – Short-lived herbaceous caulescent plants
with decussate leaf arrangement, with mostly long-pedunculate
axillary inflorescences and small, strongly bilabiate flowers.
Distributed in tropical Africa.
Notes. – Subclade “d” (Fig. 2A). A section with six species,
which forms a link between species in East and West Africa
(Hilliard & Burtt, 1975). They are characterized by a forward
directed lower lip and a reduced upper lip of the corolla, the
labellanthus-type (Fig. 7D). They are annuals with a sappy
stem and long-pedunculate axillary inflorescences. The section
includes the erstwhile genus Schizoboea. Characters shared
between this genus and Streptocarpus subg. Streptocarpella
include axillary few-flowered cymes, small corolla, two fertile stamens in anterior position (Burtt, 1974), and verruculose
seeds (Beaufort-Murphy, 1983). Initially described in Didymocarpus (Fritsch, 1894), on the basis of the straggling creeping
habit and subglabrous entire leaves, Burtt (1974) suggested that
the true affinity of the erstwhile Schizoboea does not lie with
Didymocarpus Wall., but with the little-known African Saintpaulia inconspicua, and in that he was not far off the mark. Our
molecular data suggest that both Saintpaulia and Schizoboea
have evolved from within Streptocarpus and belong in subg.
Streptocarpella (Fig. 2). Schizoboea is sister to Str. elongatus
with which it shares a bilabiate white corolla. The straight,
sometimes slightly falcate capsule, dehiscing into 4 valves and
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the lateral, sclerified vascular bundles, are seemingly another
isolated evolutionary event that Streptocarpus is prone to, but
is by no means unique in the Gesneriaceae.
Streptocarpus sect. Carnosifolii Mich.Möller & Nishii, sect.
nov. – Type: Streptocarpus saxorum Engl. in Bot. Jahrb.
Syst. 19: 154. 1894 (“1895”). — Figs. 6E, 7E.
= Streptocarpus [unranked] Pauciflori Engl. in Bot. Jahrb.
Syst. 57: 216. 1921 (“§”) – Type: Streptocarpus saxorum
Engl. in Bot. Jahrb. Syst. 57: 216. 1921.
Description. – Long-lived caulescent herbs or shrubs with
a sometimes woody stem base and succulent decussate leaves,
axillary inflorescences and twisted fruits. Seeds verruculose.
Distributed in tropical East Africa.
Notes. – Subclade “e” (Fig. 2A). A group of six species
with a distribution in Tanzania and Kenya, usually with large
corollas, but very divergent floral types, ranging from keyhole
flowers in Str. saxorum (Fig. 6E), strongly bilabiate flowers in
Str. stomandrus, to almost campanulate corollas in Str. kirkii.
The species are often long-lived plants that possess relatively
fleshy leaves with an extremely thick hypodermis of up to
seven layers in Str. saxorum (Sahasrabudhe, 1970).
Streptocarpus sect. Caulescentes (Fritsch) Engl. in Bot. Jahrb.
Syst. 57: 203. 1921 ≡ Streptocarpus [unranked] Caulescentes Fritsch in Engler & Prantl, Nat. Pflanzenfam.
IV(3b): 151. 1894 (“§ 1.”) ≡ Streptocarpus [unranked] Porrectilobati Engl. in Bot. Jahrb. Syst. 57: 214. 1921 (“§”) –
Type (designated by Hilliard & Burtt, Streptocarpus: 120.
1971): Streptocarpus caulescens Vatke in Linnaea 43: 323.
1882. — Figs. 6F, 7F.
Description. – Herbaceous caulescent plants with decussate leaves, axillary inflorescences, enantiostyly (the deflection of the style either to the left or right side of the floral axis)
with personate corollas and twisted fruits. Seeds verruculose.
Distributed in tropical East Africa.
Notes. – Subclade “f” (Fig. 2A). The species in this section
were placed in “Gruppe” Porrectilobati by Engler (1921), but
for reasons of priority the name Caulescentes has to be used
(McNeill & al., 2012). The six species in this section have a
very similar corolla form, in which the bend of the lower lip
more (Str. holstii Engl.) or less (Str. pallidiflorus C.B.Clarke,
Str. inflatus B.L.Burtt) closes the corolla tube (Fig. 6F).
Streptocarpus sect. Saintpaulia (H.Wendl.) Mich.Möller &
Haston, comb. & stat. nov. ≡ Saintpaulia H.Wendl. in
Gartenflora 42: 321, pl. 1391, fig. 66. 1893 – Type: Saintpaulia ionantha H.Wendl. in Gartenflora 42: 321. 1893 ≡
Petrocosmea ionantha (H.Wendl.) Rodigas in Ill. Hort. 42:
108. 1895 ≡ Streptocarpus ionanthus (H.Wendl.) Christenh.
in Phytotaxa 46: 6. 2012. — Figs. 6G–H, 7G–H.
Description. – Herbaceous caulescent plants with leaves in
a basal rosette (opposite as seedlings), or decussate with distinct
internodes, long petioles, axillary inflorescences, flower with
flat corollas and enantiostyly. Fruits ovoid to linear-cylindric,
not twisted. Seeds verruculose. Distributed in Tropical East
Africa.
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Notes. – Subclade “g” (Fig. 2A). This section represents
the former genus Saintpaulia which has received various treatments resulting in 21 (Burtt, 1958a, 1964), 8 (Darbyshire, 2006)
or 6 species (Christenhusz, 2012). Our phylogenetic analysis
revealed 10 distinct genetic entities which we treat as species
in sect. Saintpaulia. Saintpaulia ulugurensis Haston was previously sunk into Sa. ionantha subsp. grandifolia (B.L.Burtt)
I.Darbysh. (Darbyshire, 2006) (= Str. ionanthus subsp. grandifolius (B.L.Burtt) Christenh.; Christenhusz, 2012), although
they are distant both phylogenetically (Figs. 2A, 4) and geographically. Morphologically, they can be differentiated by the
decussate leaf arrangement, distinct internodes and bicolored
flowers of the former (Haston, 2009a), and spiral leaf arrangement, rosette habit and deep violet flowers of the latter (Burtt,
1958a). Consequently, we give it species status in Streptocarpus. Saintpaulia watkinsii was reduced to Str. afroviola subsp.
watkinsii (Haston) Christenh. (Christenhusz, 2012). However, it
is distinct from Str. afroviola in having violet-blue colouration
on the lower lobes of the otherwise white flower whereas the
flowers of Str. afroviola are white with the two upper lobes
mauve to blue. In addition, the upper leaf surface of Str. afroviola has sparse, long, erect to arching hairs, rather than the
more dense, short, appressed hairs of Str. watkinsii. Moreover, the molecular data suggest that its phylogenetic affinity
is with Str. inconspicuus (Fig. 4), from which it differs by the
short appressed hairs on the upper leaf surface as opposed to
the nearly glabrous upper leaf surfaces of Str. inconspicuus
(B.L.Burtt) Christenh. (Burtt, 1958a; Haston, 2009b). We raised
the subspecies to species rank in Streptocarpus. Saintpaulia
brevipilosa was previously included in Sa. ionantha subsp.
velutina (B.L.Burtt) I.Darbysh. (Darbyshire, 2006) (= Str. ionanthus subsp. velutinus (B.L.Burtt) Christenh.; Christenhusz,
2012), although the two species are geographically and phylogenetically distant (Fig. 4). They also differ in indumentum
and leaf texture from each other (Burtt, 1958a, 1964), thus we
raise them to species level in Streptocarpus. Saintpaulia nitida
was included within Sa. ionantha as Sa. ionantha subsp. nitida
(B.L.Burtt) I.Darbysh. (Darbyshire, 2006) (= Str. ionanthus
subsp. nitidus (B.L.Burtt) Christenh.; Christenhusz, 2012). The
two species occur in separate mountain ranges, at different altitudes and are phylogenetically distant (Fig. 4), and the former
has short appressed hairs which are not present in the latter
(Burtt, 1958a). We therefore raise Sa. nitida to species rank as
Str. nitidus in Streptocarpus (see below). The oversimplified
taxon boundaries in Darbyshire (2006) and Christenhusz (2012)
have led to unforeseen consequences in a recent biogeographic
study of Saintpaulia (Dimitrov & al., 2012), extending the distribution of Sa. ionantha across the entire range of Saintpaulia
skipping only the Uluguru Mts., and resulting in individual
taxa such as Sa. ionantha subsp. nitida or Sa. ionantha subsp.
velutina each having disjunct distributions in the Nguru Mts.
and East Usambara Mts. Given published molecular data, it is
more likely that the plants designated cf. subsp. nitida and cf.
subsp. velutina in Lindqvist & Albert (1999) belong to new taxa.
Previous studies (Möller & Cronk, 1997b; Lindqvist & Albert,
2001; Dimitrov & al., 2012) and the results presented here show
that there is no strongly supported phylogenetic structure and
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discernible taxon monophyly, except perhaps for Sa. rupicola, in the Sa. ionantha complex (= Str. ionanthus complex
in Fig. 4). Putative ancestral characters of growth form and
leaf indumentum are very homogeneous within the mountain
ranges (Fig. 4), but are mixed and combined (e.g., leaf indumentum states) in the Sa. ionantha complex, suggestive of past
hybridization events. This would explain the mixed phylogenetic signals from 5S NTS nrDNA spacer data and the base
polymorphisms detected in the ITS data (Möller & Cronk,
1997a, present study). In such a hybridization scenario, morphological heterogeneity would have resulted and taxon boundaries
would have been blurred, perhaps pointing to a reconsideration
of the need for the diverse subspecies and varieties in the Sa.
ionantha complex.
woody and herbaceous caulescents with decussate or alternate
leaf arrangement, unifoliates and rosulates, with a long seedling phyllomorphic phase and apetiolar macrocotyledons with
attenuate bases (Fig. 8F) and mostly twisted fruits. It represents
an extension of the subg. Streptocarpus concept of Hilliard
& Burtt (1971). While the subgenus clearly forms a separate
clade with distinct subclades, delimitation of all sections by
growth form alone (as in the woody Madagascan species for
example), such as herbaceous caulescents, rosulates or unifoliates, is not possible, and supporting characters from seeds,
leaves and flowers are required.
Notes. – This subgenus contains ca. 138 taxa in 119 species.
Includes the former Colpogyne, Linnaeopsis and Nodonema.
(Clade II in Fig. 2B).
Streptocarpus brevipilosus (B.L.Burtt) Mich.Möller & Haston,
comb. nov. ≡ Saintpaulia brevipilosa B.L.Burtt in Notes
Roy. Bot. Gard. Edinburgh 25: 193. 1964 – Holotype: TANZANIA: Morogoro Dist., Nguru Mts., Lulaga, Punter ref.
59/4350, cultivated at RBG Edinburgh, C.3827 (E barcode
E00009668!).
Streptocarpus sect. Lignostreptocarpus Mich.Möller & Bellstedt, sect. nov. – Type: Streptocarpus papangae Humbert
in Adansonia, sér. 2, 7: 293. 1967. — Figs. 6I, 7I.
Description. – Woody shrubs with decussate leaves, axillary inflorescences, small pouch flowers, short filaments arising at the base of the corolla with non-coherent anthers, seeds
long and pointed at both ends and distinctly larger (0.9–1.2 mm)
than in other species (0.3–0.7 mm; Jong & al., 2012). Distributed
in Madagascar.
Notes. – Subclade “h” (Fig. 2B). Seven species with 10 taxa.
This section is most readily defined by the extensive woodiness
of the stems and is equivalent to group (iii) of Hilliard & Burtt
(1971). There is not necessarily a link between the peculiarly
short filaments and the non-coherent anthers as species with
coherent anthers can have short filaments, such as Str. levis
in sect. Hova (Hilliard & Burtt, 1971; Jong & al., 2012). Section Lignostreptocarpus is sister (with strong support) to the
next section, a group of unifoliates and rosulates also from
Madagascar. This again highlights the flexible morphology (in
evolutionary terms) of Streptocarpus. The woody caulescents
in sect. Lignostreptocarpus represent a break from the correlation between caulescence and basic chromosome number
(x = 15) as previously described for subg. Streptocarpella. The
two species for which chromosomes have been counted to date
(Str. papangae and Str. suffruticosus) both have 2n = 32 (Jong
& Möller, 2000; Möller & Kiehn, 2004).
Streptocarpus nitidus (B.L.Burtt) Mich.Möller & Haston,
comb. nov. ≡ Saintpaulia nitida B.L.Burtt in Notes Roy.
Bot. Gard. Edinburgh 22: 564. 1958 ≡ Saintpaulia ionantha subsp. nitida (B.L.Burtt) I.Darbysh. in Beentje &
Ghazanfar, Fl. Trop. E. Africa: 69. 2006 ≡ Streptocarpus
ionanthus subsp. nitidus (B.L.Burtt) Christenh. in Phytotaxa 46: 7. 2012 – Holotype: TANZANIA. Morogoro
District: Nguru Mts., near Morogoro, Aug 1959, Harvey
Cox ref. A, cultivated at RBG Edinburgh, C1557 (E sheet
1 barcode E00009655!, sheet 2 barcode E00009656!; isotypes: K barcode K000378659!, P barcode P00568966!).
Streptocarpus ulugurensis (Haston) Haston, comb. nov. ≡
Saintpaulia ulugurensis Haston in Bot. Mag. 26(3): 273
(–276, pl. 656, fig.). 2009 – Holotype: Cultivated at RBGE
from wild material collected from Tanzania, Morogoro,
Uluguru Mountains, Haston 246 (NHT, n.v.; isotypes: E
barcode E00708368!, K, n.v.).
Streptocarpus watkinsii (Haston) Haston, comb. nov. ≡ Saintpaulia watkinsii Haston in Bot. Mag. 26(3): 278 (277–280,
pl. 657, fig. 1). 2009 ≡ Streptocarpus afroviola var. watkinsii (Haston) Christenh. in Phytotaxa 46: 6. 2012 – Holotype: TANZANIA. Morogoro, Uluguru Mts., Haston 99
collected with John Mejissa (NHT, n.v.; isotype: E barcode
E00201489!).
Streptocarpus subg. Streptocarpus [Streptocarpus subg.
Eustreptocarpus Fritsch, Keimpfl. Gesneriaceen: 158.
1904, nom. illeg. (Art. 21.3 & 22.2)] – Type: Streptocarpus
rexii (Bowie ex Hook.) Lindl. in Bot. Reg. 14: pl. 1173. 1828
≡ Didymocarpus rexii Bowie ex Hook., Exot. Fl. 3: t. 227.
1827. . — Figs. 5F–J, 6I–O, 7I–L, 8A–J.
Description. – Species in this enlarged subgenus share a
basic chromosome number of x = 16 (41 counts), and include
Streptocarpus sect. Colpogyne Mich.Möller & Bellstedt,
comb. & stat. nov. ≡ Colpogyne B.L.Burtt in Fl. Madagasc., Fam. 180: 150. 1971 – Type: Colpogyne betsiliensis
(Humbert) B.L.Burtt in Fl. Madagasc., Fam. 180: 150. 1971
≡ Streptocarpus betsiliensis Humbert in Adansonia, sér.
2, 7: 289. 1967. — Figs. 6K, 7K–L.
Description. – Unifoliates and rosulates, leaves with dense,
long, woolly indumentum, small flowers and deep invagination
(colpus) in the ovary walls. Seeds verruculose. Distributed in
Madagascar.
Notes. – Subclade “i” (Fig. 2B). A section with five species including the former genus Colpogyne. Colpogyne was
initially included in Streptocarpus by Humbert (1971), but then
established as a separate genus on the basis of its short and
non-twisted fruits (Hilliard & Burtt, 1971: 115). There are other
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features that led to its exclusion from Streptocarpus such as the
deep invagination in the ovary walls; however, this is also found
in Str. itremensis, Str. lanatus and Str. ibityensis (Humbert,
1971: fig. IX), the latter a species which Hilliard & Burtt (1971)
see as most similar in vegetative characters to Colpogyne, and
it is indeed this species that is phylogenetically most closely
related to Colpogyne betsiliensis (≡ Str. betsiliensis), hence its
inclusion in sect. Colpogyne. The straight fruit in Str. betsiliensis is another case of loss of fruit twist.
Streptocarpus sect. Plantaginei Mich.Möller & Bellstedt, sect.
nov. – Type: Streptocarpus hildebrandtii Vatke in Linnaea
43: 324. 1882. — Figs. 6J, 7J.
Description. – Unifoliates and rosulates with leaves in a
basal rosette, often attenuate at the base (petiole ill-defined),
veins ascending from the base and branched. Distributed in
Madagascar.
Notes. – Subclade “j” (Fig. 2B). This section includes
14 species and essentially represents group (ii) of Hilliard
& Burtt (1971: 104). It includes a few species with less conspicuous ascending venation and petiolated leaves. It is morphologically the most complex group. This is also seen in the seed
ornamentation of its members, being reticulate in most, but
verruculose in Str. brevistamineus Humbert and Str. lokohensis
Humbert. The occurrence of unifoliates and rosulates in this
Madagascan subclade is similarly mixed. This highlights again,
as previously observed by Fritsch (1904), Hilliard & Burtt (1971)
and Möller & Cronk (2001a) for African members, that transitions between the two growth forms have occurred frequently.
Intermediate forms also exist, such as Str. hildebrandtii with
two similarly sized leaves (Fig. 7J), one representing the macrocotyledon, the other a true leaf (Humbert, 1971: fig. II1–3).
Interestingly, the three chromosome counts for members of this
section all reveal polyploidy of the basic number x = 16, with
Str. perrierii Humbert being tetraploid, Str. variabilis being
hexaploid and Str. hildebrandtii octoploid (Möller & Pullan,
2015–).
Streptocarpus sect. Protostreptocarpus Mich.Möller & Bellstedt, sect. nov. – Type: Streptocarpus montanus Oliv.
in Trans. Linn. Soc. London, Bot. 2: 344. 1887. — Figs.
6L–M, 8A–G.
= Linnaeopsis Engl. in Bot. Jahrb. Syst. 28: 483. 1900 – Type:
Linnaeopsis heckmanniana Engl. in Bot. Jahrb. Syst.
28: 483. 1900 ≡ Streptocarpus heckmannianus (Engl.)
I.Darbysh., Fl. Trop. E. Africa, Gesner.: 33. 2006.
= Nodonema B.L.Burtt in Bull. Mus. Natl. Hist. Nat., B,
Adansonia, sér. 4, 3(4): 415. 1982 (“1981”) – Type: Nodonema lineatum B.L.Burtt in Bull. Mus. Natl. Hist. Nat.,
B, Adansonia, sér. 4, 3(4): 416. 1982 (“1981”) ≡ Streptocarpus lineatus (B.L.Burtt) Mich.Möller & M.Hughes
(see below).
Description. – A group of species of variable habit (rhizomatous perennial herbs, or caulescents with alternate or
opposite leaves) with extended seedling phyllomorphic phase.
Small, more or less campanulate flowers. Seeds verruculose.
Distributed in tropical Africa.
1264
Notes. – Subclade “k” (Fig. 2B). A group of 10 taxa in
8 species with a narrow distribution in Kenya and Tanzania,
and one outlier in Nigeria and Cameroon (Nodonema lineatum). The section includes a variety of vegetative forms and can
be regarded as a transition from caulescents to acaulescents.
It includes the previously recognized genera Linnaeopsis and
Nodonema. The closeness of Linnaeopsis to Streptocarpus
is emphasized by the tightly twisted fruits of one species,
Str. subscandens (B.L.Burtt) I.Darbysh. (L. subscandens
B.L.Burtt; Weigend, 2000). It had not been described in Streptocarpus since the specimens available for the description had
no intact fruits (Burtt, 1958b). The alternate leaf arrangement,
verruculose seeds, and inflorescences borne in series on the
petiole of the former Linnaeopsis and Nodonema (Weigend,
2000) are all reminiscent of Streptocarpus, and with view
to leaf arrangement, particularly of Str. bullatus. In fact this
species was found to be the closest relative of the Linnaeopsis samples included here. Another strongly supported sister
species pair found in sect. Protostreptocarpus comprises the
morphologically very similar caulescents Str. parensis and Str.
schliebenii. Streptocarpus schliebenii represents another break
from the correlation of herbaceous caulescence and x = 15
found in subg. Streptocarpella. Morphologically, the two species can readily be distinguished from other species in subg.
Streptocarpella by differences in seedling development. This
includes a long lag phase of the plumule development during
which the macrocotyledon continues to expand (Fig. 8F), by
the activity of the basal meristem, similar to the well-studied
rosulate Str. rexii (Jong, 1970; Nishii & al., 2010). Streptocarpus montanus represents a further step towards the typical
rosulate morphotype, as represented by the type Str. rexii. It
has 2n = 32 chromosomes (Ratter & Milne, 1970), and, apart
from the verruculose seeds, its leaf insertion is much looser.
The “rhizome” only superficially resembles that of “typical”
rosulates of sect. Streptocarpus (below) such as Str. primulifolius Gand., and differs by the fusion of the petiolodes (structures with functions of stem and petiole sensu Jong & Burtt,
1975) in their entirety to form the rhizome stock, whereas in
Str. primulifolius it is only the bases of successive petiolodes
that do so. The effect is that in Str. montanus old inflorescences remain attached to the rhizome and appear axillary in
origin, similar to caulescent species (Hilliard & Burtt, 1971).
Nodonema could not be included in our molecular analysis.
The verruculose seeds (Weigend, 2000), pair-flowered axillary cymes, few-flowered cymes with small infundibuliform
bilabiate white corollas, with purple lines in the throat and two
fertile stamens in anterior position (Weber & Skog, 2007–),
link the genus to Streptocarpus. The short, subglobose fruit
may suggest a closer relationship with Acanthonema or Saintpaulia, but the rhizomatous habit with apparently alternate
leaves does not occur in subg. Streptocarpella, and resembles
Str. montanus in this section.
Streptocarpus lineatus (B.L.Burtt) Mich.Möller & M.Hughes,
comb. nov. ≡ Nodonema lineatum B.L.Burtt in Bull. Mus.
Natl. Hist. Nat., B, Adansonia, sér. 4, 3(4): 416. 1982 (“1981”)
– Holotype: CAMEROUN. Crête du Nta Ali (1266 m),
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TAXON 64 (6) • December 2015: 1243–1274
Nishii & al. • Streptocarpus redefined
entre cotes 1009 et 1202, 30 km SE Mamfe, Letouzey 13873
(P barcode P00057467!; isotypes: E barcode E00632294!, P
barcodes P00057468! & P00057469!, YA barcode 0032423,
n.v.).
Streptocarpus sect. Streptocarpus – Type: Didymocarpus rexii
Bowie ex Hook. in Exot. Fl. 3: pl. 227. 1827 ≡ Streptocarpus rexii (Bowie ex Hook.) Lindl. in Bot. Reg. 14: pl.
1173. 1828.
See Figs. 6N–O, 8H–J.
= Streptocarpus sect. Rosulati (Fritsch) Engl. in Bot. Jahrb.
Syst. 57: 217. 1921 ≡ Streptocarpus [unranked] Rosulati
Fritsch in Engler & Prantl, Nat. Pflanzenfam. IV(3b): 151.
1894 (“§ 2.”) – Type (designated by Burtt in Notes Roy.
Bot. Gard. Edinburgh 21: 208. 1954): Streptocarpus rexii
(Bowie ex Hook.) Lindl. in Bot. Reg. 14: pl. 1173. 1828.
= Streptocarpus sect. Unifoliati (Fritsch) Engl. in Bot. Jahrb.
Syst. 57: 217. 1921 ≡ Streptocarpus [unranked] Unifoliati
Fritsch in Engler & Prantl, Nat. Pflanzenfam. IV(3b): 151.
1894 (“§ 3.”) – Type (designated by Burtt in Notes Roy. Bot.
Gard. Edinburgh 21: 208. 1954): Streptocarpus polyanthus
Hook. in Bot. Mag. 81: pl. 4850. 1855.
Description. – Rosulate, plurifoliate and unifoliate herbs,
with greatly extended seedling phyllomorphic phase, the formation of abscission zones, and inflorescences usually at the
base of the lamina. Seeds reticulate, with few exceptions. Distributed in Africa.
Notes. – Subclade “l” (Fig. 2B). The largest section with
85 species (99 taxa) and a distribution from Ethiopia in the
north to the Western Cape Province in South Africa in the
south. This section includes a wide range of floral forms and
sizes with a trend of a north to south speciation pattern (Möller
& Cronk, 2001b) and an accompanying increase in corolla
size southwards (Hughes & al., 2006). Section Streptocarpus
includes only acaulescent species, with the exceptions of Str.
decipiens and Str. fanniniae Harv. ex C.B.Clarke which have
unusual growth forms. Streptocarpus fanniniae is branching
and has extremely long petiolodes, but does not possess a shoot
apical meristem or axis and is acaulescent (Jong & Burtt, 1975).
Streptocarpus decipiens was initially considered to belong to
subg. Streptocarpella by Hilliard & Burtt (1971: 227), because
of the axillary inflorescences and verruculose seeds. However,
basal leaves, presence of abscission zone, origin of the “stem”
from the base of the petiolode and attenuate leaf base place it in
sect. Streptocarpus. Abscission zones are thus far only reported
for members of sect. Streptocarpus and have not been observed
in any of the Madagascan acaulescent species (Hilliard & Burtt,
1971: 118; M. Möller, pers. obs.), here placed in sect. Plantaginei
and sect. Colpogyne. Within sect. Streptocarpus, a clear evolutionary trend from verruculose to reticulate seeds is observed.
Only a few species on early-branching lineages in sect. Streptocarpus possess verruculose seed ornamentation, such as Str.
nimbicola, Str. hirtinervis, Str. decipiens, and Str. galpinii (a
few populations only). Streptocarpus dunnii, in a more nested
position, also has verruculose seeds. Growth form and other
characters (e.g., presence of abscission zone) firmly place all
these species in this section.
CONCLUSIONS
The present work is based significantly on the enormous
body of work and careful observations by O.M. Hilliard,
B.L. Burtt and K. Jong and draws on their many publications.
Since then, more species have been described in Streptocarpus, but the generic concept has changed little until recently,
principally including Afro-Malagasy taxa with twisted fruits.
The reliance on this character led Burtt (in Humbert, 1971)
to remove Colpogyne from Streptocarpus. With the advent
of molecular methods it has become clear that the evolution
of morphology is more complex, and that some characters,
such as twisted fruits, may not evolve often but can be lost
relatively easily (Möller & al., 2009, 2011a; Puglisi & al., 2011).
Burtt (1984) himself gave preference to indumentum characters over fruit characters and redefined the genus Paraboea to
include species with and without twisted fruits and was shown
to be correct in his approach (Puglisi & al., 2011). This is the
broad approach to characters we have taken here in uniting all
Streptocarpinae genera into Streptocarpus to bring taxonomy
in line with phylogeny. While the twisted fruit is no longer a
defining characteristic of the genus, it was not without exceptions before; i.e., Str. capuronii has capsules “not or scarcely
(through one turn of the spiral only) twisted” (Hilliard & Burtt,
1971: 310), and Linnaeopsis subscandens on the other hand has
strongly twisted fruits (Weigend, 2000: 49). With the inclusion
of the seven Streptocarpinae genera with non-twisted fruits,
the circumscription of the genus Streptocarpus has been widened, and can now be defined as including woody caulescents
with twisted fruits, herbaceous caulescents with twisted or
non-twisted fruits, and rosulates and unifoliates with twisted,
non-twisted cylindrical or globose fruits.
Within Streptocarpus there are strong correlations
between geographic distribution and phylogenetic relatedness, reflected in sections that are strictly either Madagascan
or African. Both subgenera include subclades from the two
regions. Hilliard & Burtt (1971) often drew morphological links
across this geographic divide (pp. 44–46), but none have been
substantiated here. Neither has Engler’s section Elongati been
found to represent a natural group, in which he included the
Madagascan Str. thompsonii R.Br. and African Str. elongatus
(Engler, 1921), here placed in separate sections. It appears that
geography rather than morphology is the key to understanding
the evolution of Streptocarpus.
Although nine species remain unplaced to section, our
work represents a huge step forwards in the classification of
Streptocarpus and a stable baseline for future refinement.
ACKNOWLEDGMENTS
The authors thank the horticulturists at the Royal Botanic Garden Edinburgh (RBGE) for their excellent work in maintaining and
accurately curating the living Gesneriaceae collections, in particular S. Barber and L. Galloway who also collected critical material
of Tanzanian species for study without which this work would not
have been possible. We are extremely grateful to J. McNeill, R. Mill
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Nishii & al. • Streptocarpus redefined
(both RBGE) for a critical revision of the nomenclatural aspects of
the manuscript, and to D. Middleton (Singapore Botanic Gardens)
for helpful comments, M. Walters, University of Canterbury, New
Zealand, for images of Str. elongatus, O. Poncy and M. Gaudeul at the
Paris herbarium and curators at BM and K for access to specimens,
and publishers for permission to use illustrations in Fig. 1. We are also
grateful to two anonymous reviewers for their constructive comments
on the manuscript. Initial research funds to M. Möller came from the
Leverhulme Trust (grant no. F/00 771 A-0735) and the Carnegie Trust
for the Universities of Scotland. Funding was granted by the South
African National Research Foundation to D. Bellstedt as part of the
South African Biodiversity Initiative (SABI). We would also like to
acknowledge the Sibbald Trust (project 2012#9) of the RBGE and the
Japan Society for the Promotion of Science (JSPS KAKENHI Grant
Number 15K18593) for support for the phylogenetic work of K. Nishii.
We thank M. Hart, L. Forrest, R. Hollands for useful comments and
help at the molecular laboratories at RBGE. RBGE is supported by
the Rural and Environment Science and Analytical Services division
(RESAS) in the Scottish Government.
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Appendix 1. Origin of the material used in the phylogenetic analyses of matrix Str226.
Taxon, locality, collector(s), collector number, herbarium code, and ITS, rpl20-rps12, and trnLF GenBank accession numbers. Asterisk indicates newly generates sequences.
Acanthonema strigosum Hook.f._1, Cameroon, Macinder B. BM491 (K), FJ501306, *KR703824, FJ501454. Acanthonema strigosum Hook.f._2, Cameroon,
Plana V. VP88 (E), *KR704038, *KR703825, *KR703921. Colpogyne betsiliensis (Humbert) B.L.Burtt_1, Madagascar: Fianarantsoa, Mt Itremo, Moeller M.
& al. MMO0122 (E), *KR704039, *KR703826, *KR703922. Colpogyne betsiliensis (Humbert) B.L.Burtt_2, Madagascar: West Betsileo, Humbert H. 28225
(E), *KR704040, –, –. Didymocarpus citrinus Ridl., Malaysia: Perlis, Kedah Peak, Davis P.H. 69437 (E), DQ912669, *KR703821, AJ492293. Haberlea rhodopensis Friv., ex cult (Bulgaria, Greece), s.n. (E), AF316898, *KR703820, AJ492296. Hovanella madagascarica (C.B.Clarke) A.Weber & B.L.Burtt, Madagascar: Antsiranana, Moeller M. & Andriantiana J., MM9880A (E), *KR704041, *KR703827, FJ501451. Hovanella vestita (Baker) A.Weber & B.L.Burtt,
Madagascar: Antananarivo, Maromiza, Moeller M. & Andriantiana J. MMO0115 (E), *KR704042, *KR703828, *KR703923. Linnaeopsis alba (E.A.Bruce)
B.L.Burtt, Tanzania: Uluguru Mts, above Morogoro town, NW side of Lupanga ridge, Pócs T. & Pócs S. 86106/D (E), AF316933, *KR703829, –. Linnaeopsis
sp._1, Tanzania: Uluguru Mts, above Morogoro town, Palata ridge, Pócs T. & Pócs S. 8656C (E), AF316932, *KR703830, *KR703924. Linnaeopsis sp._2,
Tanzania: Morogoro, N.W. Uluguru Mts, Palata track, Mabberley D.J. 1254 (E), *KR704043, –, –. Linnaeopsis sp._3, Tanzania: Uluguru Mts, above Morogoro
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Appendix 1. Continued.
town, Pócs T. & Pócs S., 8656C (E), *KR704044, *KR703831 (short), *KR703925. Paraboea rufescens (Franch.) B.L.Burtt, China: Yunnan, Möller M., Qi
Y.D. MMO01-99 (E), FJ501316, *KR703823, FJ501469. Primulina spadiciformis (W.T.Wang) Mich.Möller. & A.Weber, China: unknown locality, ex Smithsonian Institution 94-087 (E), AF316900, *KR703822, AJ492291. Saintpaulia brevipilosa B.L.Burtt_1, Tanzania: Kanga forest, Mt. Kanga (Nguru Mts), Pócs
T. s.n. (E), AF316924, *KR703832, *KR703926. Saintpaulia cf. inconspicua B.L.Burtt_1, Tanzania: Uluguru Mts, unknown locality, Watkins C. CW430/01/03
(C.Watkins), *KR704050, *KR703839, *KR703933. Saintpaulia cf. ionantha H.Wendl., Tanzania: Tanga, Sigi River, Moors D.R. s.n. (–), AF316923, –,
*KR703934. Saintpaulia difficilis B.L.Burtt_1, Tanzania: Tanga Region, Bogner J. BNR (E), *KR704045, *KR703833, *KR703927. Saintpaulia diplotricha
B.L.Burtt, Tanzania: Tanga Region, Munich Univ., s.n. (E), *KR704046, *KR703834, *KR703928. Saintpaulia goetzeana Engl._1, Tanzania: Uluguru Mts,
Evans M.J. (AGGS) s.n. (Ephoto), *KR704047, *KR703835, *KR703929. Saintpaulia grandifolia B.L.Burtt_1, ex cult. (Tanzania: E. Usambara Mts), ex Hannover B.G./Marburg B.G. MMOG-240 (E), *KR704048, *KR703836, *KR703930. Saintpaulia grotei Engl._1, Tanzania: E. Usambara Mts, Munich Univ. s.n.
(E), *KR704049, *KR703837, *KR703931. Saintpaulia inconspicua B.L.Burtt_2, Tanzania: Uluguru Mts, unknown locality, Watkins C. s.n. (E), *KR704051,
*KR703838, *KR703932 (short). Saintpaulia magungensis E.P.Roberts_1, ex cult. (Tanzania), ex RBG Kew 1983-8183 (E), *KR704052, –, *KR703935.
Saintpaulia nitida B.L.Burtt_1, ex cult (Tanzania: Nguru Mts), ex RBG Kew 1987-1566 (E), AF316925, –, *KR703936. Saintpaulia pendula B.L.Burtt, Tanzania: NE Usambara Mts, ex Phillipps Univ. Marburg B.G. s.n. (E), *KR704053, *KR703840, *KR703937. Saintpaulia pusilla Engl._2, Tanzania: Uluguru
Mts, Mabberley D.J. 1362 (E), *KR704054, –, *KR703938 (short). Saintpaulia rupicola B.L.Burtt_1, ex cult. (Kenya), ex Hannover B.G. VS2.2 (E), *KR704055,
*KR703841, –. Saintpaulia rupicola B.L.Burtt_1, Kenya: Kaloleni, ex RBG-Kew 1987-1283 (E), –, –, *KR703939. Saintpaulia shumensis B.L.Burtt_1, ex
cult. (Tanzania: W. Usambara Mts, Mt Shume), ex Clements T. s.n. (E), *KR704056, –, *KR703940. Saintpaulia teitensis B.L.Burtt_1, Kenya: Mbololo, Taita
Hills, Punter W.R. s.n. (E), AF316922, –, –. Saintpaulia tongwensis H.Wendl._1, Tanzania: Tanga Region, ex Clements T. Mather 2 (–), FJ501303, *KR703842,
FJ501446. Saintpaulia ulugurensis Haston, Tanzania: Morogoro, Uluguru Mts, Haston E. 102 (E), *KR704057, *KR703843, *KR703941. Saintpaulia velutina
B.L.Burtt, Tanzania: Usambara Mts, ex Munich University s.n. (E), FJ501304, *KR703844, AJ492303. Saintpaulia watkinsii Haston_1, Tanzania: Morogoro,
Morogoro-Rural, Lusangalala area, Tegetero, Uluguru Nature Reserve, Barber S. & Galloway L. TZBG65 (E), *KR704058, *KR703845, *KR703942. Saintpaulia watkinsii Haston_2, Tanzania: Morogoro, Uluguru Mts, Haston E. 99 (E), *KR704059, *KR703846, –. Schizoboea kamerunensis (Engl.) B.L.Burtt_1,
Burundi: unknown locality, Lewalla 6603 (E), FJ501305, *KR703847, FJ501453. Schizoboea kamerunensis (Engl.) B.L.Burtt_2, unknown origin, Drumond
& Hemsley D&H1590 (E), *KR704085, *KR703848, *KR703943. Streptocarpus andohahelensis Humbert_1, Madagascar: Tulear, Ranomafana, Moeller M.
& Rafanantsoa G. MM9717A (E), AF316903, –, FJ501449. Streptocarpus andohahelensis Humbert_2, Madagascar: Tulear, Ranomafana, Moeller M. & Rafanantsoa G. MM9717B (E), *KR704086, *KR703849, *KR703944. Streptocarpus aylae T.J.Edwards, South Africa: Eastern Cape, Msikaba, Edwards T.J. &
Styles D. s.n. (Bews Herbarium, PMB), HQ719017, HQ719134, HQ718937. Streptocarpus baudertii L.L.Britten_1, South Africa: KwaZulu-Natal, Otterspoort,
Hughes M. & al. MH1067 (E), HQ719049, HQ719166, *KR703945. Streptocarpus baudertii L.L.Britten_2, South Africa: Eastern Cape, Collywobbles, Hughes
M. & al. MH1189 (E), *KR704087, *KR703850, *KR703946. Streptocarpus beampingaratrensis subsp. beampingaratrensis Humbert_1, Madagascar: Tulear,
Ranomafana, Bellstedt D.U. DUB1234 (STE), *KR704088, –, *KR703947. Streptocarpus beampingaratrensis subsp. beampingaratrensis Humbert_2,
Madagascar: Tulear, Ranomafana, Moeller M. & Rafanantsoa G. MM9715 (E), *KR704089, *KR703851, FJ501448. Streptocarpus bindseili E.Fischer, Rwanda:
S. Nyakabuye, ex Munich B.G. [ex Bonn] s.n. (E), AF316960, *KR703852, *KR703948. Streptocarpus bolusii C.B.Clarke_1, South Africa: Satans Pass, Ngcobo,
Eastern Cape, Badenhorst L. 60 (NBG) (STE), HQ718999, HQ719116, HQ718923. Streptocarpus bolusii C.B.Clarke_2, South Africa: Satans Pass, Ngcobo,
Eastern Cape, Badenhorst L. 60 (NBG) (STE), HQ719000, HQ719117, HQ718924. Streptocarpus brevistamineus Humbert, Madagascar: Antsiranana, Maikampango forest, Moeller M. & Andriantiana J. MM9888A (E), *KR704090, *KR703853, *KR703949. Streptocarpus buchananii C.B.Clarke, ex cult. (Malawi,
S. Tanzania, N. Mocambique), ex AGGS s.n. (E), AF316919, *KR703854, *KR703950. Streptocarpus bullatus Mansf._1, Tanzania: Morogoro, Uluguru Mts,
E. of Magari peak, Pócs, Kondela & Nchimbi 6297 (E), AF316942, –, –. Streptocarpus bullatus Mansf._2, Tanzania: Morogoro, Mvomero, Beho-beho area,
Barber S. & Galloway L. TZBG41 (E), *KR704091, *KR703855, *KR703951. Streptocarpus bullatus Mansf._3, Tanzania: Morogoro, Morogoro-Rural, Kigadu
area, Barber S. & Galloway L. TZBG58 (E), *KR704092, *KR703856, *KR703952. Streptocarpus caeruleus Hilliard & B.L.Burtt_1, South Africa: Limpopo,
Lajuma, Bellstedt D.U. DUB1011a (STE), HQ719018, HQ719135, HQ718938. Streptocarpus caeruleus Hilliard & B.L.Burtt_2, South Africa: Limpopo, Blouberg, Bellstedt D.U. DUB1065 (STE), HQ719019, HQ719136, HQ718939. Streptocarpus candidus Hilliard, South Africa: KwaZulu-Natal, Ngome Forest,
Edwards T.J. s.n. (Bews Herbarium, PMB), HQ719007, HQ719124, HQ718927. Streptocarpus capuronii Humbert_1, Madagascar: Antsiranana, Mt Ambodilaitra, Moeller M. & Andriantiana J., MM9885A (E), *KR704093, –, *KR703953. Streptocarpus capuronii Humbert_2, Madagascar: unknown locality, Gautier
L. LG3936 (G), *KR704094, *KR703857, *KR703954. Streptocarpus caulescens Vatke, ex cult. (Tanzania, Kenya), Mr & Mrs I La Crox 1481 (E), AF316920,
*KR703858, *KR703955. Streptocarpus cf. mangindranensis Humbert, Madagascar: unknown locality, Gautier L. LG3477 (G), MacMaster & al. (2005), –,
MacMaster & al. (2005). Streptocarpus cf. michelmorei B.L.Burtt_1, Zimbabwe; Manicaland Province, Chimanimani National Park, Clark 458 (STE),
*KR704095, *KR703859, *KR703956. Streptocarpus cf. michelmorei B.L.Burtt_2, Zimbabwe; Manicaland Province, Chimanimani National Park, Clark 459
(STE), *KR704096, *KR703860, *KR703957. Streptocarpus compressus B.L.Burtt, Tanzania: Songea Distr., Matengo Hills, Luiri Kitesi, Redhead M &
Taylor 8982A (E), AF316957, –, –. Streptocarpus confusus Hilliard, South Africa: Natal, Near Hluhluwe Game Reserve, Makowe Mt., Harrison E.R. s.n. (E),
AF316966, –, –. Streptocarpus cooksonii B.L.Burtt, South Africa: Jamisons Drift, Edwards T.J. s.n. (Bews Herbarium, PMB), *KR704097, HE861712,
HE956758. Streptocarpus cooperi C.B.Clarke, South Africa: Natal, Nkandla distr. Qudeni forest, Edwards T.J. s.n. (E), AF316954, –, –. Streptocarpus
cyanandrus B.L.Burtt, Zimbabwe; Inyanga Downs, Wild H. 4943 (E), AF316947, *KR703861, *KR703958. Streptocarpus cyaneus S.Moore_1, South Africa:
Limpopo, Mariepskop, Hughes M. & al. MH1329 (E), HQ719024, HQ719141, HQ718943. Streptocarpus cyaneus S.Moore_2, South Africa: Mpumalanga,
Josephsdal, Hughes M. & al. MH1355_02 (E), HQ719025, HQ719142, HQ718944. Streptocarpus cyaneus subsp. longi-tommii Weigend & T.J.Edwards, South
Africa: Mpumalanga, DieGeut, Hughes M. & al. MH1232 (E), *KR704098, –, *KR703959. Streptocarpus cyaneus subsp. nigridens Weigend & T.J.Edwards,
South Africa: Limpopo, Soutpansberg, Bellstedt D.U. DUB1077_01 (STE), HQ719023, HQ719140, HQ718942. Streptocarpus daviesii N.E.Brown ex C.B.Clarke,
South Africa: Kwa-Zulu Natal, Laager Farm, Wartburg, Bellstedt D.U. DUB955 (STE), *KR704099, HE861713, HE956759. Streptocarpus davyi S.Moore,
Swaziland: Mbabane, Burtt B.L. 3551 (E), AF316946, *KR703862, *KR703960 (short). Streptocarpus decipiens Hilliard & B.L.Burtt, South Africa: E. Transvaal, Hilliard O.M. & Burtt B.L. H&B6000 (E), AF316938, *KR703863, *KR703961. Streptocarpus denticulatus Turrill_1, South Africa: Mpumalanga,
Belfast, Bellstedt D.U. DUB1096a (STE), HQ718991, HQ719108, HQ718915. Streptocarpus denticulatus Turrill_2, South Africa: Mpumalanga, Belfast,
Bellstedt D.U. DUB1096b (STE), HQ718992, HQ719109, HQ718916. Streptocarpus dolichanthus Hilliard & B.L.Burtt, Malawi: Mt. Mulanje, Litelenya Plateau,
Chapman J.D. & Chapman E.G. 8496 (E), AF316937, *KR703864, *KR703962. Streptocarpus dunnii Hook.f._1, South Africa: Mpumalanga, Uitvlugt Farm,
Hughes M. & al. MH1268 (E), HQ718988, HQ719105, HQ718912. Streptocarpus dunnii Hook.f._2, South Africa: Mpumalanga, Slaaihoek, Hughes M. & al.
MH1273 (E), HQ718989, HQ719106, HQ718913. Streptocarpus dunnii Hook.f._3, South Africa: Mpumalanga, Angle Station, Hughes M. & al. MH1371 (E),
HQ718990, HQ719107, HQ718914. Streptocarpus elongatus Engl._1, Nigeria: Taraba state, Mambilla Plateau, Nge Nyaki Forest, Spurrier M. N705 (E),
AF316913, *KR703865, *KR703963. Streptocarpus elongatus Engl._2, Nigeria: Taraba state, Mambilla Plateau, Nge Nyaki Forest, Spurrier M. N705 (–),
*KR704100, *KR703866, *KR703964. Streptocarpus erubescens Hilliard & B.L.Burtt, Malawi; Ndirandi Mts, Cram. 66-1294 (E), AF316949, *KR703867,
*KR703965. Streptocarpus exsertus Hilliard & B.L.Burtt_1, Kenya: Lolokwe Mt., Samburu District., Bellstedt D.U. & Bytebier B. 564 (NMK Nairobi),
*KR704101, –, *KR703966. Streptocarpus exsertus Hilliard & B.L.Burtt_2, Kenya, Lolokwe Mt., NE slope of Onulbeys, Gilbert 5358 (E), AF316939,
*KR703868, *KR703967. Streptocarpus eylesii S.Moore, Zimbabwe: Nyanga Mountains, Rukotso area, Clark 518 (STE), *KR704102, *KR703869, *KR703968.
Streptocarpus fanniniae Harvey ex C.B.Clarke_1, South Africa: Natal, Lion’s distr., Bridgewood, Dargle, Dale J.E. s.n. (E), AF316944, –, *KR703969. Streptocarpus fanniniae Harvey ex C.B.Clarke_2, South Africa: KwaZulu-Natal, Mt. Gilboa, Bellstedt D.U. DUB1025 (STE), HQ718982, HQ719099, HQ718906.
Streptocarpus fasciatus T.J.Edwards & C.Kunhardt, South Africa: Mpumalanga, Crocodile River Gorge, Hughes M. & al. MH1291(MMOG237C) (E),
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Appendix 1. Continued.
*KR704103, *KR703870, *KR703970. Streptocarpus fenestra-dei Weigend & T.J.Edwards, South Africa: Mpumalanga, GodsWindow, Hughes M. & al.
MH1305 (E), *KR704104, –, *KR703971. Streptocarpus floribundus M.Weigend & T.J.Edwards, South Africa: KwaZulu-Natal, *KRanskop, Edwards T.J.
s.n. (Bews Herbarium, PMB), HQ719026, HQ719143, HQ718945. Streptocarpus formosus (Hilliard & B.L.Burtt) T.J.Edwards_1, South Africa: Eastern Cape,
Mzamba, Burring J. 27 (Bews Herbarium, PMB), HQ719063, HQ719180, HQ718969. Streptocarpus formosus (Hilliard & B.L.Burtt) T.J.Edwards_2, South
Africa: Eastern Cape, Oribi Gorge, Bellstedt D.U. DUB1136_01 (STE), HQ719065, HQ719182, HQ718971. Streptocarpus galpinii Hook.f., Swaziland: Mbabane,
Burtt B.L. 3552 (E), AF316943, *KR703871, –. Streptocarpus gardenii Hook._1, South Africa: KwaZulu-Natal, Hoha Forest, Bellstedt D.U. DUB0560_01
(STE), HQ719008, HQ719125, HQ718928. Streptocarpus gardenii Hook._2, South Africa: KwaZulu-Natal, Mt. Gilboa, Bellstedt D.U. DUB1027_02 (STE),
HQ719011, HQ719128, HQ718931. Streptocarpus glandulosissimus Engl., ex cult. (Congo, Rwanda, Rurundi, Uganda, Tanzania, Kenya), Hilliard O.M. s.n.
(E), AF316918, *KR703872, *KR703972. Streptocarpus goetzei Engler, ex cult. (Tanzania, Moçambique, Malawi), La Croix I.F. s.n. (E), AF316956, *KR703872,
*KR703973. Streptocarpus grandis N.E.Brown_1, South Africa: KwaZulu-Natal, NW of Inanda Mt., Styles D.G.A. 3007_01 (STE), HQ718993, HQ719110,
HQ718917. Streptocarpus grandis N.E.Brown_2, South Africa: KwaZulu-Natal, Kingscliff Farm, Styles D.G.A. 3019_02 (STE), HQ718994, HQ719111,
HQ718918. Streptocarpus haygarthii N.E.Brown ex C.B.Clarke, South Africa, Natal, mid-Illovo, Edwards T.J. s.n. (E), AF316964, –, –. Streptocarpus hildebrandtii Vatke, ex cult. Madagascar: Parc Botanique et Zoologique de Tsimbazaza, Moeller M. & Rafanantsoa G. MM9725 (E), AF316930, *KR703874,
MacMaster & al. (2005). Streptocarpus hilsenbergii R.Brown_1, Madagascar: Asaranitra, Andringitra, Bellstedt D.U. DUB1274 (STE, TAN), *KR704105,
*KR703875, *KR703974. Streptocarpus hilsenbergii R.Brown_2, Madagascar: Ambohimahasoa, Fianarantsoa District, Bellstedt D.U. DUB1280 (STE, TAN),
*KR704106, HE861719, HE956765. Streptocarpus hirticapsa B.L.Burtt_1, Zimbabwe; Manicaland Province, Chimanimani National Park, Weigend M. s.n.
(E), AF316962, *KR703876, *KR703975. Streptocarpus hirticapsa B.L.Burtt_2, Zimbabwe; Manicaland Province, Chimanimani National Park, Clark 456
(STE), *KR704107, *KR703877, *KR703976. Streptocarpus hirticapsa B.L.Burtt_3, Zimbabwe; Manicaland Province, Chimanimani National Park, Clark
492 (STE), *KR704108, *KR703878, *KR703977. Streptocarpus hirtinervis C.B.Clarke, Malawi: Mt. Mulanje, Burtt B.L. 71-0126 (E), AF316935, *KR703879,
*KR703978. Streptocarpus holstii Engl., ex cult. (Tanzania, E. Usambara Mts), ex Cornell Univ. (Bail. Hort) 88 (E), AF316917, *KR703880, AJ492304.
Streptocarpus huamboensis B.L.Burtt, Angola: Mount Moco, Huambo Province, Harrower A. AGH4492 (Kirstenbosch NBG), *KR704109, HE861720,
HE956766. Streptocarpus ibityensis Humbert_1, Madagascar: Antananarivo, Mt. Ibity, Fischer E. 250/93 (E), AF316926, –, FJ501455. Streptocarpus ibityensis Humbert_2, Madagascar: Antananarivo, Mt. Ibity, Moeller M. & al. MMO0132b (E), *KR704110, –, *KR703979. Streptocarpus ibityensis Humbert_3,
Madagascar: Antananarivo, Mt. Ibity, Moeller M. & al. MMO0132ca (E), MacMaster & al. (2005), *KR703881, MacMaster & al. (2005). Streptocarpus
inflatus B.L.Burtt, Tanzania: Udzungwa Mts, Luke Q. (MMOG-112) (E), *KR704111, *KR703882, *KR703980. Streptocarpus itremensis B.L.Burtt_1,
Madagascar: Antananarivo, Mt. Ibity, Moeller M. & al. MMO0138Ba (E), MacMaster & al. (2005), *KR703883, MacMaster & al. (2005). Streptocarpus
itremensis B.L.Burtt_2, Madagascar: Antananarivo, Mt. Ibity, Moeller M. & Rafanantsoa G. MM9723 (E), AF316928, –, MacMaster & al. (2005). Streptocarpus itremensis B.L.Burtt_3, Madagascar: Fianarantsoa, Mt Itremo, Bellstedt D.U. DUB1302 (STE, TAN), *KR704112, –, *KR703981. Streptocarpus
itremensis B.L.Burtt_4, Madagascar: Fianarantsoa, Mt Itremo, Moeller M. & Andriantiana J. MM9898B (E), *KR704113, *KR703884, *KR703982. Streptocarpus johannis L.L.Britten_1, South Africa: Eastern Cape, Embotyi, Bellstedt D.U. DUB0840 (STE), HQ719054, HQ719170, HQ718965. Streptocarpus
johannis L.L.Britten_2, South Africa: KwaZulu-Natal, Hebron Road, Bellstedt D.U. DUB0714 (STE), HQ719056, HQ719171, HQ718966. Streptocarpus
kentaniensis L.L.Britten & Story_1, South Africa: Cape province, E. Transkei, Kei Mouth, Kunhardt, M. s.n. (E), AF316974, –, *KR703983. Streptocarpus
kentaniensis L.L.Britten & Story_2, South Africa: Eastern Cape, Kentani, Joannou J. 6 (STE), HQ719014, HQ719131, HQ718934. Streptocarpus kimbozanus
B.L.Burtt_1, Tanzania: Kimboza Forest reserve, Pócs T. 72-1700 (E), AF316911, *KR703885, *KR703984. Streptocarpus kimbozanus B.L.Burtt_2, Tanzania:
unknown locality, Polhill & Lovett 4911 (E), *KR704114, *KR703886, *KR703985. Streptocarpus kirkii Hook.f., Tanzania: Tanga region, E Usambara Mts,
Johnston-Stewart N.G.B. s.n. (E), AF316916, *KR703887, *KR703986. Streptocarpus kungwensis Hilliard & B.L.Burtt, Tanzania: W. slope of Musenabantu,
Harley & Raymond M. 9370 (E), AF316959, –, –. Streptocarpus kunhardtii T.J.Edwards, South Africa: Mpumalanga, Itala, Edwards T.J. s.n. (Bews Herbarium, PMB), *KR704115, –, *KR703987. Streptocarpus lanatus MacMaster_1, Madagascar: Fianarantsoa, Mt Itremo, Bellstedt D.U. DUB1285 (STE, TAN),
*KR704116, –, *KR703988. Streptocarpus lanatus MacMaster_2, Madagascar: Fianarantsoa, Mt Itremo, Moeller M. & al. MMO0119a2 (E), MacMaster & al.
(2005), *KR703888, MacMaster & al. (2005). Streptocarpus lanatus MacMaster_3, Madagascar: Fianarantsoa, Mt Itremo, Moeller M. & al. MMO0119a1 (E),
MacMaster & al. (2005), –, MacMaster & al, (2005). Streptocarpus levis B.L.Burtt_2, Madagascar: Tulear, Andranohela River, Moeller M. & Rafanantsoa
G. MM9709 (E), *KR704117, *KR703889, *KR703989. Streptocarpus lilliputana D.U.Bellstedt & T.J.Edwards_1, South Africa: Eastern Cape, Lupatana Gorge,
Bellstedt D.U. DUB0611 (Bews Herbarium, PMB), HQ719015, HQ719132, HQ718935. Streptocarpus lilliputana D.U.Bellstedt & T.J.Edwards_2, South Africa:
Eastern Cape, Upper Fraser Falls, FraserGorge, Bellstedt D.U. DUB0627 (Bews Herbarium, PMB), HQ719016, HQ719133, HQ718936. Streptocarpus lokohensis Humbert, Madagascar: Antsiranana, Mt Ambodilaitra, Moeller M. & Andriantiana J., MM9886A (E), MacMaster & al. (2005), *KR703890, MacMaster
& al. (2005). Streptocarpus longiflorus (Hilliard & B.L.Burtt) T.J.Edwards, South Africa: Limpopo, Blouberg, Bellstedt D.U. DUB1064_01 (STE), HQ719020,
HQ719137, HQ718940. Streptocarpus macropodus B.L.Burtt, Madagascar: Andringintra, Camp IV, Lewis & al. 1038 (E), AF316927, *KR703891, –. Streptocarpus makabengensis Hilliard, South Africa: Limpopo, Machabengberg, Truter J. s.n. (–), *KR704118, HE861721, HE956767. Streptocarpus meyeri
B.L.Burtt_1, South Africa: Eastern Cape, Bastervoetpad, Bellstedt D.U. DUB0913_01 (STE), HQ719027, HQ719144, HQ718946. Streptocarpus meyeri
B.L.Burtt_2, South Africa: Eastern Cape, Bastervoetpad, Bellstedt D.U. DUB0913_07 (STE), HQ719028, HQ719145, HQ718947. Streptocarpus meyeri
B.L.Burtt_3, South Africa: Eastern Cape, Somerset East, Glen Craig Farm, Bellstedt D.U. DUB0896_02 (STE), HQ719036, HQ719153, HQ718953. Streptocarpus meyeri B.L.Burtt_4, South Africa: Eastern Cape, Baviaanskloof, Jamieson R. s.n. (–), HQ719038, HQ719155, HQ718954. Streptocarpus meyeri
B.L.Burtt_5, South Africa: Eastern Cape, Zuurberg, Bellstedt D.U. DUB0844 (STE), HQ719040, HQ719157, HQ718956. Streptocarpus meyeri B.L.Burtt_6,
South Africa: Eastern Cape, Graaff-Reinet, Bellstedt D.U. DUB0776_01 (STE), HQ719041, HQ719158, HQ718957. Streptocarpus micranthus C.B.Clarke_1,
South Africa: Mpumalanga, Shia Longubu dam, Hughes M. & al. MH1375 (E), *KR704121, –, *KR703990. Streptocarpus micranthus C.B.Clarke_2, South
Africa: Mpumalanga, Graskop Resort, Bellstedt, D.U. DUB581 (STE), *KR704122, –, –. Streptocarpus milanjianus Hilliard & B.L.Burtt, Malawi: Mt. Mulanje,
Burtt B.L. 6385 (E), AF316936, *KR703892, *KR703991. Streptocarpus modestus L.L.Britten_1, South Africa: Eastern Cape, Upper Fraser Falls, Bellstedt
D.U. DUB0624_01 (STE), HQ719060, HQ719177, HQ718967. Streptocarpus modestus L.L.Britten_2, South Africa: Eastern Cape, Upper Fraser Falls, Bellstedt
D.U. DUB0624_02 (STE), HQ719061, HQ719178, HQ718968. Streptocarpus molweniensis Hilliard, South Africa: Kwa-Zulu Natal, Dlinza Forest, Eshowe,
Bellstedt D.U. DUB1357 (STE), *KR704123, HE861724, HE956768. Streptocarpus monophyllus Welwitsch_1, Angola: Huila Province, Humpata District,
Zootechnica, Bellstedt D.U. DUB1172 (STE), *KR704124, HE861725, HE956769. Streptocarpus monophyllus Welwitsch_2, Angola: Huila Province, Humpata District, Zootechnica, Bellstedt D.U. DUB1179 (STE), *KR704125, HE861726, HE956770. Streptocarpus montanus Oliv._1, Kenya: Taita Taveta, Mbololo
Forest, Bytebier B. BB1937 (–), HQ718981, HQ719098, –. Streptocarpus montanus Oliv._2, Tanzania: Kilimanjaro, Mvomero, S. Pare Mts, Barber S. & Galloway L. TZBG71 (E), *KR704126, *KR703893, *KR703992. Streptocarpus montanus Oliv._3, Tanzania: Morogoro, Mvomero, Kolweza Mts, Barber S. &
Galloway L. TZBG4 (E), *KR704127, *KR703894, *KR703993. Streptocarpus montigena L.L.Britten_1, South Africa: Eastern Cape, Elandsberg, McMaster
C. s.n. (STE), HQ719045, HQ719162, HQ718961. Streptocarpus montigena L.L.Britten_2, South Africa: Eastern Cape, Katberg Pass, Bellstedt D.U. DUB0897_02
(STE), HQ719048, HQ719165, HQ718964. Streptocarpus muscosus C.B.Clarke, Madagascar: Tulear, Col de Tanatana, Moeller M. & Rafanantsoa G., MM9703
(E), *KR704128, *KR703895, *KR703994. Streptocarpus nimbicola Hilliard & B.L.Burtt, Hilliard O.M., Malawi: Mt. Mulanje, Burtt B.L. 67-0252 (E),
AF316934, *KR703896, *KR703995. Streptocarpus nobilis C.B.Clarke_1, ex cult. (W. Tropical Africa), ex Vienna Univ. B. G. s.n. (E,photo), AF316912,
*KR703897, *KR703996. Streptocarpus nobilis C.B.Clarke_2 (W. Tropical Africa), Jongkiad & Abbiw 1919 (Leiden), *KR704129, *KR703898, *KR703997.
Streptocarpus nobilis C.B.Clarke_3, cult. origin, ex Berlin Botanic Garden 63-1823 (E), *KR704130, –, –. Streptocarpus occultus Hilliard, South Africa:
Mpumalanga, Warburton, Truter J. s.n. (–), *KR704131, HE861727, HE956771. Streptocarpus oliganthus B.L.Burtt_1, Madagascar: Antsiranana, Marojezy
1270
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Appendix 1. Continued.
RN12, Moeller M. & Andriantiana J. MM9866 (E), *KR704132, *KR703899, *KR703998. Streptocarpus oliganthus B.L.Burtt_2, Madagascar: Antsiranana,
Mt Ambodilaitra, Moeller M. & Andriantiana J., MM9882 (E), *KR704133, –, *KR703999. Streptocarpus pallidiflorus C.B.Clarke, Tanzania: Arusha region,
Masai distr., Longido Mts, Longido Stream, Carmichael Rev. W. s.n. (E), AF316921, *KR703900, *KR704000. Streptocarpus papangae Humbert_1, Madagascar: Tulear, Col de Beampingaratra, Moeller M. & Rafanantsoa G. MM9718 (E), AF316929, –, FJ501444. Streptocarpus papangae Humbert_2, Madagascar: Tulear, Col de Beampingaratra, Moeller M. & Rafanantsoa G. MM9718 (E), HQ718980, HQ719097, HQ718905. Streptocarpus parensis B.L.Burtt_1,
Tanzania: Kilimanjaro, Mvomero, S. Pare Mts, Barber S. & Galloway L. TZBG70 (E), *KR704134, *KR703901, *KR704001. Streptocarpus parensis B.L.Burtt_2,
Tanzanai: Kilimanjaro, Mvomero, S. Pare Mts, Barber S. & Galloway L. TZBG66 (E), *KR704135, *KR703902, *KR704002. Streptocarpus parviflorus
Hook.f._1, South Africa: unknown locality, La Croix I.F. s.n. (E), *KR704136, –, *KR704003. Streptocarpus parviflorus Hook.f._2, South Africa: Limpopo,
Magoebaskloof Hotel, Hughes M. & al. MH1292 (E), HQ719021, HQ719138, *KR704004. Streptocarpus parviflorus subsp. soutpansbergensis Weigend &
T.J.Edwards, South Africa: Limpopo, Soutpansberg, Bellstedt D.U. DUB1075 (STE), HQ719022, HQ719139, HQ718941. Streptocarpus pentherianus Fritsch,
ex cult. (South Africa: Transvaal, Swaziland, Natal), ex Chiltern Seeds s.n. (E), AF316971, –, –. Streptocarpus perrieri Humbert, Madagascar: Antananarivo,
Angavo near Ankazobe, Moeller M., Rafanantsoa G. & Irapanarivo S. MM9726 (E), AF316931, *KR703903, MacMaster & al. (2005). Streptocarpus poleevansii Verdoorn, ex cult. (South Africa: Transvaal), La Croix I.F. s.n. (E,photo), AF316950, –, *KR704005. Streptocarpus polyanthus Hook._1, South Africa:
KwaZulu-Natal, Shelter Falls, Hughes M. & al. MH1022 (E), HQ719002, –, *KR704006. Streptocarpus polyanthus Hook._2, South Africa: KwaZulu-Natal,
Ferncliff, Hughes M. & al. MH1031 (E), HQ719004, HQ719121, *KR704007. Streptocarpus porphyrostachys Hilliard_1, South Africa: E. Cape, Transkei,
Mkabati nature reserve, Kunhardt C. KNT60 (E), AF316963, –, *KR704008. Streptocarpus porphyrostachys Hilliard_2, South Africa: Eastern Cape, Mtentu
Gorge, Bellstedt D. DUB0984 (STE), HQ719001, HQ719118, HQ718925. Streptocarpus primulifolius Gand._1, South Africa: Eastern Cape, Port Shepstone,
Rooivaal, Hughes M. & al. MH1088 (–), HQ719082, HQ719199, *KR704009. Streptocarpus primulifolius Gand._2, South Africa: KwaZulu-Natal, Table Mt.,
Inanda, Edwards T.J. s.n. (Bews Herbarium, PMB), HQ719071, HQ719188, HQ718977. Streptocarpus primulifolius Gand._3, South Africa: Eastern Cape,
Msikaba, Bellstedt D.U. DUB0965_01 (STE), HQ719072, HQ719189, HQ718978. Streptocarpus primulifolius Gand._4, South Africa: KwaZulu-Natal, Stone’s
Farm, Hughes M. & al. MH1052 (E), HQ719074, HQ719191, *KR704010. Streptocarpus primulifolius Gand._5, South Africa: Eastern Cape, Endliniyokozi,
Hughes M. & al. MH1126 (E), HQ719075, HQ719192, *KR704011. Streptocarpus primulifolius Gand._6, South Africa: Eastern Cape, Dwalana Forest, Bellstedt D.U. DUB0587 (STE), HQ719079, HQ719196, *KR704012. Streptocarpus prolixus C.B.Clarke, South Africa: Natal, Pinetown Distr., Everton, Leep &
Pasche E. LEP72/2 (E), AF316973, –, –. Streptocarpus pumilus B.L.Burtt, Zimbabwe, Domboshawa, Weigend M. s.n. (E), AF316948, *KR703904, *KR704013.
Streptocarpus pusillus Harvey ex C.B.Clarke_1, South Africa: KwaZulu-Natal, Mt. Gilboa, Bellstedt D.U. DUB1029a (STE), HQ718983, HQ719100, HQ718907.
Streptocarpus pusillus Harvey ex C.B.Clarke_2, South Africa: Natal, Anne Rennies’s Mountain sunset farm, Bulwer, ex RBG Kew 1983-1816 (K), AF316945,
–, *KR704014. Streptocarpus rexii (Bowie ex Hook.) Lindl._1, South Africa: Eastern Cape, Kologha center, Hughes M. & al. MH1149 (E), HQ719088,
HQ719205, *KR704015. Streptocarpus rexii (Bowie ex. Hook.) Lindl._2, South Africa: Eastern Cape, Tsitsikamma, Hughes M. & al. MH1180 (E), HQ719096,
HQ719212, *KR704016. Streptocarpus rimicola Story_1, South Africa: Limpopo, Thabazimbi, Bellstedt D.U. DUB1047 (STE), HQ718997, HQ719114,
HQ718921. Streptocarpus rimicola Story_2, South Africa: Limpopo, Thabazimbi, Bellstedt D.U. DUB1048 (STE), HQ718998, HQ719115, HQ718922. Streptocarpus roseo-albus Weigend & T.J.Edwards, South Africa: Mpumalanga, Agnes Mine, Hughes M. & al. MH1353 (E), *KR704137, –, *KR704017. Streptocarpus sambiranensis Humbert, Madagascar: Antsiranana, Sambirano basin, Humbert H. HH18583 (E), *KR704138, *KR703905, *KR704018. Streptocarpus
saundersii Hook._1, South Africa: KwaZulu-Natal, Inanda, Styles D.G.A. DGA3013_02 (Bews Herbarium, PMB), HQ719006, HQ719123, HQ718926. Streptocarpus saundersii Hook._2, South Africa: Ozwatini Plateau, Styles D.G.A. DGA3021 (Bews Herbarium, PMB), –, HE861728, HE956772. Streptocarpus
saxorum Engl., Tanzania: Usambara, Tanga region, Mather S. 1330 (E), AF316914, *KR703906, *KR704019. Streptocarpus schliebenii Mansf._1, Tanzania:
Ukaguru Mts, Burtt B.L. s.n. (E), AF316941, *KR703907, *KR704020. Streptocarpus schliebenii Mansf._2, Tanzania: Ukaguru Mts, Mabberley D.J. 1398
(E), *KR704139, –, –. Streptocarpus schliebenii Mansf._3, Tanzania: Morogoro, Gairo, Mnyera Peak, Ukaguru Mts, Barber S. & Galloway L. TZBG37 (E),
*KR704140, *KR703908, *KR704021. Streptocarpus silvaticus Hilliard, South Africa: Natal Province, Lion’s River Distr., Karkloof range, Benvie, Hilliard
O.M. & Burtt B.L. HBT19107 (E,photo), AF316970, *KR703909, *KR704022. Streptocarpus solenanthus Mansf., Malawi: unknown locallity, La Croix I.F.
s..n. (E), AF316958, –, –. †Streptocarpus sp. nov._1, South Africa: Laager farm, Styles D.G.A. DGA3014 (Bews Herbarium, PMB), *KR704119, HE861723, –.
†
Streptocarpus sp. nov._2, South Africa: Inanda Mountain, towards Wartberg, Styles D.G.A. DGA3006 (Bews Herbarium, PMB), *KR704120, HE861722, –.
Streptocarpus stomandrus B.L.Burtt_1, Tanzania: Nguru Mts, Mabberley D.J. 687C (E), AF316915, *KR703910, *KR704023. Streptocarpus stomandrus
B.L.Burtt_2, Tanzania: Morogoro, Mvomero, Manyangu Forest, Barber S. & Galloway L. TZBG23 (E), *KR704141, *KR703911, *KR704024. Streptocarpus
suffruticosus Humbert, Madagascar: Antsiranana, Marojezy RN12, Moeller M. & Andriantiana J. MM9877A (E), MacMaster & al. (2005), *KR703912,
MacMaster & al. (2005). Streptocarpus tanala Humbert_1, Madagascar: Tulear, Analaro, Andranohela River, Moeller M. & Rafanantsoa G. MM9710 (E),
AF316906, *KR703913, *KR704025. Streptocarpus tanala Humbert_2 (C & E Madagascar), Lowry II P.P. 4364 (TAN), *KR704142, –, *KR704026. Streptocarpus thompsonii R.Brown_1, Madagascar: Analalava Forest, Andringtra, Bellstedt D. DUB1258 (STE, TAN), *KR704143, –, *KR704027. Streptocarpus
thompsonii R.Brown_2, Madagascar: Antananarivo, Angavoke, Moeller M. & Rafanantsoa G. MM9851 (E), *KR704144, *KR703914, *KR704028. Streptocarpus thysanotus Hilliard & B.L.Burtt, Tanzania: Morogoro Distr., Kimboza, ex AGGS (Moeller M. MMOG-90) (E), AF316910, *KR703915, *KR704029.
Streptocarpus trabeculatus Hilliard, ex cult. (South Africa: E. Cape, Natal), ex Chiltern Seeds s.n. (E), AF316969, –, –. Streptocarpus tsaratananensis
Humbert ex B.L.Burtt, Madagascar: unknown locality, Gautier L. LG3600 (G), MacMaster & al. (2005), –, MacMaster & al. (2005). Streptocarpus umtaliensis B.L.Burtt, Zimbabwe, Manicaland Province, Leopard Rock Hotel, Bvumba, Clark 498 (STE), *KR704145, *KR703916, *KR704030. Streptocarpus vandeleurii E.G.Baker & S.Moore_1, South Africa: Limpopo, Thabazimbi, Bellstedt D.U. DUB1055 (STE), HQ718995, HQ719112, HQ718919. Streptocarpus
vandeleurii E.G.Baker & S.Moore_2, South Africa: Limpopo, Thabazimbi, Bellstedt D.U. DUB1055 (STE), HQ718996, HQ719113, HQ718920. Streptocarpus
variabilis Humbert_1, Madagascar: Anjouan lsland, Pócs T. 9282A (E), *KR704146 (short), –, *KR704031. Streptocarpus variabilis Humbert_2, Madagascar:
Anjouan lsland, Pócs T. 9282A (E), *KR704147, *KR703917, *KR704032. Streptocarpus variabilis Humbert_3, Madagascar: Road to Beolamana, Moeller M.
MMOG32 (E), *KR704148, *KR703918, *KR704033. Streptocarpus venosus B.L.Burtt, Madagascar: Tulear, Andranohela River, Moeller M.& Rafanantsoa
G. MM9711 (E), *KR704149, *KR703919, *KR704034. Streptocarpus wendlandii Spreng._1, ex cult. (South Africa: Natal; Mtunzini Distr., Ngoye forest), ex
Hannover B.G. s.n. (E), AF316967, –, *KR704035. Streptocarpus wendlandii Spreng._2, South Africa: KwaZulu-Natal: Ngoye Forest, Mtunzini., Bellstedt
D.U. DUB1336/1345 (STE), *KR704150, HE861729, HE956773. Streptocarpus wilmsii Engl., South Africa: Mpumalanga, God’s Window, Potgieter C. CP907
(Bews Herbarium, PMB), *KR704151, HE861730, HE956774. Streptocarpus wittei DeWild., Malawi: Rumpi Distr., Nyika, La Croix I.F. s.n. (E), AF316953,
–, *KR704036. Trachystigma mannii C.B.Clarke (Gabon), deWilde, Sosef & Van Nek 10139 (Leiden), *KR704152, *KR703920, *KR704037.
† these collections from KwaZulu Natal Province, South Africa will be described “Streptocarpus medley-woodii ” in sect. Streptocarpus.
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Appendix 2. Origin of the material used in the phylogenetic analyses of matrix Sa58.
Taxon, locality, collector(s), collector number, herbarium code, and ITS and 5S NTS GenBank accession numbers. Asterisk indicates newly generates sequences.
Saintpaulia brevipilosa B.L.Burtt_1, Tanzania: Kanga forest, Mt. Kanga (Nguru Mts), Pócs, T. s.n. (E), AF316924, AF307025. Saintpaulia brevipilosa
B.L.Burtt_2, Tanzania: unknown locality, RBG Kew 1995-503 (K), –, AF108733. Saintpaulia cf. inconspicua B.L.Burtt_1, Tanzania: Uluguru Mts, unknown
locality, Watkins C. CW430/01/03 (–), *KR704050, –. Saintpaulia cf. ionantha H.Wendl., Tanzania: Tanga, Sigi River, Moors D.R. s.n. (–), AF316923, AF307034.
Saintpaulia confusa B.L.Burtt_1, Tanzania: Usambara Mts, Smith J. R6677 (E), *KR704060, AF307027. Saintpaulia confusa B.L.Burtt_2, Tanzania: unknown
locality, RBG Kew 1974-2873 (K), *KR704061, AF108728. Saintpaulia difficilis B.L.Burtt_1, Tanzania: Tanga Region, Bogner J. BNR (E), *KR704045,
AF307028. Saintpaulia difficilis B.L.Burtt_2, Tanzania, Tanga Region, Mather I.C. MAT1 (E), *KR704062, AF324926. Saintpaulia difficilis B.L.Burtt_3,
Tanzania: E Usambara Mts, Kwamkoro, Lindqvist C. Kiganga.1 (–), –, AF307019. Saintpaulia diplotricha B.L.Burtt, Tanzania: Tanga Region, ex Munich Univ.
s.n. (E), *KR704046, AF307029. Saintpaulia goetzeana Engl._1, ex cult. (Tanzania: Uluguru Mts), Evans M.J. (AGGS) s.n. (E,photo), *KR704047, AF307030.
Saintpaulia goetzeana Engl._2, Tanzania: Mbete–Lupanga peak, N Uluguru Forest Reserve, Morogoro, Lindqvist C. Lupanga.1 (–), *KR704063, AF108734.
Saintpaulia goetzeana Engl._3, Tanzania: Kilangala, South Uluguru Forest Reserve, Morogoro, Lindqvist C. Kilangala.1 (–), *KR704064, AF108735. Saintpaulia grandifolia B.L.Burtt_1, ex. cult. (Tanzania), ex Hannover B.G./Marburg B.G., MMOG–240 (E), *KR704048, –. Saintpaulia grandifolia B.L.Burtt_2,
Tanzania: Tanga Region, W Usambara Mts, Mather I.C. MAT13 (E), *KR704065, AF307031. Saintpaulia grandifolia B.L.Burtt_3, ex cult. (Tanzania), ex
Hannover B.G./Marburg B.G. s.n. (E), *KR704066, –. Saintpaulia grandifolia B.L.Burtt_4, Tanzania: W Usambara Mts, Lindqvist C. Nkoloi Stream.3 (–), –,
AF307021. Saintpaulia grotei Engl._1, Tanzania: E. Usambara Mts, ex Munich Univ. s.n. (E), *KR704049, AF307032. Saintpaulia grotei Engl._2, Tanzania:
Usambara Mts, ex RBG Kew 1995-511 (K), –, AF108729. Saintpaulia inconspicua B.L.Burtt_2, Tanzania: Uluguru Mts, unknown locality, Watkins C. s.n.
(E), *KR704051, –. Saintpaulia intermedia B.L.Burtt, Tanzania: E Usambara Mts, Kigongoi, ex Phillips Univ. Marburg B.G. s.n. (E), *KR704067, AF307033.
Saintpaulia ionantha H.Wendl. ‘Sigi Falls’, Tanzania: Tanga, Sigi Falls ex RBG Kew 1987-1280 (–), *KR704068, AF307036. Saintpaulia ionantha H.Wendl.,
Tanzania: Ulang Distr., Uzyngwa Mts, ex RBG Kew 1987-8132 (K), *KR704069, AF108732. Saintpaulia magungensis E.P.Roberts var. minima B.L.Burtt_1,
Tanzania: E Usambara Mts, Marvera, Punter W.R. 1181 (E), *KR704070, AF307038. Saintpaulia magungensis E.P.Roberts var. minima B.L.Burtt_2, Tanzania: unknown origin, ex RBG Kew 1963-42311 (K), –, AF108738. Saintpaulia magungensis E.P.Roberts_1, ex cult. (Tanzania: E. Usambara Mts), ex RBG
Kew 1983-8183 (E), *KR704072, AF307037. Saintpaulia magungensis E.P.Roberts_2, Tanzania: Magunga Forest, E Usambara Mts, Lindqvist C. Magunga.4
(–), *KR704052, AF108737. Saintpaulia magungensis var. occidentalis B.L.Burtt, Tanzania: Usambara Mts, Mather I.C. MAT15 (–), *KR704071, AF307039.
Saintpaulia nitida B.L.Burtt_1, ex cult. (Tanzania: Nguru Mts), ex RBG Kew 1987-1566 (E), AF316925, –. Saintpaulia nitida B.L.Burtt_2, Tanzania: Nguru
Mts, ex Basel Univ. B.G. CL8/99 (E), *KR704073, AF324927. Saintpaulia orbicularis B.L.Burtt_1, Tanzania: Ambangulu–Kunga Road, West Usambara
Mts, Lindqvist C. Kwabulu.5 (–), –, AF108740. Saintpaulia orbicularis B.L.Burtt_2, Tanzania: W Usambara Mts, Ambangulu Tea Estate, ex RBG Kew
1987-1370 (K), –, AF108739. Saintpaulia orbicularis B.L.Burtt_3, Tanzania: W Usambara Mts, Ambangulu Tea Estate (ex cult.), ex Munich Univ. B.G. s.n.
(E), *KR704075, AF324928. Saintpaulia orbicularis var. purpurea B.L.Burtt, Tanzania: W Usambara Mts, Ambangulu, Punter W.R. s.n. (E), *KR704074,
AF307041. Saintpaulia pendula B.L.Burtt, Tanzania: E Usambara Mts, E of Mawera Tea Estate, Smith J., JS10 (E), *KR704076, AF307043. Saintpaulia
pendula var. kizarae B.L.Burtt, ex cult. (Tanzania: NE Usambara Mts), ex Phillips Univ. Marburg B.G. s.n. (E), *KR704053, AF307042. Saintpaulia pusilla
Engl._1, Tanzania: Magari Peak, North Uluguru Forest Reserve, Morogoro, Lindqvist C. Magari.1 (–), *KR704077, AF108741. Saintpaulia pusilla Engl._2,
Tanzania: unknown locality, Mabberley D.J. 1362 (E), *KR704054, –. Saintpaulia rupicola B.L.Burtt ‘Kacharoroni’_1, Kenya: Kachororoni, Lindqvist C.
CL–97003 (–), *KR704078, AF108743. Saintpaulia rupicola B.L.Burtt ‘Kacharoroni’_2, Kenya: Kilifi Distr., Kachororoni Gorge, Smith J. R+L5126 (E),
*KR704079, AF307045. Saintpaulia rupicola B.L.Burtt ‘Mwache’, Kenya: Mwache Forest Reserve, Pearce 543-94-530.1 (–), *KR704080, AF108744. Saintpaulia rupicola B.L.Burtt ‘Mwarakaya’, Kenya: Kaloleni, Mwarakaya, Smith J. JS02 (E), *KR704081, AF307046. Saintpaulia rupicola B.L.Burtt_1, ex cult.
(Kenya), ex Hannover B.G. VS2.2 (E), *KR704055, AF307044. Saintpaulia rupicola B.L.Burtt_2, Kenya: Chasimba, Lindqvist C. CL97002.1 (–), *KR704082,
AF108742. Saintpaulia rupicola B.L.Burtt_3, Kenya: Mwarakaya, Simiyu S. 169-96-1739 (NMK), –, AF108745. Saintpaulia shumensis B.L.Burtt_1, ex
cult. (Tanzania: W. Usambara Mts, Mt Shume), Clements T. s.n. (E), *KR704056, AF307047. Saintpaulia shumensis B.L.Burtt_2, Tanzania: West Usambara
Mts, Shume Hill, Lindqvist C. Shume.1 (–), –, AF108746. Saintpaulia teitensis B.L.Burtt_1, Kenya: Mbololo, Teita Hills, Punter W.R. s.n. (E), AF316922,
AF307048. Saintpaulia teitensis B.L.Burtt_2, Kenya: Teita Hills, Mbololo, Wallace B.J. & Forlonge L.J. WFO336 (–), *KR704083, AF307049. Saintpaulia
teitensis B.L.Burtt_3, Kenya: Teita Hills, Mbololo, Lindqvist C. CL-97001 (–), *KR704084, AF108747. Saintpaulia tongwensis B.L.Burtt_1, Tanzania: Tanga
Region, Clements T. Mather 2 (–), FJ501303, AF307050. Saintpaulia tongwensis B.L.Burtt_2, Tanzania: Tongwe Mt., Tongwe Forest Reserve, Lindqvist C.
Tongwe.9 (–), –, AF108730. Saintpaulia tongwensis B.L.Burtt_3, Tanzania: Tanga Region, Pangani Falls, Lindqvist C. Pangani_Falls.97010 (–), –, AF108748.
Saintpaulia ulugurensis Haston, Tanzania: Morogoro, Uluguru Mts, Haston E. 102 (E), *KR704057, –. Saintpaulia velutina B.L.Burtt, ex cult. (Tanzania:
Usambara Mts), ex Munich University s.n. (E), FJ501304, AF307051. Saintpaulia watkinsii Haston_1, Tanzania: Morogoro, Morogoro–Rural, Lusangalala area,
Tegetero, Uluguru Nature Reserve, Barber S. & Galloway L. TZBG65 (E), *KR704058, –. Saintpaulia watkinsii Haston_2, Tanzania: Morogoro, Uluguru Mts,
Haston E. 99 (E), *KR704059, –.Streptocarpus caulescens Vatke, Tanzania: Morogoro Region, Uluguru Mts, Harris B. HRR4091 (E), AF316920, AF108727.
Appendix 3. List of species by section of Streptocarpus. * indicates placement based on morphological grounds. Remaining species included in the molecular phylogeny. Types of section names in bold italics.
Streptocarpus subg. Streptocarpella Fritsch
(“a”) sect. Trachystigma (C.B. Clarke) Mich.Möller & Nishii, comb. nov.
Streptocarpus diandra (Engl.) Nishii & Mich.Möller, comb. nov.
Streptocarpus insularis Hutch. & Dalziel
Streptocarpus mannii (C.B.Clarke) Nishii & Mich.Möller, comb. nov.
Streptocarpus nobilis C.B.Clarke in DC
Streptocarpus strigosus (Hook.f.) Nishii & Mich.Möller, comb. nov.
(“b”) sect. Parasaintpaulia Mich.Möller & Nishii, sect. nov.
Streptocarpus andohahelensis Humbert
Streptocarpus beampingaratrensis Humbert subsp. beampingaratrensis
var. beampingaratrensis
Streptocarpus beampingaratrensis subsp. antambolorum Humbert
Streptocarpus beampingaratrensis subsp. antankarana Humbert*
Streptocarpus beampingaratrensis subsp. beampingaratrensis var.
brevicarpus Humbert*
Streptocarpus mandrerensis Humbert*
1272
(“c”) sect. Hova (C.B. Clarke) Mich.Möller & Nishii, comb. nov.
Streptocarpus hilsenbergii R.Br. var. hilsenbergii
Streptocarpus hilsenbergii var. angustifolius Humbert ex B.L.Burtt
Streptocarpus integrifolius B.L.Burtt*
Streptocarpus leandrii Humbert* var. leandrii
Streptocarpus leandrii var. robustus B.L.Burtt*
Streptocarpus levis B.L.Burtt
Streptocarpus linguatus B.L.Burtt
Streptocarpus madagascaricus (C.B.Clarke) Christenh.
Streptocarpus muscosus C.B.Clarke.
Streptocarpus oliganthus B.L.Burtt
Streptocarpus prostratus Humbert* var. prostratus
Streptocarpus prostratus var. major B.L.Burtt*
Streptocarpus tanala Humbert
Streptocarpus thompsonii R.Br. var. thompsonii
Streptocarpus thompsonii var. bojeri (R. Br.) C.B.Clarke*
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Appendix 3. Continued.
Streptocarpus venosus B.L.Burtt in Humbert
Streptocarpus vestitus (Baker) Christenh.
(“d”) sect. Schizoboea (Fritsch) Mich.Möller & Nishii, comb. nov.
Streptocarpus elongatus Engl.
Streptocarpus gonjaensis Engl.*
Streptocarpus kamerunensis (Engl.) Christenh.
Streptocarpus kimbozanus B.L.Burtt
Streptocarpus muscicola Engl.*
Streptocarpus thysanotus Hilliard & B.L.Burtt
(“e”) sect. Carnosifolii Mich.Möller & Nishii, sect. nov.
Streptocarpus bambuseti B.L.Burtt*
Streptocarpus euanthus Mansf.*
Streptocarpus hirsutissimus E.A.Bruce*
Streptocarpus kirkii Hook.f.
Streptocarpus saxorum Engl.
Streptocarpus stomandrus B.L.Burtt
(“f”) sect. Caulescentes (Fritsch) Engl.
Streptocarpus buchananii C.B.Clarke
Streptocarpus caulescens Vatke
Streptocarpus glandulosissimus Engl.
Streptocarpus holstii Engl.
Streptocarpus inflatus B.L.Burtt
Streptocarpus pallidiflorus C.B.Clarke
(“g”) sect. Saintpaulia (H. Wendl.) Mich.Möller & Haston, comb. &
stat. nov.
Streptocarpus afroviola Christenh.
Streptocarpus brevipilosus (B.L.Burtt) Mich.Möller & Haston, comb. nov.
Streptocarpus goetzeanus (Engl.) Christenh.
Streptocarpus inconspicuus (B.L.Burtt) Christenh.
Streptocarpus ionanthus (H.Wendl.) Christenh. subsp. ionanthus
var. ionanthus
Streptocarpus ionanthus subsp. grandifolius (B.L.Burtt) Christenh.
Streptocarpus ionanthus subsp. grotei (Engl.) Christenh.
Streptocarpus ionanthus subsp. ionanthus var. diplotrichus (B.L.Burtt)
Christenh.
Streptocarpus ionanthus subsp. mafiensis (I.Darbysh. & Pócs) Christenh.
Streptocarpus ionanthus subsp. occidentalis (B.L.Burtt) Christenh.
Streptocarpus ionanthus subsp. orbicularis (B.L.Burtt) Christenh.
Streptocarpus ionanthus subsp. pendulus (B.L.Burtt) Christenh.
Streptocarpus ionanthus subsp. rupicolus (B.L.Burtt) Christenh.
Streptocarpus ionanthus subsp. velutinus (B.L.Burtt) Christenh.
Streptocarpus nitidus (B.L.Burtt) Mich.Möller & Haston, comb. nov.
Streptocarpus shumensis (B.L.Burtt) Christenh.
Streptocarpus teitensis (B.L.Burtt) Christenh.
Streptocarpus ulugurensis (Haston) Haston, comb. nov.
Streptocarpus watkinsii (Haston) Haston, comb. nov.
Streptocarpus subg. Streptocarpus
(“h”) sect. Lignostreptocarpus Mich.Möller & Bellstedt, sect. nov.
Streptocarpus campanulatus B.L.Burtt*
Streptocarpus coursii Humbert*
Streptocarpus glabrifolius Humbert*
Streptocarpus macropodus B.L.Burtt
Streptocarpus papangae Humbert
Streptocarpus suffruticosus Humbert var. suffruticosus
Streptocarpus suffruticosus var. hirtellus Humbert*
Streptocarpus suffruticosus var. pachycarpus B.L.Burtt*
Streptocarpus suffruticosus var. sericeus B.L.Burtt*
Streptocarpus tsaratananensis Humbert ex B.L.Burtt
(“i”) sect. Colpogyne Mich.Möller & Bellstedt, comb. & stat. nov.
Streptocarpus ibityensis Humbert
Streptocarpus itremensis B.L.Burtt
Streptocarpus lanatus MacMaster
Streptocarpus revivescens Humbert ex B.L.Burtt*
Streptocarpus betsiliensis Humbert
(“j”) sect. Plantaginei Mich.Möller & Bellstedt, sect. nov.
Streptocarpus boinensis Humbert*
Streptocarpus brevistamineus Humbert
Streptocarpus cordifolius Humbert*
Streptocarpus hildebrandtii Vatke
Streptocarpus lokohensis Humbert
Streptocarpus mangindranensis Humbert
Streptocarpus perrieri Humbert
Streptocarpus plantagineus Vatke*
Streptocarpus polyphyllus Humbert*
Streptocarpus sambiranensis Humbert
Streptocarpus stellulifer B.L.Burtt*
Streptocarpus suborbicularis B.L.Burtt in Humbert
Streptocarpus variabilis Humbert
Streptocarpus velutinus B.L.Burtt in Humbert
(“k”) sect. Protostreptocarpus Mich.Möller & Bellstedt, sect. nov.
Streptocarpus albus (E.A.Bruce) I.Darbysh. subsp. albus
Streptocarpus albus subsp. edwardsii (Weigend) I.Darbysh.*
Streptocarpus heckmannianus (Engl.) I.Darbysh. subsp. heckmannianus*
Streptocarpus heckmannianus subsp. gracilis (E.A.Bruce) I.Darbysh.*
Streptocarpus subscandens (B.L.Burtt) I.Darbysh.*
Streptocarpus bullatus Mansf.
Streptocarpus montanus Oliver
Streptocarpus parensis B.L.Burtt
Streptocarpus schliebenii Mansf.
Streptocarpus lineatus (B.L.Burtt) Mich.Möller & M.Hughes, comb. nov.
(“l”) sect. Streptocarpus
Streptocarpus actinoflorus T.J.Edwards & M.Hughes*
Streptocarpus arcuatus Hilliard & B.L.Burtt*
Streptocarpus aylae T.J.Edwards
Streptocarpus baudertii L.L.Britten
Streptocarpus bindseili Eb. Fisch.
Streptocarpus bolusii C.B.Clarke
Streptocarpus brachynema Hilliard & B.L.Burtt*
Streptocarpus breviflos (C.B.Clarke) C.B.Clarke in Dyer*
Streptocarpus burttianus T.Pócs*
Streptocarpus caeruleus Hilliard & B.L.Burtt
Streptocarpus candidus Hilliard
Streptocarpus compressus B.L.Burtt
Streptocarpus confusus Hilliard subsp. confusus*
Streptocarpus confusus subsp. lebomboensis Hilliard & B.L.Burtt
Streptocarpus cooksonii B.L.Burtt
Streptocarpus cooperi C.B.Clarke
Streptocarpus cyanandrus B.L.Burtt
Streptocarpus cyaneus S.Moore subsp. cyaneus
Streptocarpus cyaneus subsp. longi-tommii Weigend & T.J.Edwards
Streptocarpus cyaneus subsp. nigridens Weigend & T.J.Edwards
Streptocarpus cyaneus subsp. polackii (B.L.Burtt) Weigend &
T.J.Edwards*
Streptocarpus daviesii N.E.Br. ex C.B.Clarke
Streptocarpus davyi S.Moore
Streptocarpus decipiens Hilliard & B.L.Burtt
Streptocarpus denticulatus Turrill
Streptocarpus dolichanthus Hilliard & B.L.Burtt
Streptocarpus dunnii Hook.f.
Streptocarpus erubescens Hilliard & B.L.Burtt
Streptocarpus eylesii S.Moore subsp. eylesii
Streptocarpus eylesii subsp. brevistylus Hilliard & B.L.Burtt*
Streptocarpus eylesii subsp. chalensis I.Darbysh.*
Streptocarpus eylesii subsp. silvicola Hilliard & B.L.Burtt*
Streptocarpus fanniniae Harvey ex C.B.Clarke
Streptocarpus fasciatus T.J.Edwards & C.Kunhardt
Streptocarpus fenestra-dei Weigend & T.J.Edwards
Streptocarpus floribundus Weigend & T.J.Edwards
Streptocarpus formosus (Hilliard & B.L.Burtt) T.J.Edwards
Streptocarpus galpinii Hook.f.
Streptocarpus gardenii Hook.
Streptocarpus goetzei Engl.
Streptocarpus grandis N.E.Br. subsp. grandis
Streptocarpus grandis subsp. septentrionalis Hilliard & B.L.Burtt*
Streptocarpus haygarthii N.E.Br. ex C.B.Clarke
Streptocarpus hilburtianus T.J.Edwards*
Streptocarpus hirticapsa B.L.Burtt
Streptocarpus hirtinervis C.B.Clarke
Streptocarpus huamboensis Hilliard & B.L.Burtt*
Streptocarpus johannis L.L.Britten
Streptocarpus katangensis DeWild. & T.Durand*
Streptocarpus kentaniensis L.L.Britten & Story
Streptocarpus kungwensis Hilliard & B.L.Burtt
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Appendix 3. Continued.
Streptocarpus kunhardtii T.J.Edwards
Streptocarpus latens Hilliard & B.L.Burtt*
Streptocarpus leptopus Hilliard & B.L.Burtt*
Streptocarpus lilliputana Bellstedt & T.J.Edwards
Streptocarpus longiflorus (Hilliard & B.L.Burtt) T.J.Edwards
Streptocarpus makabengensis Hilliard
Streptocarpus meyeri B.L.Burtt
Streptocarpus michelmorei B.L.Burtt
Streptocarpus micranthus C.B.Clarke
Streptocarpus milanjianus Hilliard & B.L.Burtt
Streptocarpus modestus L.L.Britten
Streptocarpus molweniensis Hilliard subsp. molweniensis
Streptocarpus molweniensis subsp. eshowicus Hilliard & B.L.Burtt*
Streptocarpus monophyllus Welw.
Streptocarpus montigena L.L.Britten
Streptocarpus montis-bingae Hilliard & B.L.Burtt*
Streptocarpus myoporoides Hilliard & B.L.Burtt*
Streptocarpus nimbicola Hilliard & B.L.Burtt
Streptocarpus occultus Hilliard
Streptocarpus parviflorus Hook.f. subsp. parviflorus
Streptocarpus parviflorus subsp. soutpansbergensis Weigend &
T.J.Edwards
Streptocarpus pentherianus Fritsch
Streptocarpus pogonites Hilliard & B.L.Burtt*
Streptocarpus pole-evansii I.Verd.
Streptocarpus polyanthus Hook. subsp. polyanthus
Streptocarpus polyanthus subsp. comptonii (Mansf.) Hilliard*
Streptocarpus polyanthus subsp. dracomontanus Hilliard*
Streptocarpus polyanthus subsp. verecundus Hilliard*
Streptocarpus porphyrostachys Hilliard
Streptocarpus primulifolius Gand.
Streptocarpus prolixus C.B.Clarke
Streptocarpus pumilus B.L.Burtt
Streptocarpus pusillus Harvey ex C.B.Clarke
Streptocarpus rexii (Bowie ex Hook.) Lindl.
Streptocarpus rhodesianus S.Moore subsp. rhodesianus*
Streptocarpus rhodesianus subsp. grandiflorus I.Darbysh.*
1274
Streptocarpus rimicola Story
Streptocarpus roseo-albus Weigend & T.J.Edwards
Streptocarpus saundersii Hook.
Streptocarpus silvaticus Hilliard
Streptocarpus solenanthus Mansf.
Streptocarpus trabeculatus Hilliard
Streptocarpus umtaliensis B.L.Burtt
Streptocarpus vandeleurii Baker f. & S.Moore
Streptocarpus wendlandi Spreng.
Streptocarpus wilmsii Engl.
Streptocarpus wittei DeWild.
Incertae sedis
Streptocarpus burundianus Hilliard & B.L.Burtt — Note: No molecular
data available and morphologically incompletely known.
Streptocarpus exsertus Hilliard & B.L.Burtt — Note: See the Results
section under “Characteristics and phylogenetic relationships revealed
by the Str226 analysis.”
Streptocarpus masisiensis DeWild. — Note: No molecular data available
and morphologically incompletely known.
Streptocarpus mbeyensis I.Darbysh. — Note: No molecular data available and morphologically incompletely known.
Streptocarpus phaeotrichus B.L.Burtt — Note: No molecular data available and morphologically incompletely known.
Streptocarpus capuronii Humbert — Note: See the Results section
under “Characteristics and phylogenetic relationships revealed by the
Str226 analysis.”
Streptocarpus semijunctus B.L.Burtt — Note: No molecular data available and morphologically incompletely known.
Streptocarpus stenosepalus B.L.Burtt — Note: No molecular data available and morphologically incompletely known.
Streptocarpus tsimihetorum Humbert — Note: No molecular data available and morphologically incompletely known.
Uncertain and doubtful taxa
? Streptocarpus zimmermanii Engl.
Version of Record