South African Journal of Botany 118 (2018) 241–259

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South African Journal of Botany

journal homepage: www.elsevier.com/locate/sajb

Biosystematic revision of the native and naturalised species of Rubus L.

(Rosaceae) in the Cape Floristic Region, South Africa
M. Sochor a,⁎, B. Trávníček b, J.C. Manning c,d
a
Centre of the Region Haná for Biotechnological and Agricultural Research, Department of Genetic Resources for Vegetables, Medicinal and Special Plants, Crop Research Institute, Šlechtitelů 29,
Olomouc CZ-78371, Czech Republic
b
Department of Botany, Palacký University, Šlechtitelů 27, Olomouc CZ-78371, Czech Republic
c
Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa
d
Research Centre for Plant Growth and Development, School of Biological and Conservation Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville, Pietermaritzburg 3209, South Africa

a r t i c l e i n f o a b s t r a c t

Article history: Rubus L. is a highly diverse and taxonomically complex genus with a worldwide distribution. Although southern
Received 30 April 2018 Africa is not a major centre of Rubus diversity, a number of indigenous and naturalised taxa have been recorded
Received in revised form 22 June 2018 from the region, although their taxonomy, nomenclature and evolution are either confused or poorly studied at
Accepted 3 July 2018
best. This paper represents the first results of a wider project focusing on the biosystematics of the southern
Available online xxxx
African Rubus flora, and presents a taxonomic revision of the brambles of the Cape Floristic Region, including
Edited by I Doležalová an estimation of ploidy levels. A total of eight taxa are recognised, including the two indigenous taxa
R. pinnatus and R. rigidus agg., the four introduced species R. armeniacus, R. bergii, R. rosifolius and R. titanus,
Keywords: and two hybrids between indigenous and introduced taxa. Rubus rigidus agg. is an unresolved and variable
Hybrids species complex with at least two ploidy levels (5x, 6x). The hybrid taxon R. bergii × R. rigidus appears to be wide-
Invasive species spread and behaves as an independent apomictic complex. In addition, hybridization between R. bergii and
New records R. pinnatus is relatively common in the contact zones between the two species. The two introductions Rubus
Ploidy armeniacus and R. titanus are recorded from South Africa for the first time. The invasive potential of the intro-
Taxonomy
duced species is regarded as low but the potential for hybridization by R. bergii seems to be high. An identification
key, morphological descriptions, and notes on nomenclature, distribution and ecology are provided, and all taxa
are illustrated.
© 2018 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction
Rubus L. (brambles, blackberries, raspberries, dewberries etc.) is a
large and diverse genus in the Rosaceae with a worldwide distribution,
and includes thousands of published species names and infrageneric
taxa (Gustafsson, 1943; Kurtto et al., 2010). It is traditionally divided
into twelve subgenera (Focke, 1910–1914), although this classification
is clearly artificial as many of the subgenera have been shown to be
poly- or paraphyletic (Alice and Campbell, 1999). The species-level
taxonomy is very controversial due to the frequent occurrence of facul-
tative apomixis (asexual formation of seeds) and hybridization, particu-
larly common in some of the subgenera (Weber, 1996; Sochor et al.,
2015).
Current taxonomic treatments in Europe and North America, the
two large regions with the highest diversity of apomictic brambles, ap-
proach the subject quite differently. The leading European paradigm,
the so-called “Weberian batology” (from batos = the Greek for
⁎ Corresponding author.
E-mail address: michal.sochor@volny.cz (M. Sochor).
bramble), accepts a very narrow species delimitation that treats every
apomictic genotype with a sufficiently extensive distribution area as a
separate species (Weber, 1996; Holub, 1997), whereas the account of
the genus in the Flora of North America adopts a much broader species
concept (Alice et al., 2015) that groups several (probably many) differ-
ent, often reproductively isolated genotypes, into more broadly
circumscribed species based on morphological similarities, thereby
rejecting the earlier, more detailed treatments by Bailey (1941–1945)
and Davis et al. (1967).
Taxonomic and nomenclatural difficulties are the major reasons
why Rubus is understudied or even neglected in many regions of the
world, e.g. the southern Caucasus (Sochor and Trávníček, 2016), some
parts of Europe (Kurtto et al., 2010), and also South Africa (Stirton,
1981a, 1981b; Henderson, 2011). Data on evolutionary mechanisms,
modes of reproduction, and phylogenesis are also scarce outside
Europe. This is despite the importance of the genus, not only as an inte-
gral component of many vegetation types and as a significant ecological
element across the globe, but also as the source of valuable fruit crops
and of expansive/invasive taxa. Several Rubus species have been intro-
duced into various regions of the world, where they have become

https://doi.org/10.1016/j.sajb.2018.07.015
0254-6299/© 2018 SAAB. Published by Elsevier B.V. All rights reserved.
242 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
dangerous invaders, e. g. European R. ulmifolus Schott, R. praecox Bertol.
or Caucasian R. armeniacus Focke in western North America, South
America and Australia (Mazzolari et al., 2011; Clark et al., 2013;
Bruckart et al., 2017) or R. alceifolius Poir. in many Indian Ocean islands
and Australia (Amsellem et al., 2000).
South Africa is known for the occurrence of many alien brambles,
some of which have substantial environmental and economic
impact, particularly in the east of the region (Henderson, 2011;
Sochor and Manning, pers. obs.). Their taxonomy in the subconti-
nent, however, is still much confused and even the distinction
between native and naturalised taxa is not always clear (Stirton,
1981a, 1981b; Henderson, 2011). Literature on this topic is scarce
and inadequate, and studies on herbarium material are not very
helpful as collections are mostly incomplete, consisting usually of a
single inflorescence or only a tip of the flowering stem. For proper
determination it is necessary to study both sterile stems (i.e. first-
year stems, also called ‘primocanes’) with their associated leaves, in
combination with inflorescences, which usually develop on second-
year stems (‘floricanes’). The minimum herbarium specimens should
thus comprise parts of first-year stems with at least two well-
developed leaves attached (collected from the middle of the stem)
plus one or more well developed inflorescences/infructescences
(also from the middle of the stem).
The earliest, relatively modern account of Rubus in South Africa is
that by Ecklon and Zeyher (1836), in which they enumerate nine
species, including R. bergii (Cham. & Schltdl.) Eckl. & Zeyh.,
R. chrysocarpus Cham. & Schltdl., R. fruticosus L., R. mundii Cham. &
Schltdl., R. pinnatus Willd., R. rigidus Sm. and R. rosifolius Sm. plus the
two newly described taxa R. ludwigii Eckl. & Zeyh. and R. pappei Eckl. &
Zeyh. This number was reduced to five by Harvey (1862) in his account
of the genus for the Flora capensis, in which he recognised just three
native species, R. ludwigii Eckl. & Zeyh., R. pinnatus Willd. (including
R. pappei, treated here as an uncertain species) and R. rigidus Sm.
(including R. chrysocarpus and R. mundii as varieties), as well as the
introduced R. fruticosus var. bergii Cham. & Schltdl. and R. rosifolius Sm.
Some additional specimens remained unassigned, including at least
one potential hybrid collection. In contrast to this conservative treat-
ment, Gustafsson (1934) described a further six native species, mainly
from the eastern parts of the country, and recorded the occurrence of
an additional two introduced species, but the status of several of these
new taxa is uncertain. Since then some preliminary comments on the
genus in southern Africa have been published (Stirton, 1981a, 1981b,
1984), as well as observations on the chromosome numbers, meiotic
chromosome behaviour, reproduction and hybridization of some
species (Spies et al., 1985, 1987; Spies and du Plessis, 1985, 1986). Our
current knowledge of the genus in the subcontinent was summarised
in an informal treatment provided by Henderson (2011), which enu-
merates 13 native and introduced taxa for South Africa. Current ac-
counts of the genus in the Cape Floristic Region (CFR) list the three
taxa R. fruticosus, R. pinnatus and R. rigidus (Bond and Goldblatt, 1984;
Goldblatt and Manning, 2000; Manning and Goldblatt, 2012), without
comment on their native status, although the former is a European
introduction correctly treated under the name R. bergii.
An early reference to the occurrence of brambles in the CFR is found
in C.P. Thunberg's account of his travels in the region in 1772–1775
(Forbes, 1986) in the context of their use as quickset hedges by
European settlers in the region but it is not clear whether these were na-
tive species or plants introduced for the purpose. If the latter, then this is
likely to be one of the original reasons for the introduction of European
species into the country, presumably along with their value as a fruit
crop. Certainly R. bergii was well established in the environs of Cape
Town by the middle of the nineteenth century, when Harvey (1862) de-
scribed it as the ‘common bramble or blackberry of the Cape’. Partial
hedges of R. bergii are still evident on roadsides and the edges of
vineyards around Stellenbosch today, and may represent the remains
of these early hedges.
In view of the inadequacies in our knowledge of the biosystematics
of the South African bramble flora, we have undertaken a detailed
study of the topic using both classical and modern cytogenetic/molecu-
lar approaches. This paper presents the first results of a wider project
focusing on diversity, evolution and classification of the South African
taxa of Rubus. In this contribution, we revise the genus Rubus in the
CFR in terms of their taxonomy, nomenclature and ploidy levels, and
provide descriptions, pictures and a key to species of this greatly
overlooked plant group. No Rubus collections have been made from
Namaqualand or the Roggeveld Escarpment, i.e. the so-called Extra
Cape Flora of the CFR (Snijman, 2013), and the genus is thus restricted
to the Core Cape Flora Region within the Greater Cape Floristic Region
(Manning and Goldblatt, 2012).
2. Materials and methods
This work was based on field studies on and around the Cape Penin-
sula, and the adjacent Boland and Overberg of Western Cape, South
Africa. We examined all specimens in BOL, NBG, PRE and SAM (acro-
nyms after Thiers, 2018), the primary holdings of material from the
Western Cape, as well as all relevant types. Only taxonomically unam-
biguous specimens from the CFR are cited. Descriptions were based on
material collected in the field, supplemented by herbarium specimens
and literature sources (Launert, 1978; Lingdi and Boufford, 2003)
(only the latter two in the case of R. rosifolius).
DNA ploidy level was estimated from flow cytometric measure-
ments using a BD Accuri C6 flow cytometer (BD Biosciences, Franklin
Lakes, NJ, USA), staining with propidium iodide and Solanum
lycopersicum and Zea mays as internal standards, as described in
Sochor and Trávníček (2016). All voucher specimens are deposited in
NBG and/or OL.
3. Results and discussion
Based on our field and herbarium studies, we recognise eight taxa and
diagnosable hybrids of Rubus from the Cape Floristic Region (sensu
Manning and Goldblatt, 2012), including the two native taxa R. pinnatus
and R. rigidus agg., the four naturalised species R. armeniacus, R. bergii,
R rosifolius and R. titanus, and the two hybrids R. bergii × R. rigidus and
R. bergii × R. pinnatus. This is the first record of R. armeniacus and
R. titanus in South Africa. Many other Rubus taxa, both native and intro-
duced, are recorded from neighbouring eastern South Africa, and their
occasional presence in the CFR cannot be excluded, particularly when tak-
ing into account the long history of blackberry cultivation in the region.
We are, however, only able to recognise these eight taxa and hybrids
with certainty as occurring in the CFR.
We provide an identification key, description and notes to each of
the taxa, including the hybridogenous complex R. bergii × R. rigidus,
which behaves as an independent evolutionary unit. The other hybrid,
R. bergii × R. pinnatus, is mostly of clear intermediate phenotype and oc-
curs together with its parents, and is therefore not described separately.
3.1. Key to native and naturalised Rubus species and hybrids in the Cape
Floristic Region
1a. Petals white, flowers solitary or few in sparse inflorescence;
sepals with long caudate tip; stem villous with long patent
hairs; aggregate fruit bright red, usually of hundreds of small
drupelets … . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.R. rosifolius
1b. Petals pink to white, flowers usually many in ± rich panicles; sepals
acute to acuminate but not caudate; stem glabrous to tomentose
with short hairs; aggregate fruit black, of up to tens of large
drupelets:
2a. Leaves discolourous, grey/white stellate-tomentose beneath:
3a. Leaves pinnate or ternate, never palmate; stem tomentose
with stellate hairs … . . . . . . . . . . . . . . . . . . . . . 2.R. rigidusagg.
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
3b. Leaves palmate or ternate, sometimes pinnately palmate
with reduced or smaller basal leaflets; stem subglabrous to
sparsely hairy:
4a. Inflorescence usually dense and compact with hooked
prickles; sepals erect or spreading to slightly reflexed
after anthesis; primocanes usually terete or obtusely angu-
lar, with prickles of the same colour as stem or slightly red-
dish … . . . 3.R. bergii×R. rigidushybridogenous complex
4b. Inflorescence broadly paniculate with straight,
declinate prickles; sepals reflexed after anthesis;
primocanes usually sharply angular, with red prickles
along angles contrasting with green primocanes in shade
… . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.R. armeniacus
2b. Leaves not markedly discolourous, not tomentose beneath:
5a. Leaves pinnate, mostly 7-foliolate; primocanes pruinose,
pubescent; petals small, ±4–8 mm long, shorter than sepals
… . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.R. pinnatus
5b. Leaves ternate, palmate or pinnate-palmate; primocanes gla-
brous or sparsely hairy, not distinctly pruinose; petals larger,
longer than sepals
6a. Leaves ternate; prickles on primocanes dense, 20 to 40 per
50 mm, unequal in size … . . . . . . . . . . . . . . . . . 7.R. titanus
6b. Leaves on well-developed primocanes 5- to 7-foliolate;
prickles on primocanes dispersed, 4 to 18 per 50 mm,
subequal:
7a. Leaves mostly palmately 5-foliolate; prickles on
primocanes mostly straight (to slightly curved); styles
yellowish … . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.R. bergii
7b. Leaves often pinnate-palmately 7-foliolate; prickles on
primocanes mostly strongly to slightly curved; styles
pink … . . . . . . . . . . . . . . . . . . . . . . . R. bergii×R. pinnatus
3.2. Taxonomic account
Rubus L., Sp. Pl.: 492 (1753). Type: R. fruticosus L.
Note: Characters occurring in extra-southern African taxa are inserted
in square brackets.
Shrubs [subshrubs or perennial herbs], often with biennial stems;
first-year stem (primocane) usually without flowers, simple or (later)
branched, erect, arching, creeping or scrambling, rooting or not at
nodes or tips, terete, angled or furrowed, glabrous or hairy with simple
or stellate hairs, eglandular, stipitate-glandular or sessile-glandular,
usually bearing prickles; prickles sparse to dense, equal or unequal in
size and shape, strong to weak, broad-based to narrow-based, straight
to curved, erect to declinate; second-year stem (floricane) floriferous,
inflorescences usually axillary, sometimes terminal. Stipules filiform to
ovate, glabrous or hairy, glandular or not. Leaves differing according to
the position on the plant (primocane vs. floricane vs. branch vs. inflores-
cence, base vs. middle vs. tip of stem), [simple or] usually compound,
ternate, 5- to 9-pinnate or 5-palmate, glabrous or hairy with simple
hairs to tomentose (then usually with short stellate plus long simple
hairs and leaves discolourous), hairy to touch or not; leaflets (at least
terminal ones) petiolulate, wide-lanceolate to orbicular, margin usually
serrate with serration even or periodic. Inflorescences paniculiform,
racemiform or cymiform, leafy or not; inflorescence axis and branches
terate to angular, usually armed and hairy, glandular or not. Flowers
bisexual [rarely unisexual]; sepals 5, erect, spreading or reflexed, trian-
gular or ovate to long-caudate, unarmed or armed, usually hairy to
tomentose, eglandular or glandular; petals 5 (rarely more), white to
pink or magenta, suborbiculate to elliptic, obovate, or spatulate; sta-
mens >20, in several series, shorter to longer than carpels, sometimes
hairy. Carpels free, usually many (>20), glabrous or hairy to tomentose,
each with two ovaries, but only one of them developing; styles subter-
minal to terminal, slender, caducous after anthesis; stigma bifid or
243
clavate. Fruits drupelets in rich aggregates, but sometimes most of
them aborted; aggregate fruits globose to hemispheric or cylindric, usu-
ally weakly to strongly coherent, separating with or without torus
attached, green turning yellow to red or black at maturity, sometimes
pruinose, with hypanthium and sepals usually persistent.
A large and diverse genus with a cosmopolitan distribution (except
Antarctica). The taxonomy is controversial, with the number of cur-
rently recognised species estimated to be ±1000–1500, depending on
the species concept (Bailey, 1941–1945; Lingdi and Boufford, 2003;
Kurtto et al., 2010; Alice et al., 2015), classified in 12 subgenera. The
larger subgenera (mainly subg. Rubus) are subdivided into sections,
subsections and series. Infraspecific taxa are usually not accepted in
the current taxonomy of apomictic brambles.
The CFR species are currently placed in two subgenera. Subg. Rubus
(most of the European and North American taxa) is recognised by a
shrubby habit, usually erect to arching stems, palmate (or ternate)
primocane leaves, usually distinct prickles, and mostly black aggregate
fruits separating with the torus attached. Subg. Idaeobatus Focke
(South African native taxa but also North American, European and
Asian taxa) is recognised by a shrubby habit, usually erect or scrambling
stems, distinct but sometimes small prickles, imparipinnate leaves (very
rarely simple or palmate in Asian species) and yellow to red or black
aggregate fruit, more or less easily separating from the torus.
3.2.1. Indigenous taxa
1. R. pinnatus Willd., Sp. Pl., 2(2): 1081 (1799). Type: Without lo-
cality, 9879 (B [W09879], lecto.—image!), designated by Cham.
and Schlecht. in Linnea 2: 19 (1827).
Shrub, usually 1.5–2.0 m tall. Primocanes arching or scrambling
(then more than 4 m tall), terete, usually 4–7 mm diam., vinaceous,
greyish pruinose and densely hairy (to sparsely tomentose) with short
simple or stellate and longer simple hairs, with 7 to 25 prickles per
50 mm stem length; prickles unequal in size, strong and broad-based,
slightly to distinctly curved, declinate, 1–6 mm long, 1–5 mm broad at
base, ± concolourous with stem but less pruinose/tomentose, with
brown or yellowish tip. Stipules filiform to narrowly lanceolate, 8–10 ×
± 0.5–1.0 mm, adaxially sparsely hairy with long simple and rarely
also stellate hairs to almost glabrous. Leaves on primocanes pinnate,
exceptionally palmate-pinnate (i.e. with small additional leaflets),
5- to 7(9)-foliolate, usually light green, glabrous above and with only
sparse simple hairs on veins beneath; leaflets broadly to narrowly
ovate or elliptic, usually rugose, with rounded (to shallowly cordate)
base, apex acute to long-acuminate, serration sharp, somewhat irregu-
larly periodic to even (at apex), with primary incisions 1.5–2.0(−4.0)
mm deep; petioles, petiolules and leaf axis pruinose and tomentose,
petioles of varying length, longer or shorter than basal leaflets, with 9
to 18 broad-based curved prickles per 50 mm, petiolules prominent,
7–12% of lamina length of lateral leaflets and 14–32% of lamina length
of terminal leaflet. Inflorescence densely paniculate, 70–180 mm long,
narrowly cylindrical to slightly pyramidal with erect to erecto-patent
branches; leaves in inflorescence usually absent but sometimes up
to six, pinnate, ternate or simple, gradually becoming smaller distally,
similar to primocane leaves but more evenly dentate; bracts lanceolate
to trifid; inflorescence axis densely hairy with subadpressed tufted hairs
and patent longer hairs, with ± 12 to 35 prickles per 50 mm of axis
length; prickles slender, sometimes slightly broad-based, distinctly
curved and declinate, (0.5–)1.0–3.0 mm long; pedicels 4–8 mm long,
densely hairy with short adpressed intertwined hairs and long
semipatent hairs, with 3 to 34 slender subulate unequal prickles.
Flowers many; sepals 6–9 mm long, usually erect to spreading after
anthesis to ± adpressed to fruit, densely hairy with short adpressed
intertwined hairs and long (sub)adpressed hairs, unarmed or with up
to ± 10 unequal pricklets (some immersed in the hairs); petals pale
pink, slightly to distinctly shorter than sepals, narrowly obovate,
obovate or elipsoid, sometimes shallowly emarginate. Stamens unequal,
244 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259

mostly ± shorter than styles (inner series) or as long as or rarely longer
(outer series); filaments white to pale pinkish. Carpels sparsely to
densely hairy with stellate hairs on abaxial side, ovaries green; styles
and stigmas pale to dark pink. Aggregate fruit semiglobose, green turn-
ing red and finally black at maturity (Fig. 1).
DNA-ploidy level: 4x (four individuals analysed: Sochor RSA02/17 [OL,
NBG], RSA04/17 [OL], Sochor & Manning RSA26/17 [NBG, OL], RSA47/17
[NBG]). Spies and du Plessis (1985) report both diploidy and tetraploidy
in the species but we are unable to locate their vouchers to confirm the
identification.
Distribution: reported as widespread from South Africa to tropical Africa
(Guinée to Kenya, Zambia, Mozambique, Zimbabwe: Friis and Vollesen,
1998; Launert, 1978). In South Africa occurring in Limpopo, Mpumalanga,
KwaZulu-Natal, Eastern Cape (but in slightly deviating morphotypes, see
below) and the southwestern parts of Western Cape, where it ranges
along the southern coast mountains as far west as Table Mt on the Cape
Peninsula, with isolated populations inland on the Swartberg (Fig. 2).
Ecology: Rubus pinnatus occurs in a variety of habitats, typically in or
along the margins of primary or secondary forest or woodland, usually
in half-shade amongst trees or tall shrubs but also in full sun or under
closed canopy, and then usually scrambling through the canopy, but
also in disturbed places, including clearings after the felling of planta-
tions, roadsides, and parks.
Diagnostic characters: readily recognised by the pinnate, usually
7-foliolate, subglabrous leaves, pruinose hairy stems, and small pinkish
petals.
Rubus pinnatus is traditionally included in subg. Idaeobatus. It is a
taxonomically distinct species with (presumably) strictly sexual repro-
duction (Sochor, preliminary FCSS data; Spies and du Plessis, 1986). Its
sexuality is reflected in the relatively large morphological variation, as
well as relatively frequent hybridisation with the introduced R. bergii
in the contact zones between the two species. However, introgression
in either direction has not been implied by our preliminary molecular
data (cpDNA and ITS markers).

Fig. 1. Rubus pinnatus: A, primocane leaves; B, immature infructescence; C, detail of primocane; D, F, flowers; E, underside side of leaf; G, fruits.
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Fig. 2. Distribution of Rubus pinnatus in the Cape Floristic Region.
In eastern South Africa, a few morphotypes similar to R. pinnatus
occur, which differ in their glabrous-pruinose or glossy primocanes,
yellow fruits and often 4-jugate leaves. These are usually treated as
R. pinnatus, including plants described as R. pappei Eckl. & Zeyh.
Another morphotype with markedly villous to tomentose
primocanes and branches, ± discolourous leaves that are distinctly
hairy above but villous beneath (thus resembling R. rigidus agg.),
and yellow fruits, is commonly treated as R. apetalus Poir. Some of
these morphotypes likely occur in the easternmost part of CFR,
particularly around Port Elizabeth. Nevertheless, herbarium speci-
mens alone do not allow us to draw any taxonomic conclusions yet
and further studies are necessary.
Additional specimens examined:
WESTERN CAPE: 3318 (Cape Town): Constantia, Cecilia Planta-
tion (–CD), 29 Sep 2011, Ramjukadh C-LR18 (NBG); montis
Tafelberg prope et infra Platteklipp (–CD), Oct/Nov, Ecklon & Zeyher
1705 (SAM); in umbrosis ad Wynberg et Rondebosch (–CD), Jan,
Ecklon & Zeyher 35390 (SAM); Kirstenbosch (–CD), Zeyher 17432
(NBG); Cape Peninsula, Newlands (–CD), Wolley-Dod (BOL); Cape
Town, Kirstenbosch Arboretum, S margin at stream (–CD), 16 Mar
2017, Sochor RSA10/17 (OL); Cape Town, Rondebosch, 300 m NW
of Rhodes Memorial along road (–CD), 16 Mar 2017, Sochor & Man-
ning RSA06/17 (NBG, OL); Table Mt, Skeleton Gorge, 300 m SE of
Hely-Hutchinson Dam (–CD), 11 Mar 2017, Sochor RSA02/17 (OL,
NBG); Table Mt, Woodhead Dam, 60 m W of Waterworks Museum
(–CD), 12 Mar 2017, Sochor RSA04/17 (OL); Kirstenbosch, by Nurs-
ery Stream (–CD), 27 Jan 1941, Barker 846 (PRE); Newlands Station
(–CD), 1916, Page s.n. (PRE); Jonkershoek State Forest, Upper
Biesievlei (–DD), 31 Jan 1984, Richardson 87 (PRE). 3320 (Mon-
tagu): Grootvadersbosch (–DD), 14 Dec 1952, Taylor 5/5 (NBG);
Voormansbosch und in Grootvadersbosch (–CD), Zeyher 2452
245
(SAM). 3322 (Oudtshoorn): Swartbergpas, woudgemeenskap in
nou rotsagtige klofie (–AC), 4 Mar 1985, Geldenhuys 971 (PRE);
George District, Wilderness (–CD), 20 Dec 1928, Mogg 11783
(PRE); Oudtshoorn, Homtini River crossing on the Homtini Pass
between Karatara and Knysna (–DD), 21 Oct 1990, Henderson 1099
(PRE); Millwood (–DD), 26 Sep 2011, Paijmans DMP076 (NBG).
3323 (Willowmore): Buffelsnek Forest Reserve (–CC), 16 Dec
1953, Taylor 1071 (NBG). 3418 (Simonstown): Constantia, between
Spilhaus Ravine and Constantia Nek, along Contour Path (–AB), 26
Mar 2017, Sochor RSA33/17 (OL); Hout Bay, NE end of town along
road to Constantia (–AB), 18 Mar 2017, Sochor RSA13/17 (NBG);
Bergvliet Farm, Constantia (–AB), 2 Nov 1918, Purcell 90038
(SAM); Jonkershoek (–AB), 3 Nov 1941, M.R. Levyns 8486 (BOL); be-
tween Constantia Nek and Hout Bay (–BB), 9 Sep 1976, Stirton
6186A (PRE); Kogelberg State Forest, left of road to Oudebos, at
forking of road next to quarry (–BD), 22 Sep 1992, Kruger 792
(NBG); Betty's Bay botanical garden, 500 m NW of entrance (–BD),
4 Apr 2017, Sochor & Manning RSA47/17 (NBG); entrance to Disa
Gorge, Betty's Bay (–BD), 27 Jun 1977, Stirton 7217 (PRE);
Kogelberg Nature Reserve, 500 m NW of Visitors Centre (–BD), 22
Mar 2017, Sochor & Manning RSA26/17 (NBG, OL). 3419 (Caledon):
Mossel River (–AD), Dec 1912, Potb. 5047 (SAM); Caledon Dist.,
Witvoetskloof (–DD), 31 May 1946, Jordaan 963 (NBG). 3420
(Bredasdorp): Suurbraak, about 2 km out of Suurbraak on road to
Swellendam (–BA), 30 Nov 1991, Henderson 1135 (BOL, PRE).
3423 (Knysna): Knysna (–AA), 10 Nov 1894, Schlechter 1344 (BOL,
PRE). 3424 (Humansdorp): Humansdorp (–AB), Aug 1926, Thode
A820 (PRE); Duine Veld, Slang River (–BA), Nov 1921, Fourcade
1884 (BOL). EASTERN CAPE: 3325 (Port Elizabeth): Loerie Planta-
tion (–CC), Oct 1934, Dix 133 (BOL); on the road between the R75
and St Albans Prison (–CD), 30 Nov 1988, Henderson 1031 (PRE).
246 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
2. R. rigidus agg.
R. rigidus Sm. in Rees, Cyclopedia: 30 (1819). Type: South
Africa, [Western Cape], ‘Cape of Good Hope’, without collector (LINN
[HL653-23], holo.—image!).
R. chrysocarpus Cham. & Schltdl. in Linnea 2: 17 (1827). R. rigidus var.
chrysocarpus (Cham. & Schltdl.) Harv. in Fl. cap. 2: 287 (1862). Type:
South Africa, [Western Cape], ‘In Promontorio bonae spei ad fluvium
Hexrivier’, Mund & Maire s.n. (B?).
R. mundii [as ‘mundtii’] Cham. & Schltdl. in Linnea 2: 18 (1827).
R. rigidus var. mundii (Cham. & Schltdl.) Harv. in Fl. cap. 2: 287 (1862).
Type: South Africa, “Prom. bon. spei, prope Hanglip”, Jul 1815, J.L.Mund
& C.Maire s.n. (HAL [0098161], holo.—image!).
R. discolor sensu E. Mey. in Drége, Zwei pflanzengeogr. Dokum.
(1843), non Weihe & Nees (1824).
The complex also includes the following species currently still
recognised as distinct:
R. atrocaeruleus Gust. in Arkiv Bot. 26A, 7: 54 (1934). Type: Kenya,
‘Kenia orient., inter Meru et Kaseri in frutices’, 22 Feb 1922, E.Rob. & C.E.
Fries 1802 (UPS?, not located).
R. inedulis Rolfe in J. Linn. Soc., Bot. 37: 514 (1906). Type: Uganda,
‘Buddu District at 1230 m alt.’, Dec 1904, E. Brown 133 (K
[000312542], syn.—image!); Uganda, ‘Koki District at 1200 m alt.’,
6 Sep 1905, Dawe 388 (K [000312540], syn.—image!).
R. intercurrens Gust. in Arkiv Bot. 26A, 7: 61 (1934). Type: South
Africa, ‘Transvaal, Fort Cunynghame, in forest on hill side’, 28 May
1898, Galpin 2430 (PRE, holo.!).
R. immixtus Gust. in Arkiv Bot. 26A, 7: 14 (1934). Type: South Africa,
[Eastern Cape], ‘Victoria East, Hogsback’, G.Rattray s.n. (herb. unknown).
R. longepedicellatus (Gust.) C.H. Stirt. in Bothalia 15, 1: 106 (1984).
R. rigidus var. longepedicellatus Gust. in Arkiv Bot. 26A, 7: 59 (1934).
Type: South Africa, [Limpopo], ‘Transvaal, Pietersburg, Duivels Kloof’,
12 Jun 1929, E.E.Galpin 10107 (K, holo.; PRE, iso.—image!).
Shrub, 1–2 (>3) m high. Primocanes arching or scrambling, obtusely
angular with flat or convex sides, 6–7 mm diam., greyish-green to dark
vinaceous on side exposed to sun, densely brownish or whitish tomen-
tose with short stellate hairs, sometimes with dispersed sessile glands;
prickles mildly unequal (especially in size), 8 to 11 per 50 mm of stem
length, usually strong and broad-based, sometimes slender and almost
subulate, usually strongly curved, slightly to distinctly declinate,
4–7 mm long and 2–5 mm broad at base, ± concolourous with stem
or slightly reddish, with yellowish tip. Stipules filiform-linear, ± 12 ×
< 1 mm, densely hairy with stellate and simple hairs. Leaves on
primocanes pinnately 5-foliolate but ternate on terminal part of stem,
usually vivid green, almost glabrous above with sparse hairs on main
veins, whitish/greyish tomentose beneath with short stellate hairs and
long simple hairs, softly hairy to touch; petiole as long as basal leaflets
or much shorter; terminal leaflet ovate with rounded base, acuminate,
serration periodic, primary incisions 2–3 mm deep, petiolule 20–45%
as long as leaflet lamina; lateral leaflets ± obliquely ovate to elliptic.
Inflorescence usually dense, narrowly paniculate, lower branches dis-
tinctly erect to erecto-patent, upper branches short, patent; leaves 1 to
3, simple or ternate, present only in lower half of inflorescence; bracts
lanceolate to trifid; inflorescence axis ± densely hairy with stellate
hairs, with 6 to 9 prickles per 50 mm of axis length; prickles slender,
sometimes broad-based, distinctly to slightly curved, 1–5 mm long;
pedicels very short, usually < 5(− 10) mm long, densely hairy with
long hairs, without prickles. Flowers many; sepals mostly erect or
spreading after anthesis, 5–7 mm long, greyish tomentose and also
with longer ± patent hairs and yellowish sessile glands and 0(− 2)
pricklets; petals pink, narrowly to broadly obovate to suborbicular, ±
as long as sepals or slightly longer. Stamens pink, unequal, ± as long
as styles (inner series often shorter, outer series somewhat longer).
Carpels glabrous, styles pink, stigmas yellowish. Aggregate fruit
semiglobose, green turning red and finally black (sometimes orange?)
at maturity (Fig. 3).
DNA-ploidy level: 5x, 6x (Sochor & Manning RSA27/17 [NBG, OL],
Sochor & Manning RSA40/17 [NBG, OL], respectively).
Distribution: a widely distributed species complex recorded from
tropical Africa (Cameroon, Zaire, Ethiopia, Uganda; Launert, 1978) to
South Africa, where it is reported from all provinces except Northern
Cape (Henderson, 2011); in the CFR it is recorded from the Cederberg
along the western mountains to the Cape Peninsula and eastwards
along the southern coastal mountains to Plettenberg Bay, extending
inland onto the Swartberg and Kouga Mrns (Fig. 4).
Ecology: mainly on roadsides and other secondary habitats, along
rivers or waterlines, in both shady and sunny situations.
Diagnostic characters: readily distinguished from the other CFR
species by the tomentose primocanes with discolourous, mostly
5-pinnate leaves that are whitish tomentose beneath, and the mostly
erect or spreading sepals.
Rubus rigidus is traditionally included in subgenus Idaeobatus and in the
current sense comprises a variable complex of several (possibly many)
morphological types (morphotypes) with at least two ploidy levels in
CFR. Diploidy and tetraploidy were also reported from the eastern South
African R. longepedicellatus (Spies and du Plessis, 1985). Our preliminary
flow cytometry seed screen (FCSS) data indicate that apomixis occurs in
the complex, together with frequent sexuality, at least in hexaploids (see
also Spies and du Plessis, 1986). Further data on reproduction will be
crucial for defining a proper species concept. If apomixis is the dominant
reproductive mode in some genotypes (expected mainly in pentaploids),
it may lead to a uniclonal population structure across a wide geographic
area. Such genotypes often exhibit a distinct combination of morphological
traits and usually fulfil specific ecological roles, and can therefore be con-
sidered as species (microspecies; Weber, 1996).
For the present, we adopt the name R. rigidus agg. for this species
complex, which includes R. immixtus Gust. and R. longepedicellatus
Gust. from eastern South Africa as well as other taxa from tropical
Africa. Some of the taxa that we include in the complex (based on liter-
ature survey and scans of the type material) may represent distinct
microspecies but this is impossible to establish given the present state
of knowledge. Due to the mostly incomplete specimens in herbaria,
which hardly ever include primocanes, extensive field studies will be
necessary to adequately deal with the taxonomy of this complex.
Additional specimens examined: South Africa. WESTERN CAPE: 3219
(Wuppertal): Boontjes River, (–AA), 25 Nov 1910, Pearson 5167
(NBG); Matjesrivier, Cederberg, (–AC), Jan 1944, Wagener 320 (NBG);
Citrusdal, Theerivier (–CA), 7 Sep 1968, Hanekom 1183 (PRE). 3318
(Cape Town): Rondebosch (–CD), Nov 1831, Zeyher 15646 (SAM);
Claremont (–CD), 22 Jul 1893, Schlechter 2977 (BOL); Bosboukloof,
Jonkershoek (–DD), May 1967, Kerfoot K5833 (NBG); Stellenbosch,
Jonkershoek, Swartbas (–DD), 20 Aug 1963, Bos 449 (NBG); Biesiesvlei,
Jonkershoek, edge of swamps (–DD), 2 Feb 1941, Levyns 7539 (BOL);
upper Biesievlei (–DD), 31 Jan 1984, Richardson 87 (NBG); along mill-
stream at Stellenbosch (–DD), Jul 1917, Marloth 7762 (PRE); Stellen-
bosch, Swartboskloof (–DD), 20 Oct 1960, Rensburg 2183 (PRE);
Stellenbosch, Waterkloof Farm (–DD), 25 Feb 1994, Nel & Boucher 71
(PRE). 3319 (Worcester): inter frutices ad flumen Baviaansrivier apud
Genadenthal (Genadendal), tum prope Waterfall haud procul a Tulbagh
(–AC), Ecklon s.n. (SAM); Winterhoek, Tulbagh (–AC), Zeyher 15652
(SAM); Mitchells Pass, near Waverley Mills (–AC), Nov 1879, Bolus s.n.
(BOL); Worcester Div., Buffelshoek, river bed (–AD), 29 Apr 1942,
Esterhuysen 7816 (BOL); banks of the Hex River (–BC), 11 Oct 1939,
Pillans 9064 (BOL, PRE); Fairy Glen, lower slopes of Audensberg Mts.
(–CB), 18 Sep 1981, Stirton 9122 (PRE); Worcester, at waterfall (–CB),
Dec 1830, Ecklon & Zeyher s.n. (PRE). 3321 (Ladismith):
Seweweekspoort in Klein Swartberg near Ladismith, Farm Aristata
(–AD), 18 Jan 1991, Henderson 1109 (NBG). 3322 (Oudtshoorn):
Boomplaas, Cango Valley, foot of slope below excavation site (–AC), 5
Feb 1975, Moffett 608 (NBG); Agric. Exp. Station, George (–CD), 18
Nov 1935, Baker s.n. (PRE); Oudtshoorn, Wilderness (–DC), 21 Oct
1990, Henderson 1097 (PRE); Wilderness, Fairy Knowe, at the Ebb and
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259 247

Fig. 3. Rubus rigidus agg. A, habit; B, primocane leaves; C, underside of leaf; D, detail of primocane; E, immature infructescence; F, flower; G, detail of flower with carpels.

Flow River (–DC), 13 Jan 1929, Mogg 11740 (PRE); sandy hills, Ruigte
Vlei, near Zwart River, Div. George (–DD), 10 Oct 1921, Fourcade 1528
(BOL). 3419 (Caledon): Caledon, Riviersonderend Valley (–AB), 18 Feb
1992, Henderson 1152 (PRE); in montibus Genadendal (–BA), Ecklon &
Zeyher s.n. (SAM); inter frutices ad flumen Baviaansrivier apud
Genadenthal [Genadendal], tum prope Waterfall haud procul a Tulbagh
(–BA), Ecklon s.n. (SAM); along R44 (Kleinmond to Bot River), N of
Arabella Country Estate (–AC), 22 Mar 2017, Sochor & Manning RSA27/
17 (NBG, OL); along R43, 6.2 km NNE of Fisherhaven (–AC), 29 Mar
2017, Sochor & Manning RSA40/17 (NBG, OL). 3421 (Riversdale):
Garcia's Pass (–AA), 20 Jan 1991, Henderson 1115 (BOL). 3422 (Mossel
Bay): Leermansdriftrivier, W of Plettenberg Bay (–BB), 12 Nov 1979,
Hugo 2100 (NBG, PRE). 3423 (Knysna): Knysna (–AA), Zeyher 15651
(SAM); Feb 1933, Duthie s.n. (PRE); road to Avontuur (–AB), 7 Oct
2011, Paijmans DMP136 (NBG); Plettenbergsbay [Plettenberg Bay]
(–AB), Jan [without year], Zeyher s.n. (SAM). EASTERN CAPE: 3323
(Willowmore): Buigpoort S of Willowmore (–AD), 19 Dec 1950, Theron
1006 (PRE). 3324 (Steytlerville): Kareedouw Pass, 12 Nov 1928, Hutch-
inson 1421 (BOL). 3325 (Port Elizabeth): Zuurberg National Park (–AC),
10 Jan 1986, Van Wyk & Van Wyk 1275 (PRE); prope Port Elizabeth, in
monte Zuurberg (–AD), 7 Sep 1930, Fries, Norlindh & Weimarck 709
(PRE); Zuurberg (–BC), Mar 1914, Holland 162 (PRE); near Van Staadens
Pass (–CC), 1949, Theron 568 (PRE); Uitenhage, Ado (–CD), Oct 1830,
Ecklon & Zeyher s.n. (PRE); prope Port Elizabeth, in Convalla Bethelsdorp
Kloof (–CD), 31 Aug 1930, Fries, Norlindh & Weimarck 303 (PRE); Target
Kloof, Baakens River (–DC), 7 May 1974, Olivier 1073 (NBG); Target
Kloof, Baakens River (–DC) Port Elizabeth, 30 Jan 1974, Olivier 957
(NBG); 7 May 1974, Olivier 1073 (PRE); Humewood, Port Elizabeth
(–DC), 30 Aug 1966, Dahlstrand 451 (PRE). 3424 (Humansdorp):
Clarkson (–AB), Oct 1926, Thode A822 (PRE).
248 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Fig. 4. Distribution of Rubus rigidus agg. in the Cape Floristic Region.
3. R. bergii × R. rigidushybridogenous complex
Shrub, usually 1–2 m high. Primocanes arching, terete to angular with
flat or (rarely) slightly furrowed sides, usually 7–10 mm diam.,
yellowish/greyish-green to dark vinaceous on side exposed to the sun,
densely to sparsely hairy with short stellate hairs to almost glabrous,
without stalked glands but sometimes densely covered by sessile glands;
prickles subequal or somewhat unequal, 5 to 21 per 50 mm of stem
length, usually strong and broad-based, sometimes slender and almost
subulate, straight to distinctly curved, declinate, 4–9 mm long and
2–9 mm broad at base, ± concolourous with stem or slightly reddish,
with yellowish tip. Stipules filiform-linear to narrowly lanceolate,
10–15 × ±1 mm, densely hairy with stellate and simple hairs. Leaves
on primocanes palmate or rarely palmate-pinnate, 5(−7)-foliolate but
ternate in terminal part of stem, usually deep green, subglabrous
above, whitish/greyish tomentose beneath, softly hairy to the touch; pet-
iole as long as basal leaflets or longer; terminal leaflet broadly ovate or
elliptic to almost orbicular with cordate (palmate leaves) or acute (pin-
nate-palmate leaves) base, apex acute to acuminate, often with one or
more acute lateral lobes, serration periodic, primary incisions mostly
1.5–2 mm deep, petiolule medium to long (30–55% as long as leaflet
lamina); lateral leaflets ovate to elliptic, sometimes slightly oblique,
petiolules (0–)1–6 mm long. Inflorescence paniculate, narrowly pyrami-
dal, cylindrical to broadly obovoid with long basal branches, commonly
with second main axis growing from base of primary inflorescence in
late season; branches erect to rarely erecto-patent; leaves 1 to 4, simple
or ternate, present only in lower quarter to half of inflorescence; bracts
lanceolate to trifid; inflorescence axis ± densely hairy with stellate
hairs, with 6 to 25 prickles per 50 mm axis length; prickles slender,
often broad-based, usually distinctly curved, declinate, 3–5(−7) mm
long; pedicels short, usually 4–10 mm long, densely hairy with long
hairs and sometimes with short stalked glands, with 1 to 4 slender,
curved prickles. Flowers many; sepals erect, spreading or slightly reflexed
after anthesis, 5–8 mm long, greyish tomentose and also with longer ±
patent hairs, with 0 to 2 (rarely more) small pricklets; petals pale to
deeper pink, rarely almost white, narrowly to broadly obovate to almost
orbicular, much longer than sepals (almost double their length), up to
±13 mm long. Stamens dark pink to pinkish, usually longer than styles,
inner series sometimes somewhat shorter. Carpels glabrous, styles rich
pink, stigmas yellowish pink. Aggregate fruit semiglobose, green turning
red and finally black at maturity (Fig. 5).
DNA-ploidy level: 5x (seven individuals analysed: Sochor & Manning
RSA18/17 [NBG, OL], RSA20/17 [NBG, OL], RSA28/17 [NBG, OL], RSA30/17
[NBG, OL], RSA39/17 [NBG, OL], RSA41/17 [NBG, OL], RSA44/17 [NBG,
OL]).
Distribution: common in the south-western part of Western Cape,
South Africa, especially between Paarl and Stellenbosch but not
recorded from the Cape Peninsula, with scattered records along the
south coast as far east as Knysna and near Kouga in Eastern Cape and
an isolated collection in the Cederberg in the northwest (Fig. 6); only
a few incomplete, and thus rather ambiguous specimens likely belong-
ing to other hybrid taxa, were seen from elsewhere (Eastern Cape, Gau-
teng, Limpopo).
Ecology: mainly on roadsides and other secondary habitats in both
shady and sunny situations, mostly in or near wooded vegetation (sec-
ondary forests, tree plantations, orchards etc.).
Diagnostic characters: similar to the Euro-Caucasian series Discolores,
e.g. R. armeniacus, in having palmate leaves that are white tomentose
beneath but differing in the spreading to erect sepals and sometimes
also by the occurrence of pinnately palmate (6- to 7-foliolate)
primocane leaves. Plants are further characterised by the dense inflores-
cence with distinctly curved prickles on the inflorescence axis, glabrous
carpels, and often curly, sharply serrate leaflet margins. From R. rigidus
agg., the complex differs mainly in absence of pinnate leaves, in having
subglabrous to sparsely (rarely densely) hairy but not distinctly tomen-
tose primocanes, and larger flowers with the petals much longer than
the sepals (vs. as long as or slightly longer). Herbarium specimens that
lack flowers and primocanes may therefore be difficult to determine.
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259 249

Fig. 5. Rubus bergii × R. rigidus. A, B, primocane leaves; C, detail of primocane; D, E, inflorescences; F, carpels and stamens.

This is a highly variable complex that shares morphological features
with both R. rigidus agg. (R. subg. Idaeobatus) and “true” blackberries
(R. subg. Rubus). Plants are often not, however, morphologically inter-
mediate between these two groups in many characters and have there-
fore usually been determined as either R. rigidus or ‘R. fruticosus’
(actually R. bergii) in herbaria. Despite the fact that this taxon is quite
common in the Boland of the Western Cape, no published name belong-
ing to it is known to us. The hybrid origin of the plants was confirmed by
our preliminary DNA sequence data, in which alleles of both putative
parental taxa were found at the nuclear ITS locus. Plastid sequencing
further revealed multiple independent origins from R. rigidus agg. serv-
ing as the pistillate parent. The second parent is a member of R. subsect.
Rubus, a group represented mainly in Europe and eastern North America
(Focke, 1910–1914; Kurtto et al., 2010). The molecular data imply a
close relationship to the European complex. The only species of this
group known so far in CFR is R. bergii (see below), which is therefore
the most probable candidate for the staminate parent. The common
occurrence of the complex, its apparent polytopic origin, and the spon-
taneous spreading of the complex in the region may imply that it has
passed through a period of independent evolution.
Additional specimens examined: South Africa. WESTERN CAPE: 3219
(Wuppertal): Cederberg, Matjiesrivier (–AC), Jan 1944, Wagener 319
(NBG). 3318 (Cape Town): Riebeek Kasteel (–BD), Oct 1958, Braam 1
(NBG); Dwarsriviershoek Farm, Banhoek on road to Kylemore School
(–DD), 17 Dec 1985, Morris 3 (NBG); Stellenbosch, Ida's Valley
Dam (–DD), 23 Aug 1973, Kok H29 (NBG); 0.6 km SW of Idasvallei
Dam (–DD), 21 Mar 2017, Sochor & Manning RSA18/17 (NBG, OL); Stel-
lenbosch, Vredenburg Farm, on dam wall (–DD), 17 Dec 1985, Morris 1
(NBG); Zevenrivierenpad, Banhoek (–DD), 17 Dec 1985, Morris 2
(NBG); along R45 between Simondium and Drakenstein (–DD), 21
250 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Fig. 6. Distribution of Rubus bergii × R. rigidus in the Cape Floristic Region.
Mar 2017, Sochor & Manning RSA21/17 (NBG, OL); Kylemore, along
roadside Stellenbosch to Pniel (–DD), 21 Mar 2017, Sochor & Manning
RSA20/17 (NBG, OL); in collibus prope Paarl (–DB), 16 Nov 1896,
Schlechter 9210 (BOL). 3319 (Worcester): Bain's Kloof, on mountainside
opposite Mr. Small's flower stall (–CA), 15 Nov 1980, Walters 2389
(NBG). 3321 (Ladismith): Riversdale, between Platkloof and
Kortefontein, foothills of Langeberg (–CC), 21 Jan 1991, Henderson
1116 (BOL, PRE). 3322 (Oudtshoorn): Karatara near Sedgefield (–DD),
21 Sep 2011, Paijmans DMP032 (NBG). 3419 (Caledon): along river mid-
way between Genadendal and Helderstroom (–AB), 29 Mar 2017,
Sochor & Manning RSA44/17 (NBG, OL); roadside between Sandbaai
and Caledon, Creation (–AD), 29 Mar 2017, Sochor & Manning RSA41/
17 (NBG, OL); along N2 between Patryslaagte and Houhoek (E of
Grabouw) (–AC), 29 Mar 2017, Sochor & Manning RSA39/17 (NBG,
OL); along R44 from Kleinmond to Bot River, N of Arabella Country
Estate (–AC), 22 Mar 2017, Sochor & Manning RSA28/17 (NBG, OL);
Elgin Valley, 10 km NNE of Kleinmond, amongst orchards at the stream
(–AC), 22 Mar 2017, Sochor & Manning RSA30/17 (NBG, OL);
Genadendal, crossroads on SW margin of the town (–BA), 29 Mar
2017, Sochor & Manning RSA42/17 (NBG, OL). 3422 (Mossel Bay):
Mossel Bay, Moordkuil River Valley, between Klein-Brakrivier and
Ganna Kraal (–AA), 19 Oct 1990, Henderson 1092 (BOL); Belvidere
(–BB), Jan 1926, A. Duthie 863a (NBG). EASTERN CAPE. 3324
(Steytlerville): Keerom (–CB), 1 Oct 1981, Stirton 9566 (PRE).
3.2.2. Naturalised taxa
4. R. armeniacus Focke in Abh. Nat. Ver. Bremen 4:183 (1874).
Type: ‘Ungarn, Armenien, Transkaukasien’ without date, W.O.
Focke s.n. (BREM, lecto. [not seen]), designated by Weber in
Abh. Westf. Mus. Naturk. 47(3): 222 (1985).
R. procerus sensu auct. mult., non P.J.Müller ex Boulay, Ronces Vosg.
1: 7 (1864).
Shrubs, under favourable conditions vigorous and able to form large
thickets, in W Cape usually not taller than 1 m. Primocanes arching,
rooting at apex, ±10 mm diam., distinctly angular with flat to furrowed
sides, green or vinaceous when exposed to sun, sparsely hairy with
short stellate hairs (especially young stems) to almost glabrous, without
stalked glands, with 4 to 9 prickles per 50 mm stem length arranged
along stem angles; prickles straight, erect or slightly declinate, strong
and broad-based, 6–9 mm long and 6–10 mm wide at base, usually
red and contrasting with green stem (living specimens). Stipules linear,
± 12–14 × < 1 mm, moderately to rather sparsely hairy with stellate
hairs to glabrescent adaxially. Leaves on primocanes palmate, usually
5-foliolate, leaflets usually touching or slightly overlapping one another,
dark green, subglabrous above, white/greyish tomentose beneath with
many short stellate and sparse long simple hairs, slightly hairy to
touch; petiole ± as long as basal leaflets, with scarce to scattered stellate
hairs and 6 to 15 slightly curved and strongly declinate prickles; termi-
nal leaflet with medium to long petiolule 25–45% length of lamina,
lamina broadly obovate or elliptic to suborbicular, base rounded or
very shallowly cordate, apex shortly acuminate (tip ± 10–15 mm
long), margins ± flat, slightly periodically to almost evenly dentate,
teeth broad, ± acute, incisions 2–4 mm deep; basal leaflets elliptic to
broadly elliptic with petiolules 2–7 mm long. Inflorescence usually richly
branched, broadly paniculate, ± pyramidal, truncate, inflorescence
leaves 1 to 11, simple or palmately 3- to 5-foliolate, present usually in
lower half to three quarters of inflorescence; bracts simple to deeply
trilobed, narrowly to broadly lanceolate; inflorescence axis angular,
hairy with dense subadpressed stellate hairs and longer patent hairs;
prickles 5 to 13 per 50 mm of axis length, mostly thin, straight to slightly
curved, usually somewhat declinate, 4–9 mm long, with wide red base;
pedicels 6–17 mm long, tomentose with adpressed stellate hairs and
with dense long patent simple hairs, with 5 to 15 unequal, thin, straight
to somewhat curved prickles 0.5–3.0 mm long. Flowers many; sepals
greyish tomentose with short stellate hairs and dense long simple
hairs, distinctly reflexed during and after anthesis, 5–8 mm long; petals
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259 251

pale to mid-pink, much longer than sepals, 15–20 mm long, broadly
elliptic to suborbicular, shallowly crenate and ciliate on margins. Sta-
mens much longer than styles; filaments white; anthers glabrous or
sparsely hairy. Carpels densely hairy; styles and stigmas glabrous, pink-
ish. Aggregate fruit large, usually longer than wide, green turning red
and finally black at maturity, juicy and sweet (Fig. 7).
DNA-ploidy level: 4x (two individuals analysed: Sochor RSA14/17
[NBG, OL], RSA34/17 [NBG, OL]; agrees with records of Krahulcová
et al., 2013).
Distribution: native to the southeastern Caucasus (Sochor et al., 2017)
but introduced as a fruit crop to Western and central Europe and western
North America, where it has become a significant invasive weed (Ingham,
2014). Some general sources report its occurrence also in South America,
Australia, New Zealand and South Africa, although rather tentatively and
without reference to primary data (Holub, 1995; Kurtto et al., 2010).
Many of these reports were later doubted due to close similarity of
R. armeniacus to, and the resulting frequent confusions with, other
European species (Rejmánek and Richardson, 2013). Except for three col-
lections from Swaziland (labelled as “R. affinis”), we found no specimens
of R. armeniacus in the herbaria that we searched, and our collections of
the species are the first documented for the CFR. Thus far we have
found the species only on Constantia Nek on the Cape Peninsula (Fig. 8).
Ecology: the preferred habitats of the species in its native range are
poorly known. In Europe, R. armeniacus occurs mostly along roads and
railways, on abandoned lands in cities and industrial areas, and in
other ruderal habitats (Weber, 2005), whereas in north-western
North America it spreads vigorously also in pastures, forests and ripar-
ian habitats (Ingham, 2014). On the Cape Peninsula, the most vigorous

Fig. 7. Rubus armeniacus. A, primocane leaf; B, fruits; C, old infructescence; D, detail of primocane; E, detail of inflorescence axis; F, carpels; G, flower. Photos F and G are of a specimen from
Olomouc, Czech Republic.
252 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Fig. 8. Distribution of Rubus armeniacus in the Cape Floristic Region.
shrubs were found along roadsides in Pinus or Eucalyptus plantations in
half-shaded situations but many smaller shrubs were also seen in adja-
cent disturbed fynbos vegetation.
Diagnostic characters: R. armeniacus resembles morphotypes of
the R. rigidus × R. bergii hybridogenous complex in its palmately
5-foliolate primocane leaves that are tomentose beneath but differs in
having straight, robust, red prickles (especially on the inflorescence
axis), usually a rather loose inflorescence with long patent or erecto-
patent branches, and flowers with reflexed sepals and larger petals
15–20 mm long.
R. armeniacus belongs to R. subg. Rubus ser. Discolores. It is also a
member of the so-called R. fruticosus agg. as usually understood, i.e. in-
cluding almost all European and Caucasian species from subgen. Rubus.
Nevertheless, delimitation of this highly diverse and species-rich taxon
is rather vague and artificial and its use is therefore not recommended.
Assessment of invasive potential: we have located no historical
records of Rubus armeniacus in the CFR and know it only from a single
locality covering less than 1 km2. It therefore does not seem to represent
a significant threat at this moment. Nevertheless, its locally common
occurrence at the margin of primary fynbos implies it is naturalised
and may spread in the future even into natural vegetation, particularly
in areas with suppressed fires.
Additional specimens examined: South Africa. WESTERN CAPE: 3418
(Simonstown): Hout Bay, along road from Kirstenbosch to Hout Bay,
0.5 km W of Constantia Neck (–AB), 18 Mar 2017, Sochor RSA14/17
(NBG, OL); 450 m NNE of Constantia Nek (–AB), 26 Mar 2017, Sochor
RSA34/17 (NBG, OL).
5. R. bergii (Cham. & Schltdl.) Eckl. & Zeyh., Enum. Pl. Afric. Austral. 2:
262 (1836). Rubus fastigiatus var. bergii Cham. & Schltdl. in Linnaea 2:
16 (1827). Type: South Africa, without precise locality or date, Ber-
gius s.n. (B?†, syn.). Neotype: South Africa, ‘Dornhoogde, in planitie
capensi’, Jun, K.L.P.Zeyher s.n. (P, neo. [not seen]), designated by
Van de Beek in Gorteria 36: 177 (2014).
R. vigorosus P.J. Müll. & Wirtg. Exsicc. Herb. Rub. Rhenan. 2: 53. 1860.
Type: Zw. Hillscheid und Höhr, 1 Jul 1859, F.P.Wirtgen Herb. Rub. Rhen. 2:
no. 53 (JE, lecto. [not seen]), designated by Weber in Bot. Jahrb. Syst.
106: 300 (1986).
R. affinis Weihe & Nees, Rub. Germ. 18 (1822), pro parte., nom. illeg.
superfl. pro R. fruticosus L. (see Weber, 1985).
R. fruticosus sensu auct. (incl. Bond and Goldblatt, 1984; Goldblatt
and Manning, 2000; Henderson, 2011; Manning and Goldblatt, 2012),
non L., Sp. Pl. 1: 493 (1753).
Shrub, usually 1–2 m tall. Primocanes arching, angular with flat or
slightly furrowed sides, ±5–7 mm diam, green to vinaceous on sides
exposed to sun, glabrous or nearly so, without stalked glands; prickles
almost uniform, 4 to 12 per 50 mm stem length, straight to slightly
curved, somewhat declinate, 7–10 mm long, 5–9 mm broad at base, suf-
fused red, with yellowish tip. Stipules narrowly lanceolate, 10–13 × ±
1 mm, ± densely hairy with simple hairs on both surfaces. Leaves on
primocanes palmately 5-foliolate, rarely incompletely palmate-
pinnately 6- or 7-foliolate, ternate in terminal part of stem, sun-leaves
three-dimensional with leaflets often concave and strongly rugose, leaf-
lets contiguous to overlapping one another, dark green and glabrous or
nearly so above, paler green or yellowish-green and sparsely to moder-
ately hairy (especially on veins) beneath, only slightly hairy to the
touch, without stellate hairs; petioles usually 50–70 mm long, as long
as or shorter than basal leaflets, sparsely to moderately hairy, without
glands, with 9 to 15 slender, strongly declinate and distinctly curved
prickles; terminal leaflet with relatively short petiolule ± 25–35%
length of leaflet lamina, lamina mostly ovate, cordate at base, apex grad-
ually or more abruptly acuminate, margins lightly crisped, serration
slightly periodic, with incisions mostly 2–3 mm deep; basal leaflets
ovate to elliptic, with petiolules 2–4 mm long. Inflorescence paniculate,
narrowly pyramidal or cylindrical, with erecto-patent branches, ± leaf-
less or with up to five leaves usually in lower quarter to two thirds of
inflorescence (late-summer inflorescence may differ); leaves palmately
4- or 5-foliolate to ternate, uppermost leaf simple; leaflets with similar
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259 253

serration and vestiture as on primocanes; bracts linear to lanceolate,
simple to trifid; inflorescence axis ± flexuous, moderately to densely
hairy (especially in upper part), with 5 to 13 prickles per 50 mm of
axis length; prickles slender, slightly to strongly curved, declinate,
5–7 mm long; pedicels usually 5–15 mm long, tomentose with stellate
and simple hairs, with 2 or 3(4) acicular prickles 2–4 mm long. Flowers
many; sepals 5–7 mm long, reflexed after anthesis, tomentose, (greyish-
)green beneath and white felted on margins, with 0 to 2 prickles; petals
pink to pale pink, broadly elliptic to orbicular, much longer than sepals,
±>10 mm long. Stamens longer than styles; filaments white, anthers
glabrous. Carpels glabrous, styles whitish or yellowish. Aggregate fruit
globose, green turning red and finally black at maturity, shiny, juicy
(Fig. 9).
DNA-ploidy level: 4x (three individuals analysed: Sochor & Manning
RSA17/17 [NBG, OL], RSA19/17 [NBG], Sochor RSA38/17 [NBG]; agrees
with the record of Krahulcová et al., 2013 [as R. vigorosus]).
Distribution: this species is native to north-western Europe
(Germany to Ireland and northern France to southern Sweden; Kurtto
et al., 2010 [as R. vigorosus]), and is an early introduction to South
Africa, from where it was later described. In South Africa R. bergii
appears to be concentrated in the Western Cape around old settlements,
including farms and mission stations, particularly on the Cape Peninsula
and the adjacent Boland and on the Piketberg, extending along the
southern coast to Knysna and inland on the Swartberg (Fig. 10). Its pos-
sible wider occurrence in Eastern Cape and other provinces (as reported
for R. fruticosus agg. in Henderson, 2011) needs to be determined.

Fig. 9. Rubus bergii. A, habit; B, primocane leaf; C, immature infructescence; D, flowers; E, mature fruits; F, carpels; G, underside of leaf.
254 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Ecology: R. bergii is the most common of the introduced bramble
species in Western Cape, and many stands are apparent remnants of
former cultivation as hedges or fruit crop plantings along roads,
amongst vineyards, and in villages or other settlements. It is neverthe-
less widely naturalised, and we have recorded probable or apparently
spontaneous occurrences in a variety of secondary habitats, including
roadsides, tree plantations, pastures etc., and also in indigenous bush
and forest margins, mostly with some degree of disturbance (roads,
waterworks, tourist paths) and a long history without fires, in both
shady and sunny situations.
It is likely that R. bergii was at least one of the brambles used by early
settlers in and around Cape Town as quickset hedges and fruit crops, as
recorded by Swedish botanist C.P. Thunberg during his travels in the
region in the late eighteenth century (Forbes, 1986). This raises the
intriguing possibility that some of the remnant plantings should be
considered as part of the cultural heritage of the region.
Diagnostic characters: similar to R. armeniacus and the R. bergii ×
R. rigidus hybridogenous complex in having palmately 5-foliolate
primocane leaves but differing in lacking tomentum beneath. The leaf-
lets are often plicate, usually with a long, distinctly acuminate tip. It is
a richly fruiting species, even in dry seasons, bearing medium to large
juicy berries. This was marked in several localities where we observed
the species growing together with R. rigidus × R. bergii, which set fruit
poorly or not at all.
Rubus bergii belongs to R. subg. Rubus ser. Rubus and is a member of
so-called R. fruticosus agg. (see also a note at R. armeniacus).
Assessment of invasive potential: due to its wide and common occur-
rence in both secondary and (semi-)natural habitats, R. bergii may
represent a potential threat. Significantly, it was the only species (native
or introduced) that we observed setting ripe fruits in very dry summers.
Nevertheless, we did not observe significant spontaneous stands in the
extreme south-west Western Cape, possibly due to the mostly infertile
soils and dry summer climate. Hybridization and introgression with na-
tive taxa might represent another potential threat. R. pinnatus × R. bergii
Fig. 10. Distribution of Rubus bergii in the Cape Floristic Region.
hybrids were encountered relatively frequently and some of them
appeared to be fertile. A different case is the R. rigidus × R. bergii
hybridogenous complex, in which the hybrids seem to be locally at
least as successful as either of the parents and possess a great invasive
potential. Although common deviations in shape of leaves and hairiness
were observed in both R. bergii and native taxa (R. pinnatus, R. rigidus
agg.), introgression has not yet been implied by plastid haplotype
patterns or any other approach.
Additional specimens examined: South Africa. WESTERN CAPE: 3218
(Clanwilliam): Piketberg, Avontuur, below Levant (–DD), 26 May
1974, Linder 336 (BOL). 3318 (Cape Town): Rhodes Estate (–CD),
30 Nov 1980, Stirton 8429 (PRE); Table Mt, Disa Gorge, near tunnel
(–CD), Dec 1921, Marloth 11046 (PRE); Kirstenbosch, behind Women's
Hostel (–CD), 25 Nov 1940, Compton s.n. (BOL); Table Mt (–CD), Dec
1919, Marloth 9412 (NBG); in dumetis inter mont. Leonis & Tabul.
(–CD), Jun, Zeyher 15655 (SAM); in dumetis ad Rondebosch (–CD),
Oct, Zeyher 15654 (SAM); inter frutices ad viam inter montes
Leeuwen- et Tafelberg (Cap) et prope Klapmuts (Stellenbosch) (–CD,
–DD), Ecklon & Zeyher 35401 (SAM); Kirstenbosch (–CD), Nov, Zeyher
s.n. (SAM); prope catarractas in latere orientali montis “Duyvelsberg”
(–CD), Oct Nov, Zeyher s.n. (SAM); Cape Town, Rondebosch, 750 m W
of Rhodes Memorial (–CD), 16 Mar 2017, Sochor & Manning RSA08/17
(NBG); Table Mt, Back Table, in front of the Overseer's Cottage (–CD),
12 Mar 2017, Sochor RSA03/17 (NBG, OL); Table Mt, Valley of Isolation,
±800 m NW of Waterworks Museum, at S oriented rock wall at a tour-
ist path (–CD), 11 Mar 2017, Sochor RSA01/17 (NBG, OL); Table Mt.
Woodhead Dam, 60 m W of Waterworks Museum (–CD), 26 Mar
2017, Sochor RSA38/17 (NBG); Wellington (–DB), 9 Aug 1982, Knobel s.
n. (PRE); Paarl Mt., 2 km S of summit, under Victoria Dam (–DD),
21 Mar 2017, Sochor & Manning RSA24/17 (NBG); Paarl Mt., at picnic
place 0.9 km NE of summit (–DC), 21 Mar 2017, Sochor & Manning
RSA23/17 (OL); along R45 between Simondium and Cillie (–DD),
21 Mar 2017, Sochor & Manning RSA22/17 (NBG); Kylemore, along
road Stellenbosch–Pniel (–DD), 21 Mar 2017, Sochor & Manning
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
RSA19/17 (NBG); Stellenbosch, base of Pappegaaiberg, S of summit
(–DD), 21 Mar 2017, Sochor & Manning RSA17/17 (NBG, OL); Stellen-
bosch, Devon Valley, along main road to Cape Town (–DD), 21 Mar
2017, Sochor & Manning RSA16/17 (NBG); Stellenbosch, oewer van die
[bank of the] Eersterivier (–DD), 31503, Wahl 68 (NBG); base of
Pappagaaiberg next to the river (–DD), 20 Oct 2011, Paijmans DMP161
(NBG); Elsenberg, dist. Stellenbosch (–DD), Sep 1903, Marloth 3304
(NBG, PRE). 3319 (Worcester): Ceres (–AD), Dec 1929, Thode A2236
(PRE); Skoongesig, Ceres, Koue Bokkeveld, 11 Jan 1968, Hanekom 1037
(PRE). 3322 (Oudtshoorn): Swartberg Pass, between Oudtshoorn and
Prince Albert (–AC), 19 Jan 1991, Henderson 1110 (PRE); George Dist.,
Silver River (–DC), Oct 1921, Fourcade 1582 (BOL). 3323 (Willowmore):
Uniondale Dist., Prince Alfred's Pass (–BB), 30 Nov 1942, Fourcade 5858
(BOL); Knysna Dist., Deepwalls (–CC), Dec 1923, Phillips s.n. (PRE);
Prince Alfred's Pass, between Avontuur and Kruisvallei (–CC), 1 Mar
1990, Henderson 1058 (PRE). 3418 (Simonstown): Bergvliet Farm, vlei
north of Tokai Road (–AB), 21 Nov 1918, Purcell 276 (SAM); Bergvliet
Farm, below upper brick pond (–AB), 16 Dec 1915, Purcell s.n. (SAM);
Wynberg (–AB), Dec 1907, Kensit 12978 (BOL). 3419 (Caledon):
±8.7 km NE of Kleinmond, Elgin Valley, amongst vineyards (–AC), 22
Mar 2017, Sochor & Manning RSA29/17 (OL); Genadendal, bushes
240 m SW of Moravian Church (–BA), 29 Mar 2017, Sochor & Manning
RSA43/17 (NBG, OL). 3420 (Bredasdorp): just outside Swellendam on
the road to Bonnievale, river crossing (–AB), 4 Dec 1991, Henderson
1145 (BOL). 3422 (Mossel Bay): Knysna Dist., Belvidere (–BB), 15 Feb
1933, Duthie s.n. (PRE); 1927, Duthie 1060 (NBG). 3423 (Knysna):
gully NE of Royal Hotel, Knysna (–AA), 22 Dec 1919, Schonland 3492
(PRE); Knysna forest (–AA), 20 Sep 1897, Galpin 3990 (PRE); Knysna vil-
lage (–AA), Dec 1922, Keet 1088 (NBG, PRE).
6. R. rosifolius Sm. [as rosaefolius], Pl. Icon. Ined. 3: 60 (1791). Type:
Mauritius, ‘Isle de France’ [Mauritius], Commerson s.n. (LINN
[HS902.63], holo.—image!).
Erect or climbing shrub, up to 2 or 3 m tall. Stems terete, softly hairy
with long patent hairs or subglabrous, dotted with sessile yellowish
glands, sparsely prickly, prickles erect, straight to curved, broad-based
± gradually tapering to apex. Stipules linear or lanceolate, softly hairy.
Leaves pinnately 5- to 7-foliolate, petiole ± as long as basal leaflets or
usually shorter, leaflets ovate or ovate-elliptic to lanceolate, base
rounded, apex acute to acuminate-attenuate, margin periodically (dou-
bly) serrate, distinctly incised with primary incisions ± 2.0–3.5 mm
deep, both surfaces pilose to subglabrescent; petiolules and leaf rachis
softly hairy and sparsely and minutely prickly, sometimes subglabrous,
petiolule of terminal leaflet rather short, 10–25% length of leaflet lam-
ina, petiolules of lateral leaflets very short, 1–4 mm long. Inflorescences
terminal or axillary, few-flowered; bracts linear or lanceolate,
puberulous; pedicels (10–)20–35 mm long, ± softly hairy and sparsely
and minutely prickly. Flowers 20–30(−50) mm diam.; sepals spreading
to slightly reflexed after anthesis, ovate-lanceolate, caudate, tomentose
Fig. 11. Rubus rosifolius, A, inflorescence; B, infructescence. Specimen from Sarawak, Malaysia.
255
and villous; petals white, obovate to suborbicular, somewhat shorter
than to slightly longer than sepals, shortly hairy abaxially. Stamens
shorter than styles. Carpels very numerous, glabrous. Aggregate fruit
compressed-globose, ovoid, obovoid to oblong, composed of very
many (more than 100) small drupelets, up to 20 mm long, green turning
red at maturity (Fig. 11).
Distribution: indigenous to south-east Asia and south-eastern
Australia but widely introduced in the tropics and subtropics, including
Hawaii, Polynesia, Melanesia, Caribbean, Reunion, Saint Helena (GISD,
2017) and Africa, predominantly in the eastern region, from KwaZulu-
Natal in South Africa to Zambia and Malawi (Launert, 1978;
Henderson, 2011). In the CFR, the species is known historically only
from the bottom of Platteklip Gorge above Cape Town (Fig. 12), where
it was first collected by C.F. Ecklon and C.L. Zeyher in the first half of
the nineteenth century and sporadically thereafter until the early twen-
tieth century but has not been recorded there since and can therefore be
considered extinct in Western Cape.
Ecology: in its primary distribution area, R. rosifolius grows on
rainforest margins, in wet eucalypt forests and in pastures (Bean,
1997). In its only historical locality in Western Cape, it was recorded
along a stream under trees.
Diagnostic characters: readily recognised by the few-flowered inflo-
rescences of large, white flowers 20–30(− 50) mm diam., pinnately
5- to 7-foliolate leaves that are pilose to subglabrescent on both sur-
faces, and aggregate fruit composed of very many small drupelets, up
to 20 mm long, red at maturity.
R. rosifolius is traditionally included in R. subg. Idaeobatus, although it
may be unrelated to the African taxa of the subgenus (R. rigidus agg.,
R. pinnatus, etc.; Alice and Campbell, 1999).
Assessment of invasive potential: the last herbarium record of
R. rosifolius in Western Cape is more than a century old and we could
not re-locate it despite the fact that apparently suitable habitats in
Platteklip Gorge are very limited in extent. Considering that a number
of planted ornamental species (e.g. Hibiscus sp., Opuntia sp.) are thriving
at the locality, we assume that this bramble has never naturalised there.
Additional specimens examined: South Africa. WESTERN CAPE: 3318
(Cape Town): Platteklip (–CD), Aug 1915, R. Marloth 7021 (BOL, NBG,
PRE); stream below Platteklip (–CD), Jun, F.C.Kolbe 3206 (BOL);
Table Mt, Platteklip, under trees on stream (–CD), May 1906,
R. Marloth 4338 (NBG); Platteklip, Table Mountain, (–CD), Penfold 9
(NBG); Platteklip (–CD), Jul, Ecklon & Zeyher 3586 (SAM); Table Mt
(–CD), Dümmer 1094 (SAM); in umbrosis silvis clivis montis Tabularis,
prope Platte Klipp (–CD), Apr 1908, Dümmer 1113 (PRE); in dumotis
ad viam in pede mount. Tabular (–CD), Ecklon & Zeyher 35386 (SAM).
7. R. titanus Bailey in Gent Herb I: 222 (1925). R. vitifolius var. titanus
(Bailey) Bailey, Sp. Batorum 1: 50 (1941). Type: California, Sebasto-
pol, Monitz s.n. (BH [92301], lecto.—image!), effectively designated
by Bailey in Sp. Batorum 1: 50 (1941). [Note: Bailey, 1925 did not
mention any specimens in the protologue for his R. titanus but
256 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Fig. 12. Historical distribution of Rubus rosifolius in the Cape Floristic Region.
cited Monitz s.n. from Sebastopol, CA under his later combination
R. vitifolius var. titanus (Bailey, 1941–1945). We regard this as effec-
tive lectotypification based on presumed original material.]
Common names: Mammoth Blackberry, Pacific Dewberry
Shrub, under favourable conditions vigorous and able to form canes
up to 13 m long (Bailey, 1941–1945), in Western Cape plants usually
smaller, arching, ±1.0–1.5 m high. Primocanes terete to indistinctly an-
gular, ± 5–8 mm diam., green to light vinaceous, with sparse to dis-
persed long simple hairs and sessile or shortly stalked glands, with
±35 to 70 prickles per 50 mm stem length; prickles strongly unequal,
straight, erect to slightly declinate, very slender thin with low broad
base, 1–7 mm long, up to 7 mm broad at base, ± concolourous with
stem or somewhat more reddish. Stipules narrowly lanceolate, 13–15
× ±1–2 mm, with dispersed to rather dense long simple hairs and dis-
persed to scarce sessile to shortly stalked glands. Leaves on primocanes
mostly ternate, sometimes palmately 4-foliolate, vividly green to partly
vinaceous in full sun, subglabrous above, sparsely hairy beneath with
long simple hairs, softly hairy to the touch; petiole markedly shorter
than basal leaflets, distinctly furrowed adaxially, sparsely villous and
with ±20 to 40 unequal prickles; terminal leaflet ovate, distinctly cor-
date at base, apex triangular acute and gradually tapering, margins
very coarsely dentate with serration irregularly periodic to almost
even distally, teeth often gradually merging into acute lobes up to
±20 mm long; petiolule of medium length, 20–30% of leaflet lamina,
lateral leaflets narrowly ovate with long triangular tip and with one
prominent accessory lobe on lower (basiscopic) side, petiolules rather
short, 2–7 mm long. Inflorescence sparse and few-flowered, leafy up to
top with ternate or simple leaves of similar shape as those on
primocanes but thinner and more coarsely dentate, branches erect to
erecto-patent, inflorescence axis ± densely hairy with long simple
hairs, with 6 to 25 prickles per 50 mm axis length, prickles unequal,
mostly straight and somewhat declinate, with short wide base,
1–4 mm long, up to 3 mm wide at base. Flowers few; pedicels
tomentose, (10–)20–45 mm long, with ± 7 to 17 mostly slender
unequal prickles up to 4 mm long; sepals tomentose, erect, spreading
to reflexed; petals and stamens not seen. Carpels hairy. Aggregate fruit
large, long cylindrical, of many large drupelets, black at maturity
(Fig. 13).
Ploidy level: 6x (two individuals analysed: Sochor RSA12/17 [NBG,
OL], Sochor & Manning RSA48/17 [NBG])
Distribution: this hybridogenous cultigen is not known from the
wild. Since its formation, it has been planted in California and probably
many countries with suitable climate worldwide (Waldo and Darrow,
1948). In South Africa it has been recorded only at two localities in
Western Cape, on the Cape Peninsula and naturalised in the Harold Por-
ter Botanic Garden at Betty's Bay (Fig. 14).
Ecology: the general ecological requirements of the species can only
be assessed based on its origin in California and its occurrence in Western
Cape. The CFR localities are in disturbed vegetation either along the road-
side between a vineyard and Eucalyptus plantation, or along a stream in a
botanic garden. The plants were, however, rather depauperate.
Diagnostic characters: distinguished from other Western Cape bram-
bles by its mostly ternate, non-tomentose leaves, numerous thin
straight unequal prickles, few-flowered inflorescence, and long, cylin-
drical aggregate fruits.
The taxon was derived by J.H. Logan in around 1883 from the cross of
“California wild blackberry” (probably octoploid R. vitifolius Cham. &
Schltdl.) and the ‘Texas Early’ (‘Crandall’, i.e. a variety of an unknown
east-American bramble, possibly R. flagellaris Willd., R. plicatifolius
Blanchard or some other tetraploid member of sect. Flagellares Bailey;
Bailey, 1941–1945; Waldo and Darrow, 1948, Water, 1901). Bailey
later reconsidered its status as a species and treated it as a variety of
R. vitifolius. Nevertheless, because of its hybridogenous origin and due
to the general avoidance of infraspecific ranks in modern batology, we
prefer the species status.
The plants from the Cape Peninsula and the Harold Porter Botanic
Garden correspond well to the description by Bailey (1925, 1941) but
 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Fig. 13. Rubus titanus. A, primocane leaves; B, detail of primocane; C, old infructescence; D, old fruit; E, habit.
another specimen that we found in the Kogelberg Nature Reserve near
the Visitors' Centre differed in its pinnate-palmate leaves, denser
prickles, acuminate leaflets with shorter tip, and in its ploidy level
(7x). Although it is likely closely allied to R. titanus, it is probably a hy-
brid of R. titanus and some species of R. subgen. Idaeobatus (possibly
R. pinnatus) and should therefore be treated as such.
Assessment of invasive potential: the degree of naturalisation in West-
ern Cape is unclear but is probably low as both occurrences may be
remnants of former cultivation. Although another population (not typi-
cal, see above) was located in a protected nature reserve in relatively
little-disturbed fynbos vegetation, it also does not appear to be
naturalised. Therefore, its invasive potential appears low at the
moment.
Additional specimens examined: South Africa. WESTERN CAPE. 3418
(Simonstown): Cape Town, along road from Kirstenbosch to Hout
Bay, behind the crossroads to Alphen (–AB), 18 Mar 2017, Sochor
RSA12/17 (NBG, OL); between Constantia Nek and Hout Bay (–AB), 9
Sep 1976, Stirton 6187A (PRE); Betty's Bay, Harold Porter National
Botanical Garden, 170 m NW of the entrance (–BD), along stream,
4 Apr 2017, Sochor & Manning RSA48/17 (NBG).
257
Anomalous specimen (perhaps hybrid R. titanus × R. pinnatus):
WESTERN CAPE. 3418 (Simonstown): Kogelberg Nature Reserve,
300 m W of Visitors Centre (–BB), 22 Mar 2017, Sochor RSA25/17
(NBG, OL); Southern Hottentots Holland Mountains, Kogelberg State
Forest (–BB), 6 Oct 1992, Kruger 825 (NBG, PRE).
3.2.3. Excluded species
R. pappei Eckl., Enum. Pl. Afric. Austral. [Ecklon & Zeyher] 2: 263
(1836). Type: South Africa, [Eastern Cape], inter frutices nemorum
Krakakamma et Ado, tum ad Zwartehoogdens et prope Katrivier
(Uitenhage, Albany), Ecklon s.n. (L[0019831, syn.—image!]). This speci-
men is probably not identical to R. pinnatus s. str. (cf. Harvey, 1862)
and may represent one of the eastern morphotypes of unknown taxo-
nomic value (see notes under R. pinnatus). Nevertheless, the original
description is inconsistent and the type specimen likely consists of
two different taxa. It is therefore provisionally excluded from our
treatment.
R. trifoliatus Suess. in Mitt. Bot. Staatssamml. München 2: 54 (1950).
Type: South Africa, [Western Cape], Franschhoek Pass, 17 Nov 1946,
S. Rehm sub W. Giess 1391 (M [0214132], holo.–image!). This species is
258 M. Sochor et al. / South African Journal of Botany 118 (2018) 241–259
Fig. 14. Distribution of Rubus titanus in the Cape Floristic Region.
based on a single incomplete collection. We are unable to match it with
any others from the region or elsewhere in South Africa and conclude
that it probably represents a North American taxon from the subgen.
Rubus. Its current occurrence in the region is questionable and we pro-
visionally exclude it.
Acknowledgements
We thank the curators of the studied herbaria for allowing us access
the Rubus material and Bram Van de Beek for consultations. Material
was collected under permit from CapeNature. Michelle Smith prepared
the electronic maps. This research was supported by grant No. LO1204
(Sustainable development of research in the Centre of the Region Haná)
from the National Program of Sustainability I, by project no. IGA PrF-
2018-001 from the Internal Grant Agency of the Palacký University,
and by a grant from the National Research Foundation, South Africa.
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