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Bothalia 32.1: 9-13 <2(X)2)
The genus Buglossoides (Boraginaceae) in southern Africa
E. RETIEF* and A.E. VAN WYK**
Keywords: Boraginaceae. Boraginoideae. Buglossoides Moench , Lithospermeae. nutlets, pollen, southern Africa, taxonomy
ABSTRACT
The genus Buglossoides Moench, a m ember o f the fam ily Boraginaceae. subfamily Boraginoideae. tribe Lithospermeae.
is revised tor the Horn o f southern A frica (FSA) region. Buglossoides comprises about seven species of annual, biennial or
perennial herbs and subshrubs native to Asia, southern Europe and northern Africa. B. anensis (L.) I.MJohn st.. a natural
ized weed in many parts o f the world, is the only member of the genus represented in the flora o f southern Africa, presum
ably introduced with imported cereal seed. Although the similar polyaperturate pollen grains of Buglossoides and
Lithospennum support the view that they are con generic, they are retained as separate entities here. Diagnostic characters,
a full description, various illustrations and a distribution map of B. an'e nsis in southern Africa are given.
INTRODUCTION
In 1794 Moench established the genus Buglossoides
(Boraginaceae/Boraginoideae/Lithospermeae), separat
ing it from the closely related genus Lithospemnun L.
Ever since, the generic status of Buglossoides has been a
matter of controversy. De Candolle (1846), Boissier
(1879), Giirke (1897), Wright (1904). Riedl (1967) and
Mabberley (1997) regarded the genus as congeneric w ith
Lithospennum. not accepting or unaware of Moench’s
view. In 1954 Johnston published a taxonomic revision
of Buglossoides. He used the corolla throat, decorated
inside w ith five well-developed vertical lines of hairs, to
distinguish Buglossoides from Lithospennum. which is
characterized by a corolla throat lacking these hairs but
bearing localized faucal appendages or groups of stipi-
tate glands. Buglossoides was accepted by various
authors like Ingram (1958). Fernandes (1972).
Edmondson (1978). Qaiser (1979 >. Brummitt (1992). Al-
Shehbaz (1991). Verdcourt (1991). Herman (1993).
Retief & Herman (1997). Lebrun & Stork (1997) and
Retief (2000). and this view is also taken here.
Buglossoides is a genus of about seven species, dis
tributed primarily in the European portion of the
Mediterranean region and in adjacent southeastern Asia,
with a single species indigenous to China. Korea and
Japan (Al-Shehbaz 1991). B. an ens is (L.) I.M Johnst. is
the only member of the genus represented in the flora of
southern Africa, scattered across the region. B. a n ensis
most probably reached southern Africa with imported
cereal seed. It is widespread as a naturalized weed in
many parts of the world. As yet it is not known from the
Flora zambesiaca region, but is recorded in Tanzania and
North Africa. Fernandes (1971. 1972) reduced three
Buglossoides species to subspecies o f B. a n ensis. How -
* National Herbarium. National Botanical Institute. Private Bag X 10 1.
0001 Pretoria.
** H.G.W.J. Schweickerdt Herbarium. Department of Botany.
University of Pretoria. 00 02 Pretoria.
MS. received: 2001 -06-18.
ever, typically B. ane n sis differs from these taxa in its
infundibuliform corolla (not hypocrateriform), a calyx
that usually equals or exceeds the corolla tube in flower
(not with calyx distinctly shorter) and in its distribution
(not confined to the Mediterranean region). It is here re
cognized as a separate species. One of the three species
concerned. B. sibthorpianum Griseb.. is regarded as con-
specific with B. a ne n sis by Edmondson (1978) and
Qaiser (1979).
De Candolle (1846) already cited Burchell 1686 under
Lithospennum an e nse L. Burchell collected his speci
men at Kloof village in the Asbestos Mountains.
Northern Cape in 1811. Wright (1904) in his revision of
Lithospennum in Flora capensis cited several localities
in South Atrica where 'L. an en se' had been collected.
Bolus & Wolley-Dod (1904) also mentioned the occur
rence of Buglossoides a n ensis because Wolley-Dod had
collected it at a few places on the Cape Peninsula.
Although Wright (1904) and Ross (1972) cited Gerrard
230 tor KwaZulu-Natal. B. anen sis is not recently
recorded from the region.
In this paper, diagnostic characters, a full description,
various illustrations and a distribution map of Buglos
soides a n ensis in southern Africa are given. The descrip
tion is based exclusively on local collections. This paper
forms part of a revision of the. Boraginaceae for the Flora
o f southern Africa (FSA) which is currently in progress.
MATERIALS AND METHODS
Herbarium specimens of Buglossoides arvensis
housed in BM. BOL. E. GRA. K. NBG. NH. N’MB. NU,
PRE. PRU and SAM w ere studied to assess morpholog
ical variation as well as phenological and distributional
attributes. Acetolysis of pollen followed the standard
method of Erdtman ( I960». For scanning electron micro
scope studies, samples were coated with gold and stud
ied with an ISI-SX-25 SEM. Measurements of pollen
grains were taken with the light microscope from acetol-
ysed grains mounted in glycerine jelly.
10 Bothalia 32,1 (2 002)
FIGURE 1.— Bug losso ides arvensis. A -C, rigid setae: A, on calyx lobes; B, on upper le af surface; C, basal part. D, E: inner corolla; D, lines of
hairs; E, close up o f hairs and p ollen grains. F, part o f annulus and pollen grains; G, nutlet; H, I, rugose, tuberculate nutlet surface. A-E,
H, I, Acocks 1774 0; F, Aco cks 19 035; G, Acocks 17805. Scale bars: 10 mm. A, 122 pm; B, 190 pm; C, 492 pm; D, 130 pm; E, 36 pm; F,
71 pm; G, 5 28 pm; H. 269 pm; I, 59 pm.
MORPHOLOGICAL CHARACT ERS OF TAXONOMIC
SIGNIFICANCE
Macromorphology
Species of Buglossoides are divided into two groups:
perennials with corollas 12-20 mm long, nutlets smooth
or punctate-reticulate, white or yellowish and shiny, and
annuals with corollas up to 10 mm long, nutlets tubercu
late, brownish or greyish and not shiny. B. arvensis
belongs to the latter group. The species is distinguished
from others in the genus by its calyx with a whitish indu
mentum and by the calyx lobes which are strongly
accrescent in fruit, narrow, acute and covered with long,
rigid setae (Figure 1A) and hairs (Leistner 2000: 759).
The indumentum of the leaves is appressed-setulose
with rigid setae, and the multicellular bases of the setae
are 1-layered (Figure IB, C). B. arvensis is distinguished
from all species of Lithospermum, and from all other
southern African representatives of Boraginaceae, by a
combination of the following characters: five longitudi
nal bands of hairs extending from the base of the corolla
lobe downwards to the tips of the anthers, as is typical for
the genus (Figure ID, E); an annulus of scale-like lobes
present near the base of the corolla tube (Figure IF); and
nutlets that are rugose and tuberculate (Figure 1G-I).
Pollen
Pollen of Buglossoides arvensis is isopolar and pro
late, with P = 12.5-18.8 pm and E = 8.3-12.6 (jm. In
equatorial view, the grains are rectangular to elliptic,
with the long sides more or less straight and the poles
convex, but frequently collapsing inwards (Figure 2A,
B). Ectoapertures are positioned at the equator, rhombic
in outline, regularly granular and the margins are not
thickened. Endoapertures are situated on or near the
equator and are ± oval and lalongate. The tectum is psi-
late. Orbicules are present.
Bothalia 32,1 (2002) 11
f t .
i *
i & j >,*>
M i.
m m m
* V f
r %
i © *
FIGURE 2.— Buglossoides arvensis. A , pollen grains; B, pollen grain with collapsed poles; C, granular aperture and psilate tectum. All grains ace-
toly sed. A, Ac ocks 17 805 ; B, C, Acocks 17 740. Scale bars: 10 mm. A, 4 pm: B. 2 pm: C, 1 pm.
The polyaperturate pollen grains of Buglossoides,
similar to those of Lithospermum, support the merging of
these genera. In their studies on pollen, Clarke (1977)
and Diéz et al. (1986) accepted Buglossoides as a sepa
rate genus. Ahn & Lee (1986), in a palynotaxonomical
study of the Korean Boraginaceae, did not accept
Buglossoides. Johnston (1954) divided the genus into
two sections and concluded that the pollen of section
Eubuglossoides (e.g. B. arvensis) is isopolar, whereas
pollen of section Margarospermum (e.g. B. purpuro-
caerulea (L.) I.M.Johnst.) is heteropolar and shows simi
larity to pollen of Lithospermum. Pollen morphology
thus suggests that whereas the section Margarospermum
can perhaps be included in Lithospermum , members of
the section Eubuglossoides are from Lithospermum and
best retained in Buglossoides. The white, smooth, shiny
nutlets of Buglossoides purpurocaerulea similar to those
of Lithospermum afrom ontanum Weim., for example, as
opposed to the rugose, tuberculate ones of B. arvensis,
support Ahn & Lee’s (1986) view. Heterostyly is absent
in B. arvensis and a vast amount of pollen is released on
to the corolla and between the lines of hairs (Figure
1D-F).
Pollen of southern African Boraginaceae is classified
into seven pollen types (Retief & Van Wyk 1999). Two
genera Lobostemon and Echiostachys, mainly confined
to Western Cape of South Africa, and the two species of
Echium introduced into southern Africa, are often
regarded as belonging to the tribe Lithospermeae in the
subfamily Boraginoideae (Johnston 1954). The triapertu
rate, heterocolpate pollen grains with a reticulate tectum
found in these genera are very different from those of
Buglossoides and Lithospermum , which are polyapertu
rate with a psilate tectum. Palynology supports the recog
nition of a separate tribe, Echieae, for Lobostemon,
Echiostachys and Echium.
Buglossoides Moench, Methodus plantas horti
botanici et agri marburgensis: 418 (1794); R.Fem.: 87
(1972); Al-Shehbaz: 129 (1991); Verde.: 79 (1991);
Retief: 180 (2000). Type: B. tenuiflora (L.f.) I.M.Johnst.
Lithospermum L.: 132 (1753), pro parte.
The genus name is derived from Buglossum Miller,
the name of a genus now reduced to a section of Anchusa
L,, and eidos = appearance, alluding to the superficial
resemblance of the two genera; the name Buglossum is
derived from the Greek bous. an ox and glossa, a tongue,
in reference to the broad, rough leaves (Al-Shehbaz
1991).
Buglossoides arvensis (L.) I.M.Johnst. in Journal
of the Arnold Arboretum 35: 42 (1954); Edmondson: 316
(1978); Qaiser. 82 (1979); Meikle: 1148 (1985); Tolken:
1152 (1986); Verde.: 79 (1991); Gibbons & Brough: 204
(1992); Retief & Herman: 350 (1997). B. arvensis (L.)
I.M.Johnst. subsp. arvensis sensu R.Fem.: 87 (1972).
Type: Europe, Herb. Linn. 181/4 (LINN, syn.-PR E,
microfiche!).
Lithospermum arven se L.: 132 (1753): A.DC: 7 4 (1846); Boiss.:
216 (1879); C.H.Wright: 24 (19 04); I.M.Johnst.: 7 (1927 ); Levyns: 688
(19 50); Levyns: 2 34 (1966); Jacot Guill.: 23 5 (1971); J.H.Ross: 298
(1972).
Illustrations: Qaiser: 83 (197 9); Tolken: 1153 (1986); Verdcourt: 80
(1991 ); Gibbons & Brough: 205 (199 2).
Annual herb up to 0.6 m high, greyish green, hispidu-
lous-strigillose. Roots purplish red. Stems erect or
decumbent, densely covered with appressed setae,
0.7-1.0 mm long. Leaves sessile, narrowly ovate or nar
rowly obovate, 15-45 x 3-8 mm. somewhat clasping at
base, apex acute or slightly obtuse, margin entire, dense
ly setulose; setae rigid, with multicellular, 1-layered
bases. Inflorescences terminal; cymes scorpioid,
bracteate. Calyx divided almost to base into 5 narrow
lobes; lobes with apices acute, strongly strigose, accres
cent, up to 12 mm long in fruit. Corolla white, infundibu-
liform: tube cylindric. 5-7 mm long, throat with 5 well-
differentiated, longitudinal bands o f hairs extending from
bases of corolla lobes downwards to tips of anthers;
lobes 5, rounded or truncate. ±2x1 mm; annulus near
base o f corolla tube of scale-like lobes. Stamens 5,
included, arising from corolla below middle of tube; fil
aments equal, shorter than anthers; anthers oblong, with
short apical appendage. Ovary 4-lobed. 4-locular,
gynobase disc-like; stigmas 2, subterminal. Fruit of 4
nutlets; nutlets erect, ovoid, flattened on dorsal side,
keeled on ventral side, with ± straight beak, 2.5-3.0 mm
long, rugose, tuberculate, attachment scar basal or nearly
so; fruiting pedicel ± 2.5 mm long. Flowering time:
August to December. Figure 3.
12 Bothalia 32,1 (2002)
FIGURE 3.— Buglo ssoides arv ensis.
A, habit, x 0.65; B, stigma, x
13.6; C, tuberculate nutlet, x
12; D, coro lla laid open, x
3.3; E, setae on upper leaf
surface, x 34.5. A, C -E, Acocks
19035; B, Acocks 17740. Illu
strations by Gillian Condy.
Distinguishing characters: annual herb, branching
from base, appressed hairy; corolla infundibuliform,
white, hairy inside; stigmas 2, subterminal; nutlets
rugose, tuberculate.
Distribution: North-West, Gauteng, M pumalanga,
Free State, Lesotho, Northern Cape, Western Cape,
Eastern Cape (Figure 4); a native of Eurasia, naturalized
as a weed in many parts of the world.
Habitat: weed of waste places; also in disturbed areas
such as in and along edges of cultivated lands.
Com m on nam es: sheepweed, white ironweed,
gromwell, com gromwell, grom well-comcockle, puc-
coon; naelbossie, naaldjiebossie, nagelbossie (Afrikaans)
(Smith 1966; Wells et al. 1986).
SPECIM ENS EXA MINE D (southern Africa on ly)
Acoc ks 17 740 K. PRE: 17805 PRE: 19035 K. PRE: 408 9 7 PRE. Acock s
á Roux 1594 8 K. PRE.
Bolus 101 BOL; 122 K; 274 GRA: 9201 BOL. B radfield 20 0 PRE.
Burchell 1686 K.
Diete rlen 1290 PRE.
Flanagan 162 9 K. PRE. Fourcad e 941 BOL, GRA.
Ge rrar d 23 0 K.
Holub s.n. K.
Jenkins TRV7232 PRE
Lee ndertz TRV 9748 PRE. Lov emore GRA-A1642 GRA.
MacChxan 19 26 GRA . Marloth 7216, 9670, P RE40898 PRE. Mogg
15292 PRE. Muir 790 PRE.
Schle chter 35 50 GRA. S harpe 9145 PRE. Silk 52 PRE. Smith 884 PRE.
Tyson 378 GRA. Trollope GRA-A1640 GRA.
Wardle PR E40893 PRE. Wilman 19817, P RE59952 PRE. Wilms 1006
BM. W olley-Do d 1551 K; 27 94 BOL; 3611 BM, BOL, K.
ACKNOWLEDGEMENTS
Dr O.A. Leistner, Miss G. Condy, Mmes E. du Plessis
Bothalia 32.1 (2<H)2) 13
FIGURE 4.— Known distribution o f Buglo ssoides arve nsis in the
Flora of southern Africa region.
and J. Ready are thanked for assistance with the prepara
tion of the manuscript. The directors, curators and staff
of cited herbaria are thanked for allowing the authors to
study their material and for sending specimens on loan.
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