Bothalia 35,1 (2005)
78
Omalycus (1814) predates Calvatia Fr. (1849) by 35 years,
and its adoption to cover species of Calvatia would require
a considerable number of new combinations, something
which is highly undesirable. Since Calvatia is already a
nomen consen’andum, it would be logical to add Omalycus
to the list of rejected names against it, which would not pre­
clude the use of Omalycus for a segregate including C.
cyathiformis. A formal proposal to that effect has been sub­
mitted to the journal Taxon.
AC KNO WLEDGEMENTS
The first author wishes to express his gratitude to Dr
B. Hein (B); Dr M. Pignal and Prof. P. Morat (P); and
Dr B. Buyck and Prof. A. Coute (PC) for their hospitali­
ty and friendly assistance during visits to the respective
institutions. Dr Vincenzo Migliozzi is thanked for pro­
viding us with a copy of his paper on Calvatia cyathi­
formis. We are also indebted to Dr V. Demoulin,
Universite de Liege, for his valuable comments on an
earlier draft of this paper. Funding received from the
Peninsula Technikon research committee is acknow­
ledged with gratitude.
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J.C. COETZEE* and A.E. VAN WYK**
* Department of Horticulture and Food Technology. Bellville Campus,
Cape Peninsula University of Technology. P.O. Box 1906, 7535 Bellville,
Cape Town.
** H.G.W.J. Schweickerdt Herbarium. Department of Botany, University
of Pretoria, 0002 Pretoria.
MS. received: 2004-05-20.
BORAGINACEAE
CODONOIDEAE, A NEW SUBFAMILY BASED ON CODON
The genus Codon was formally established by Carl
Linnaeus (1767) in the second volume of the 12th edition
of his Systema naturae. He placed the genus in his Class
X: Decandria, Monogynia. The generic name is derived
from the Greek word kodon, a bell (although the flowers
do not hang down), and alludes to the shape of the flow­
ers of C. royenii L., which are deeply cup-shaped. Codon
comprises two described species, C. royenii and C.
schenckii Schinz, both endemic to Namibia and South
Africa. A possible undescribed third species is found in
the southern part of Namibia and is currently under
investigation.
It was in France that a move towards more ‘natural’
groupings of plants was first made. It is clear from his
writings that Linnaeus recognized natural affinities, but
that ease of classification and identification were his
main objectives (Gunn & Codd 1981). Michel Adanson’s
Families des plantes (1763-64) can be regarded as the
first ‘logically and philosophically sound basis for a clas­
sification of plants’ (Stafleu & Cowan 1976). In 1789
Antoine-Laurent de Jussieu followed with his Genera
plantarum. He published the description of ‘Borragineae'
as one of 1(X) orders (i.e. families). Many of his families
are still maintained in modem classifications. De Jussieu
based ‘Borragineae’ on the genus Borago L. He divided
28 genera into three different groups using fruit mor­
phology as a distinguishing character: 1, berry-like fruits;
2, one- or two-locular capsules; and 3, four separate nut­
lets. He regarded Codon as a genus of uncertain position.
O f the five genera of Hydrophyllaceae known to him.
De Jussieu (1789) assigned Hydrophyllum L., Phacelia
Juss. and Ellisia L. to ‘Borragineae’ and Nama L. and
Hydrolea L. to ‘Convolvuli’. R. Brown separated the for­
mer trio of genera as the natural order Hydrophylleae in
1810, and the latter two as the natural order ‘Hydroleae’
in 1818. Choisy (1833) treated the Hydroleae in a mono­
Bothalia 35,1 (2005)
graph, recognizing the genera Hydrolea, Nama, Wigandia Kunth and Romanzoffia Cham.; to these he added
Eriodictyon Benth. in 1846, and at the same time vigor­
ously defended the distinctness of the Hydroleae. De
Candolle (1846: 589) was the first to place Codon in the
family Hydrophyllaceae. Gray (1875) united all the gen­
era mentioned in the family Hydrophyllaceae, which he
divided into four tribes. Baillon (1890) merged Hydro­
phyllaceae under Boraginaceae, but his view was not fol­
lowed at the time. Hydrophyllaceae was restored by Brand
(1913) in a monograph of the family.
Until recently, most authors accepted the Hydrophylla­
ceae as a separate family. A comparison between Hydro­
phyllaceae and Boraginaceae in southern Africa based on
pollen and macromorphological characters, however,
shows a strong overlap of features. The surface structure of
pyrenes of Ehretia P.Browne shows similarity with the
seeds of Nama (compare Retief & Van Wyk 2001: 15 and
Chance & Bacon 1984: 832), although this may not be
meaningful, as the outer surfaces in these two structures are
obviously not homologous. O f more significance is the
likeness between pollen grains of Wellstedia Balf.f. and
those of Eriodictyon, Nama and Phacelia (compare
Constance & Chuang 1982 and Retief & Van Wyk 2005);
tapetal orbicules or Ubisch bodies— sporopollenin particles
usually lining the inner tangential tapetal cell walls of secre­
tory tapetums— of Wellstedia and Codon show similarity in
morphology (Retief et al. 2001). The broad family concept
of Baillon (1890) is followed here, and we agree with the
Angiosperm Phylogenetic Group (APG) (i 998, 2003) and
Langstrom & Chase (2002) who regard Hydrophyllaceae
and Lennoaceae as synonyms of Boraginaceae s.I.
Modem views on the delimitation of Boraginaceae
differ, for example, 1, segregating a separate family,
Heliotropiaceae (Diane et al. 2002) from Boraginaceae
s./.; or 2, recognizing several segregate families: Bora­
ginaceae s. str., Cordiaceae, Ehretiaceae, Heliotropiaceae.
Hydrophyllaceae. Lennoaceae and Wellstediaceae (Lebrun
& Stork 1997; Gottschling et al. 2001; Gottschling 2003).
However, in neither of these two approaches has the
position of Codon been considered.
In Ferguson’s (1999) phylogenetic analysis of evolu­
tionary relationships within the Hydrophyllaceae, it is
concluded that the family is nested within a paraphyletic
Boraginaceae s.I., excluding Codon and Hydrolea.
Hydrolea is placed in a family of its own (APG II 2003).
Codon is included in Boraginaceae s.I. in a treatment of
this family for the Cape flora (Retief & Buys 2000: 374,
706). The genus Codon has traditionally been assigned to
the Hydrophyllaceae, where it seems to be unusual geo­
graphically, as the family is otherwise largely restricted
to the New World. Similarities between Codon and other
members of Boraginaceae indicate that Codon should be
placed in a subfamily of its own within Boraginaceae s.I.
A new subfamily, Codonoideae, is established here to
accom m odate this southern African genus within
Boraginaceae s.I., the other local subfamilies being Wellstedioideae. Ehretioideae. Cordioideae. Heliotropioideae
and Boraginoideae (Retief 2(XX): 179; Retief & Van Wyk
2001). The precise classification of the other, mainly
New World genera (see Brand 1913) of Hydrophyllaceae
s.str. within Boraginaceae s.I. has not yet been addressed
79
but they would most probably also require placement in
one or more additional subfamilies.
Phytogeographically the restriction of Codon to the arid
southwestern comer of Africa is of special interest. Its nearest
relatives appear to be those members of Boraginaceae s. I. pre­
viously placed in Hydrophyllaceae s. str., found mainly in
North America, but with members of both Nama and
Phacelia also occurring in South America (Deginani 1999).
Among plants, African-New World distributions are rather
unusual, but for southern Africa, involve as many as seven
families and many more genera (Goldblatt 1978). It is intrigu­
ing that several other plant groups with a distribution pattern
comparable to that of Codon, namely with a disjunct presence
in southwestern and in northeastern Africa, show links with
taxa in the New World. Among these are the boraginaceous
genus Wellstedia as well as members of Calliandra, Caesalpinia, Haematoxylon. Hoffinannseggia. Parkinsonia, Xerocladia (all Fabaceae), Nicotiana (Solanaceae), Thartmosma
(Rutaceae) and Tumera (Tumeraceae) (Van Wyk & Smith
2001). These African-New World disjuncts may have been
established in different ways, but one possible explanation is
based on the proximity of Africa and South America during
Gondwana times and for quite some time after the break-up
of the supercontinent.
Members of the new subfamily Codonoideae, estab­
lished here, show the strongest affinity with the subfamilies
Wellstedioideae and Ehretioideae in Boraginaceae s.l.— in
similar pollen, inflorescence and trichome morphology.
However, the Codonoideae differ from Wellstedioideae in
the 10 or 12 (not 4) corolla lobes and in the seed which is
subglobose and glabrous (not truncate and pubescent). They
differ from Ehretioideae in habit in that they are annual or
short-lived perennial herbs (not shrubs or trees) and in the
fruit which is a capsule (not a drupe).
C odonoideae Retief & A.E. van Wyk, subfam. nov.
Type: Codon royenii L.
Herbae annuae vel breviter perennes, patentes ad erectae.
interdum basi lignescentes. Partes vegetativae aculeatopubescentes trichomatibus spiniformibus, et setis trichomatibusque multicellularibus non ramosis. Folia
petiolata. Flores cymis scorpioideis, ebracteati. Calyx
profunde lobatus, lobi lineares. Corolla 10- vel 12-lobata. tubus cylindricus vel campanulatus, fauce glabra.
Stylus terminalis, linearis, paene ad medium fissus, persistens; stigma capitatum. Fructus capsula multiseminalis. Semina globosa vel irregulariter angulosa, omata.
Spreading to erect, annual or short-lived perennial herbs,
sometimes woody at base. All vegetative parts prickly
pubescent with spine-like trichomes, also with setae and
unbranched, multicellular trichomes. Leaves petiolate. In­
florescence scorpioid. Calyx deeply lobed. lobes linear.
Corolla 10- or 12-lobed. tube cylindric or campanulate.
throat naked. Style terminal, linear, cleft to almost halfway,
persistent; stigma capitate. Fruit a capsule, many-seeded.
Seeds globose or irregularly angled, ornamented.
Genus: Codon L.
ACKNOWLEDGEMENT
We wish to thank Dr O.A. Leistner for translating the
subfamily description into Latin.
Bothalia 35,1 (2005)
80
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E. RETIEF*+and A.E. VAN WYK**
* National Herbarium, South African National Biodiversity Institute,
Private Bag X 101, 0001 Pretoria.
t Student affiliation: Department of Botany. University of Pretoria,
0002 Pretoria.
** H.G.W.J. Schweickerdt Herbarium. Department of Botany, Univer­
sity of Pretoria. 0002 Pretoria.
MS. received: 2004-06-30.
POACEAE
NOTES ON ERAGROSTIS
A v a ria n t of Eragrostis gummiflua Nees?
Eragrostis gummiflua occurs in Botswana. Lesotho,
Mozambique, Namibia, Swaziland, South Africa and
Zimbabwe, usually on sand. Up to now, it has been one of
the easier Eragrostis species to identify in these regions as
it is a perennial with large, sticky, glandular patches below
the collar on the leaf sheaths and often at the nodes as well.
Sand grains or other pieces of material usually stick to these
areas, making them easy to see. The nodes and area below
the collar are often flushed purple, though sometimes the
glandular patch below the collar is yellow or brown. The
spikelet is purple to straw-coloured, with distinct, thick
nerves on the lemmas. At maturity, the palea and lemma
curve away from each other, leaving only their bases and
apices touching and then resembling the pincers of a crab.
In December 1985 the author collected specimens in
northeastern KwaZulu-Natal (2732 BA) that bore a close
resemblance to E. gummiflua but without any indication
of sticky glandular patches. A further search in the PRE
(National Herbarium, Pretoria) collection yielded two
more specimens without these glandular patches, one
from the Manzibomvu area (2732 DA) and a another
from southern Mozambique between Bela Vista and
Umbeluzi (2632 ?). The main differences between these
specimens and E. gummiflua are provided in Table I.
To date, the non-sticky specimens seen by the author are
from the biogeographical region known as Maputaland, an
area that has been identified as an important centre of
endemism and biodiversity in southern Africa (Siebert et al.
2004). It is bound by the Inkomati-Limpopo River in the
north, the Indian Ocean in the east, the Lebombo Mountains
in the west and by the St Lucia estuary in the south. Much
of the area is a flat, low-level coastal plain with infertile
soils consisting of geologically recent fine-grained aeolian
sands. Climatically it lies within a transitional zone between