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Nord. J. Bot. - Section of tropical taxonomy
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The genus Colpodium (Gramineae) in Africa
0. Hedberg and I. Hedberg
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Hedberg, 0. & Hedberg, I, 1994. The genus Colpodium (Gramineae) in Africa. - Nord.
J. Bot. 14: 601-607. Copenhagen. ISSN 0107-055X.
Recent collections of the genus Colpodium from Ethiopia and South Africa made a
renewed revision of the African material of this genus desirable. Apart from the two
earlier recognized African species (C. chionogeiton and C. hedbergii) a third one is
described (C. drakensbergense), restricted to the Drakensberg area of eastern Lesotho
and amply distinct both in morphology and chromosome number. The phytogeographical position of Colpodium is briefly discussed.
0. Hedberg and I. Hedberg. Department of Systemutic Botany, Uppsala University,
Villuvagen 6, S-752 36 Uppsala, Sweden.
new revision of the African species, a need which was
accentuated by the work on the Flora of Ethiopia.
Introduction
When Tzvelev (1964) published his revision of the genus
Colpodium Nevski, I9 species were known to him; later 6
more have been described (Bor 1970; Malyschev 1971).
He also made a subdivision of Colpodium into five subgenera (Hyulopoa Tzvel., Puracolpodium Tzvel., Nevskia Tzvel., Catubrosella Tzvel., and Colpodium s. str.).
In a later publication three of these were segregated as
independent genera (Tzvelev & Bolchovskieh 1965).
Both Colpodium s. str. and Puracolpodium were later
revised by Alexeev (1980, 1981). Clayton & Renvoize
( 1986) reestablished the genus Colpodium in its earlier
wide sense, and their circumscription of the genus is
followed here.
The discovery of the somatic chromosome number 8 in
grass collections from Mts Kenya and Kilimanjaro referred to Agrostis chionogeiton Pilger, and in a collection
of a closely related taxon from Mt Elgon (0.Hedberg
1952) caused Melderis (1956) to create the genus Keniochlou to harbour these taxa. Later Tzvelev (in Tzvelev &
Bolchovskieh 1965) united Keniochlou with Colpodium
Nevski s.str. In his account for Flora of Tropical East
Africa Clayton (1970) accepted this transfer and provided
a detailed revised account. The discovery of this genus
both in Ethiopia (0. Hedberg, unpublished) and South
Africa (van Zinderen Bakker & Werger 1974) prompted a
Accepted 16-6-1994
0
Material and methods
The present revision is mainly based on herbarium material obtained on loan from EA, K, PRE, S and UPS
(abbreviations according to Holmgren et al. 1990), but it
was also possible to study specimens in their natural
habitats both in Ethiopia and in Lesotho. The figures used
for the diagrams represent means of measurements of five
spikelets and (as far as possible) five panicles. Chromosome counts were partly made in sections of root tips (cp.
0. Hedberg 1952), partly in squash preparations (cp. I.
Hedberg 1970: 154).
Results and discussion
Morphology
The Tropical African taxa treated here are - like those
Eurasian species with which they were united by Tzvelev
& Bolchovskieh (1965: 1320) in the subgenus Colpodium
- characterized by one-flowered spikelets with soft tex-
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NORDIC JOURNAL OF BOTANY
NORD. J . BOT. 14: 601407
39
Nord. 1. Bot. 14 ( 6 ) (1994)
60 1
I
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1
I
2
I
3
I
I
4
5
Fig. I . Scatter diagram illustrating correlated variation in lemma
length in mm (vertical axis) and palea length (horizontal axis) in
a representative material of Colpodium chionogeiton (dots) and
C. hedbergii (circles). Collections measured were of the former:
Bie 35, 117, 135 and 141, Hedberg 1253, 1902 and 3759, King
6; of the latter: Hedberg 5361, 5492, 5618 and 5654, and
Phillips 28; all at UPS.
J-
0
0
0
4-
0
0
3-
The somatic chromosome number 8 reported by 0. Hedberg (1952) for Colpodium (Keniochloa) chionogeiton
and C. hpdbergii was confirmed in four additional collections of the latter species from Ethiopia (Tab. 1 and Fig. 4
A). Material from the Drakensberg in Lesotho, on the
other hand, deviated not only in basic number but also in
ploidy, with the presumably tetraploid somatic number
20 (Fig. 4 B).
It is interesting to note that the Eurasian species C.
versicolor (Stev.) Woron., considered by Tzvelev & Bolchovskieh (1965) most closely related to the African
species, approaches them in its basic chromosome number, which according to Sokolovskaja & Probatrova
(1977), Gviniadnidze & Avaznelii (1982) and Davliadnidze (1985) is 2 - half of that found in the African
species, and the lowest number found in angiosperms.
From the cytological and morphological data summarized above it is evident the the Drakensberg material
represents a distinctive species, characterized by its deviating chromosome number as well as by it decumbent
habit and its short and unequal glumes (Fig. 3). It is
described below as Colpodium drakensbergense.
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2-
1-
Chromosome numbers
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6-
5-
ture, awnless lemma with nerveless tip, and frequent
occurrence of clavate-tipped hairs on palea and lemma.
The main morphological characters used for specific
distinction in the tropical African material by Melderis
(1956) and Clayton (1970) were panicle shape (contracted in C. chionogeiton, open in C. hedbergii) and
spikelet length (longer than 3.5 mm in C. chionogeiton,
shorter than 3.5 mm in C. hedbergii).
In the material earlier known there seemed to be a
clear-cut distinction between these species both in panicle shape and in spikelet length, but some recent collections deviated so much that it was tempting to question
interspecific distinction. Further detailed studies disclosed, however that the two taxa also differ in floret
morphology: in C. chionogeiton the lemma is distinctly
longer than the palea, whereas in C. hedbergii they are of
comparable length (Fig. I). The combined variation pattern summarized in Fig. 2 therefore supports interspecific
distinction between them.
The South African material is distinguished from that
of Tropical Africa by its decumbent and rooting stem and
by its distinctly unequal glumes, shorter than palea and
lemma (Fig. 3).
Fig. 2. Pictorialized scatter diagram illustrating correlated variation in some characters studied in tropical African material of
Colpodium: spikelet length in mm (vertical axis) and ratio of
panicle length to panicle width (horizontal axis). An open circle
corresponds to a specimen where palea and lemma are equilong
(C. hedbergii). a filled circle to a specimen with palea and
lemma of unequal length (C.chionogeiton). A tail means that at
least some panicle branches are reflexed.
602
Ecology
All three African Colpodium species are definitely depending on a high water level, either along streams or in
small shallow pools, and are restricted in East Africa to
the afroalpine belt, in Lesotho to the austroafroalpine
belt.
Nod. J. Bot. 14 (6) (1994)
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4
I
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Fig. 3. Colpodium drakensbergense drawn from Hedberg 82009 (UPS). - A. habit x 1.3. - B. Ligule x7. - C. Spikelet x 15. D. Flower x30. - E. Lodicules x30. - F. Palea x 15. - G. Lemma x 15. - H. Upper glurne x 15. - I. Lower glume x 15.
39' Nod. J. Bot. 14 (6) (1994)
603
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Tab. I . Chromosome numbers in African Colpodium (cp. 1. Hedberg & 0. Hedberg, 1977)
Taxon
2n
C. chionogeiron
8
C. hedhergii
8
C. drukensbergense
20
Locality
Voucher
Mt Kenya, 4250 m
Hedberg 1902
Hedberg 1759
Hedberg 1253
Bie 35
Hedberg 908
Hedberg 5361
Hedberg 5492
Hedberg 5645
Hedberg 56 I8
Hedberg 82088
Mt Kenya, 4450 m
Kilimanjaro, 4830 m
Kilimanjaro, 3800 in
Mt Elgon, 3600 m
Simen Mts, 3600 m
Simen Mts, 3700 m
Bale Mts, 4100 m
Bale Mts, 4150 m
Drakensberg 2900 m
Phytogeography
The genus Colpodium s. lat. occurs from Turkey and
Caucasus to Nepal and eastern Siberia, extending southwards through the Ethiopian and East African high
mountains to the Drakensberg in Lesotho (Fig. 5). Since
the diversity of the genus is much larger in Asia than in
Africa - from Iran alone 12 species have been reported
(Bor 1970) - the actual distribution pattern must obviously have resulted through dispersal from the north. The
southward migration has evidently been accompanied by
a rise in basic chromosome number, to 4 in East Africa
and 5 in the Drakensberg. The karyotypes are not very
helpful in tracing this evolution, however, each of the
three African species has two SAT chromosomes and the
other chromosomes are more or less metacentric with
little size variation (Fig. 4).
Additional support for Iran and surrounding countries
as centre of diversity (if not origin) of this group is the
occurrence there of the genus Zingeriu P. Smirn., which
comprises annual species only and was believed by Tzvelev & Bolchovskieh (1965) to be closely related to and
perhaps derived from Colpodium subgenus Colpodium.
Bjorkman ( 1 956) also found great similarities in lemma
epidermis and other spikelet characters between Zingeria
and “Keniochloa” (cp. Bjorkman 1960). It is particularly
interesting in this connection that Zingeria also resembles
Colpodium subgenus Colpodium in the occurrence of the
extremely low basic chromosome number 2 (Sokolovskaja & Probatova 1979; Clayton & Renvoize 1986).
In the afroalpine flora Colpodium belongs to the Himalayan flora element (0.Hedberg 1965, 1986), whereas in
the austro-afroalpine flora of the Drakensberg it is
brought to the “northern temperate group” (van Zinderen
Bakker & Werger 1974: 47).
Taxonomy
Colpodium Trinius
zy
(1820: 1 19); Tzvelev 1964: 5-19; Clayton 1970: 49; Alexeev
1980: 4; Clayton & Renvoize 1986: 103. - Type species: Cul-
podium versicolor (Stev.) Schmalh.
Keniochlou Melderis 1956: 538; 0. Hedberg 1957: 53. Type: K. chionogeiron (Pilg.) Meld.
Cutuhrosellu (Tzvel.)Tzvelev 1965: 1320. -Type: C. hurnilis
(Bieb.) Tzvel.
Hvu/(ipoa (Tzvel.) Tzvelev loc. cit. - Type: H. pontica (Bal.)
Tzvel.
Paracolpodium (Tzvel.) Tzvelev loc. cit.; Alexeev 1981: 86.
- Type: P ulruirum (Trin.) Tzvel.
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Fig. 4. Metaphase plates from root tip squashes of Colpodium,
x 1200. - A: C. hedbergii; - B: C. drukensbergetue. - A from
Hedberg 5492 (Ethiopia, Simen), B from Hedberg 82088
(Lesotho, Sani Pass).
604
Small perennials. Leaves flat or folded, with boat-shaped
tip. Panicle spiciform or spreading. Spikelets in African
species 1-flowered, in some Asian species up to 4-flowered, awnless, with persistent firmly membranous
glumes with hyaline margins and obtuse or subacute tip.
Nord J. Bot. 14 ( 6 ) (1994)
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Fig. 5. Known distribution
of the genus Colpodium in
Africa. Filled circles C. chionogeiron, open circles
- C. hedbergii, triangle C. drakensbergense.
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Key to species in Africa
1. ~l~~~~ distinctly unequal,
than lemma and
palea; stems decumbent, rooting at the nodes . . . . . .
.......................
3. C.drakensbergense
- Glumes equal, overtopping lemma and palea; stems
erect and tufted . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2
2. %)jkelets at least 3.7 mm long; panicle usuah'
contracted with all branches erect and lengtwwidth ratio
Nord. J. Bot. 14 ( 6 )(1994)
above 3; palea distinctly shorter than lemma. . . . . . .
..........................
1. C. chionogeiton
Spikelets at most 3.5 mm long; fully developed
panicle open with at least some branches reflexed and
lengthlwidth ratio below 3; palea and lemma equal or
subequal ......................
2. C. hedbergii
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Lemma thinly membranous, 3-5-nerved, glabrous or pubescent with appressed clavate-tipped hairs.
The genus comprises about 20 species, occurring from
Turkey and Caucasus to Nepal and eastern Siberia, and
furthermore on African high mountains from Ethiopia to
Lesotho. Restricted to high mountain areas (Fig. 5).
-
1. C. chionogeiton (Pilg.) Tzvel.
Tzvelev in Tzvelev & Bolchovskieh 1965: 1319; Clayton 1970:
5 I , Fig. 18; Alexeev 1980: 8, Fig. 2: 1-6. - Agrostis chionogeiton Pilger 1926: 5 10. - Keniochloa chionogeiton (Pilg.) Melderis 1956: 540, Fig. I : 2; 0. Hedberg 1957: 53. Type: Kenya, Mt
Kenya, w alpine region, Fries 1400a (S holotype, K, UPS
isotypes).
Agrostis oreades Peter 1930: 87. - Keniochloa chionogeiton
var. oreades (Peter) Melderis 1956: 540, Fig. 1: 3; 0. Hedberg,
1957: 53. - Colpodium oreades (Peter) Alexeev 1980: 8, Fig. 2:
605
7-1 I . -Type: Tanganyika, Kilimanjaro, Peter’s Hut to Mawenzi
saddle, Peter 41928 (B holotype, not seen).
appressed pubescence of clavate-tipped hairs. Palea equilong with lemma.
Tufted perennial with erect or ascending culms 4-40 cm
high. Leaf blades flat or folded, up to 17 cm long and 4
mm wide. Ligule 2-6(-14) mm long. Panicle usually
contracted, often spike-like, 1.5-1 2 cm long, its basal
part often concealed in the slightly inflated sheath of the
uppermost leaf. Spikelets 3.7-6.5 mm long, narrowly
elliptic, light green to purple. Glumes equal, longer than
lemma. Lemma 3.3-4.9 mm long, 3-nerved, glabrous or
with appressed clavate hairs. Palea 2.9-4.2 mm long,
distinctly shorter than lemma.
Chromosome number. 2n = 8 (cf. Tab. I). Fig. 4A.
Distribution. Ethiopia, Gondar, Simen, Geech, Hedberg
& Getachew 5492,5361 (EA, ETH, FI, K, PR, ROML, S,
SRGH, UPS). Bale, Bale Mts National Park: on the crest
S of Garba Goracha camp site, Hedberg 5645 (EA, ETH,
K, PR, UPS); Saneti Plateau, Hedberg 5618 (EA, ETH,
FI, K, PR, ROML, S, SR, GH, UPS) and 9067 (UPS), S.
M. Phillips 28 (ETH, K, UPS).
Kenya, Mt Elgon, at Maji ya Moto, Hedberg 908 (EA,
K, S, UPS). Mt Kenya, Teleki valley, Aichhorn 7 (EA);
Kaziba valley, Hanid 157 (EA).
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The population on Kilimanjaro was described by Peter as
a separate species (Agrostis oreades), treated by Melderis
as a variety of Keniochloa chionogeiton, and by Alexeev
as an independent species, Colpodium oreades. It was
claimed by Alexeev to differ from C. chionogeitiun by
having the lemma more or less hairy in the lower part,
glumes 1-3-veined, and anthers 1.2-1.3 mm long (as
against glabrous lemma, glumes 3-nerved and anthers
0.6-0.8 mm in C. chionogeiton on Mt Kenya). Investigation of a larger material demonstrated that the pubescence
of the lemma is too variable to be of taxonomic use,
however. In all collections examined the lower glume is
practically always I-nerved, the upper one 3-nerved. The
only difference remaining is that the anthers studied measured 0.9- l .2 mm in Mt Kenya material and l .O-2. l mm
in Kilimanjaro material, a difference which does not
merit taxonomic recognition.
Habitat. In shallow water along streamsides or in small
pools, 3 6 0 W l S O m.
3. Colpodium drakensbergense 0. & I. Hedberg n.
SP.
C. hedbergii affine sed glumis distincte inaequalibus palea brevioribus, caulibusque repentibus nodis radicantibus praecipue
differt. - Orig. coll. Lesotho, Qacha’s Nek Distr., Sani Pass
lodge, 2860 m, in small dried-out pools, Hedberg 82088 (UPS
holotype, EA, ETH, K, PRE, ROML isotypes).
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Chromosome number. 2n = 8 (cf. Tab. 1).
Distribution. Kenya, Mt Kenya, alpine region, Fries
1400a (S); Hedberg 1759, 1902 (EA, K, S, UPS).
Tanzania, Kilimanjaro, Mawenzi, Hedberg 1253 (EA,
K, S, UPS); King 6 (EA, UPS); saddle between Kibo and
Mawenzi, Goyns 18 (PRE); W slope in upper Umbwe
Valley, Bie 35, 117, 135 and 141 (all UPS).
Carabrusa aguarica auct. non (L.) Beauv.: Andersson 1969: 73.
Colpodium hedbergii auct. non (Meld.) Tzvel.: van Zinderen
Bakker & Werger 1974: 41, 47.
Colpodiurn sp. nov. in Killick
1978: 551.
Perennial with decumbent-ascending stems, branching
and rooting at the nodes. Leaf blades up to 1 1 cm long
and 5 mm wide, usually flat. Ligule 2 4 mm long. Panicle ovate, open, 3-10 cm long, its basal part often remaining in the uppermost sheath. Spikelets 2.3-3.5 mm
long, narrowly elliptic, light green with purplish tinge.
Glumes distinctly unequal and much shorter than lemma,
obtuse. Lemma and palea equilong, 2.3-3.5 mm, both
denticulately obtuse and glabrous.
Habitat. On moist ground along small brooks, 38005000 m.
Chromosome number. 2n = 20 (cf. Tab. I). Fig. 4B.
2. C. hedbergii (Meld.) Tzvel.
Tzvelev in Tzvelev & Bolchovskieh 1965: 1319; Clayton 1970:
51; Alexeev 1980: 9, Fig. 1: 14-19. - Keniochloa hedbergii
Melderis 1956: 542, Figs. I : upper row, and 2; 0. Hedberg
1957: 54. - Type: Kenya, Elgon, in the crater, Hedberg 908
(UPS holotype, EA, K, S isotypes).
Collecrions orher rhun fype: Qacha’s Nek Distr., Sani Pass, near
the lodge, in small rivulet, 3.11.1982, 0. Hedberg 82146 (PR,
ROML, UPS). Lesotho, Butha Buthe Distr. ,along the road from
Oxbow Lodge to the diamond mine, at a pond, 27.1.1982. 0.
Hedberg 82009 (EA, ETH, K, PRE, ROML, S, UPS). Natal,
Underberg, in seepage area on summit of Drakensberg, 9000’.
25.1.1966, D. J. B. Killick & J. Vahrmeyer 3730 (PRE). Basutoland, Buthe Buthe, Pone Valley, Mothae Mts., in pools 9700’,
8.1.1958, Liitjeharms (PRE). Lesotho, Drakensberg, Werger
I614 (K).
Tufted perennial with erect culms 7-40 cm high. Leaf
blades up to 11 cm long and 4 mm wide. Ligule 2-8 mm
long. Panicle ovate, 4-20 cm long, with spreading
branches, some of which are normally reflexed. In young
specimens the panicle may still appear contracted, as in
Aichhorn 7 (EA). Spikelets 2.4-3.4 mm long, narrowly
elliptic, light green or purplish. Glumes equal, longer
than lemma. Lemma 2.3-3 mm long, 3-nerved, with
606
Acknowledgements - Herbarium material was obtained on loan
from - or studied at EA, K, PRE, S and UPS. For the
arrangement of loans we are indebted to the Herbarium at
Uppsala university, and for care of living material to the Botanic
Garden of Uppsala University. Living material and additional
herbarium specimens were kindly sent by Mr S. Bie, Dr A. Jacot
Guillarmod and Dr D. J. B. Killjck. Dr Killick also organized an
excellent opportunity for us to see and collect material in the
-
N o d . J. Boi. 14 ( 6 ) (1994)
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Drakensberg. We are also indebted to Ms Inger Sorensen for
drawing Fig. 3 and to Ms Eva Persson for preparing the other
illustrations and assisting in various other ways. Assistance on
cytological preparation was given by Mrs W. Olsson. The Latin
diagnosis was checked by Prof. L. Holm. Valuable comments on
the manuscript were given by Dr D. J. B. Killick, Pretoria, Mrs
S. Phillips, Kew, and an anonymous reviewer.
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