Chromosome numbers of South African Umbelliferae (Apiaceae)

Abstract

Chromosome numbers are reported for 12 species from nine genera of South African Umbelliferae, of which seven species and one genus (Itasina) are recorded for the first time. A detailed list of all published chromosome counts for southern African species is also presented, together with a review of the literature. The new data obtained are briefly discussed in the context of the taxonomy and relationships of local Umbelliferae. The counts agree with previous reports except that Annesorhiza appears to have 2n=22, with or without one additional B-chromosome, and not 2n=24 as reported in the literature. The number for Itasina (2n=24) is of considerable interest and indicates that a detailed chromosome study of the South African genera Annesorhiza and Chamarea may yield valuable taxonomic information.

Full text

Short communication
Chromosome numbers of South African Umbelliferae (Apiaceae)
J.V. Shner
a
, T.V. Alexeeva
a
, M.G. Pimenov
a
, B.-E. Van Wyk
b,
⁎
a
Botanical Garden, Moscow State University, Moscow 119899, Russia
b
Department of Botany and Plant Biotechnology, University of Johannesburg, Johannesburg, South Africa
Received 31 December 2009; received in revised form 15 July 2010; accepted 19 October 2010
Abstract
Chromosome numbers are reported for 12 species from nine genera of South African Umbelliferae, of which seven species and one genus
(Itasina) are recorded for the first time. A detailed list of all published chromosome counts for southern African species is also presented, together
with a review of the literature. The new data obtained are briefly discussed in the context of the taxonomy and relationships of local Umbelliferae.
The counts agree with previous reports except that Annesorhiza appears to have 2n = 22, with or without one additional B-chromosome, and not
2n = 24 as reported in the literature. The number for Itasina (2n = 24) is of considerable interest and indicates that a detailed chromosome study of
the South African genera Annesorhiza and Chamarea may yield valuable taxonomic information.
© 2010 SAAB. Published by Elsevier B.V. All rights reserved.
Keywords: Anginon; Apiaceae; Apioideae; Cape Floristic Region; Chromosome counts; Itasina
1. Introduction
Chromosome numbers are part of a multidisciplinary
approach to the characterization of taxa. The South African
Umbelliferae include several anomalous and taxonomically
isolated genera that are of importance in unravelling the
relationships within the family as a whole (Burtt, 1991; Magee
et al., 2010a,b; Van Wyk and Tilney, 2004), yet chromosome
counts are available only for a limited number of species.
Among the approximately 205 species of southern African
Umbelliferae (Allison and Van Wyk, 1997; Burtt, 1991; Liu et
al., 2007; Magee et al., 2008a,b, 2009a,b; Schubert and Van
Wyk, 1997; Tilney and Van Wyk, 2002; Van Wyk and Tilney,
2003, 2004; Winter et al., 2008), chromosome counts based on
local material are available for 25 species only, belonging to 19
genera (Table 1). Most of them are South African endemics or
they are at least endemic or subendemic to the African
continent. The only exceptions are the more widely distributed
Apium graveolens L. and Torilis arvensis (Huds.) Link. This
means that chromosome numbers of only 23 South African
species (indigenous or naturalised) have been determined up to
now (Pimenov et al., 2002).
The most important sources of data include two publications
by L. Constance et al. (Constance and Chuang, 1982; Constance
et al., 1976). Some additional references on the identity of the
species that were counted can be found in Burtt (1991). The
only other counts reported are those of Riley and Hoff (1961)
for Steganotaenia araliacea Hochst., Cauwet (1971) and
Cauwet-Marc (1979) for Bupleurum mundtii Cham. & Schltdl.
and Winter and Van Wyk (1996) for some South African
species of Heteromorpha Cham. & Schltdl.
The aim of this paper is to (1) report chromosome numbers
for an additional 12 South African species from nine genera of
Umbelliferae, and (2) to review all available data on southern
African species and the associated literature.
2. Materials and methods
Plant material (flower buds and fruits) was collected during
two short excursions before and after the 4th International
Apiales Symposium in Pretoria, 3 to 19 January 2003 (Table 2).
Author citations for the scientific names given in Tables 1 and 2
are not repeated from here on. Fixations were made by M.G.
Pimenov and voucher specimens were collected and identified
A
vailable online at www.sciencedirect.com
South African Journal of Botany 77 (2011) 497 –502
www.elsevier.com/locate/sajb
⁎Corresponding author. Tel.: +27 115592412; fax: +27 115592411.
E-mail address: bevanwyk@uj.ac.za (B.-E. Van Wyk).
0254-6299/$ - see front matter © 2010 SAAB. Published by Elsevier B.V. All rights reserved.
doi:10.1016/j.sajb.2010.10.009

by M.G. Pimenov, B.-E. Van Wyk, E.V. Kljuykov and T.A.
Ostroumova; they are kept in MW.
Meiotic chromosomes were examined in pollen mother cells
(Fig. 1). Flower buds were fixed in the field in 3:1 96%
ethanol–glacial acetic acid mixture and then stained with 1%
acetocarmine for 3 to 4 days at about 5 °C. After staining, the
anthers were squashed and mounted. Somatic chromosomes
were examined in meristem cells of root tips from germinating
seeds (Fig. 2). After a pretreatment in 0.01% aqueous colchicine
solution for 3 h at room temperature, the root tips were fixed in
3:1 96% ethanol–glacial acetic acid mixture and then stained
with 1% acetocarmine at 100 °C for 3 to 5 min. Both the anthers
and the root tips were squashed on a glass microscope slide in
chloral hydrate (Hoyer's solution).
3. Results and discussion
Chromosome numbers were determined for 12 species from
nine genera (Table 2). These counts include a first record for the
genus Itasina Raf. and also first counts for seven other species
(Afrosciadium caffrum,A. magalismontanum,Anginon dif-
forme,A. paniculatum,A. swellendamense,Lichtensteinia
lacera and Notobubon capense). In addition, a first correct
count for Annesorhoza macrocarpa is presented.
The chromosome number of Itasina filifolia (n = 12) is rather
unusual in Apioideae. It is known for only 24 species of the
subfamily (representing 1.6% of species investigated) and was
found in the traditional tribes Scandiceae (two species),
Caucalideae (four species), Smyrnieae (one species), Apieae
(14 species), Peucedaneae (one species) and Tordylieae (two
species). Itasina traditionally belongs to the tribe Apieae but
molecular systematic evidence recently showed that it is related
to the genera Annesorhiza Cham. & Schltdl., Astydamia DC.,
Chamarea Eckl. & Zeyh., Ezosciadium B.L.Burtt and Molo-
pospermum W.J.D. Koch (Magee et al., 2010b) and that these
genera should be recognized as a new tribe, the Annesorhizeae
Magee et al. (Magee et al., 2010b). These genera (especially
Ezosciadium) would be worth sampling in further cytological
studies. Counts of n = 12 (or 2n = 24) occur sporadically in
species of unrelated genera, including Ammoselinum Torr. & A.
Gray, Apium L., Bunium L., Bupleurum L., Carum L., Cnidium
Cusson ex Jussieu, Pimpinella L., Spuriopimpinella (H.
Boisseu) Kitag. and Tilingia Regel (Pimenov et al., 2002).
The count of n = 12 in Umbelliferae deserv es special attention as
it is common in Araliaceae, the reputedly related family.
Moreover, x = 12 or x = 6 is regarded as the most plaus ible basic
and ancestral chromosome number in Apiales (Sharma and
Chatterji, 1964; Yi et al., 2004). In connection with this putative
close affinity, n = 12 in the woody African species S. araliacea
Table 1
Previously determined chromosome numbers for 19 genera and 25 species of South African Umbelliferae.
Genera and species Chromosome number References
Afroligusticum wilmsianum (H. Wolff.) P.J.D.Winter [as Peucedanum wilmsianum H.Wolff] n = 11 Constance and Chuang (1982)
Afrosciadium platycarpum (Sond.) P.J.D.Winter [as Peucedanum platycarpum E.Mey ex Sond.] n = 22 Constance and Chuang (1982)
n=11 Burtt (1991) [Eastern Cape specimen]
Agrocharis melanantha Hochst. n = 22 Constance and Chuang (1982);
Burtt (1991, p. 152; Sani Pass specimen)
Alepidea natalensis J.M.Wood & Evans n = 8 Constance and Chuang (1982)
Anginon rugosum (Thunb.) Raf. [= A. uitenhagense (Eckl. & Zeyh.) B.L.Burtt], as n = 11 Burtt (1991)
Annezorhiza macrocarpa Eckl. & Zeyh. n= 12 Burtt (1991)
Apium graveolens L. n = 11 Constance et al. (1976);
Constance and Chuang (1982)
Arctopus echinatus L. n = 9 Constance et al. (1976)
Bupleurum mundtii Cham. & Schltdl. 2n = 16 Cauwet (1971); Cauwet-Marc (1979)
Conium chaerophylloides (Thunb.) Sond. n = 11 Constance and Chuang (1982)
C. fontaneum Hilliard & B.L.Burtt n = 11 Constance and Chuang (1982)
Cynorhiza typica Eckl. & Zeyh. [as Peucedanum olifantianum (Koso-Pol.) M.Hiroe] n = 11 Constance and Chuang (1982)
Dasispermum capense (Lam.) Magee & B.-E.Van Wyk [as Stoibrax capense (Lam.) B.L.Burtt] n = 9 Constance et al. (1976)
D. humile (Meisn.) Magee & B.-E.Van Wyk [as Sonderina humilis (Meisn.) H.Wolff] n = 8 Burtt (1991)
D. suffruticosum (Berg.) B.L.Burtt n = 9 Constance et al. (1976);
Constance and Chuang (1982)
Glia prolifera (Burm.f.) B.L.Burtt n = 11 Constance et al. (1976)
Hermas villosa (L.) Thunb. n = 7 Constance et al. (1976)
Heteromorpha arborescens (Spreng.) Cham. & Schltdl. n = 11 Constance and Chuang (1982)
H. arborescens var. frutescens P.J.D.Winter 2n = 22 Winter and Van Wyk (1996)
H. involucrata Conrath 2n = 22 Winter and Van Wyk (1996)
H. pubescens Burtt Davy 2n = 22 Winter and Van Wyk (1996)
Lichtensteinia interrupta (Thunb.) Sond. [as L. kolbeana Bolus] n = 11 Constance and Chuang (1982)
Notobubon galbanum (L.) Magee [as Peucedanum galbanum (L.) Benth. & Hook.f.] n= 11 Constance and Chuang (1982)
N. laevigatum (Aiton) Magee [as Peucedanum capense (Thunb.) Sond.] n = 11
Steganotaenia.araliacea Hochst. 2n = 22 Riley and Hoff (1961);
Constance and Chuang (1982)
Torilis arvensis (Huds.) Link n = 6 Constance et al. (1976)
498 J.V. Shner et al. / South African Journal of Botany 77 (2011) 497–502

(Constance and Chuang, 1982) could be of considerable
interest. Steganotaenia and Polemanniopsis B.L. Burtt (both
woody shrubs or trees) have recently been recognized as a new
tribe, the Steganotaeniae C.I. Calviño & S.R. Downie (Calviño
and Downie, 2007). The count of n = 12 in the herbaceous
Itasina however, seems to be of independent origin, as there is
no other evidence of a direct connection with Steganotaenia
Hochst. or with Araliaceae. It is interesting to note that Itasina
and Steganotaenia both form part of the early diverging and
predominantly African lineages of the subfamily Apioideae
(Magee et al., 2010b).
Our determination does not confirm the reported chromosome
number of n= 12 for A. macrocarpa,publishedbyBurtt (1991)
with a reference to Constance and Chuang, cited as “n =12 (C & C
ined. based on Batten 590 from E. Cape, E. London). Our data
showed that the species has n= 11, but sometimes its karyotype
contains one additional B-chromosome. It is therefore likely that
earlier researchers incorrectly counted a B-chromosome as one of
the basic set. Since Annesorhiza,Chamarea and Itasina are now
considered to be closely related (Magee et al., 2010b; Vessio,
2001), a wider survey of these genera seems desirable and will
yield valuable information.
There is no straightforward evidence for the chromosome
number of any species of the woody genus Anginon Raf. in the
karyological literature. However, Burtt (1991) showed n = 11
for A. uitenhagense on the basis of a re-identification of the
Bayliss 8877 collection studied by Constance and Chuang
(1982) and reported as “Peucedanum zeyheri”.Allison and Van
Wyk (1997) reduced A. uitenhagense to the synonymy of A.
rugosum but did not cite Bayliss 8877 among the material
examined. We determined the same count of n = 11 for three
correctly identified species of Anginon (see Table 2). This
means that all counts available for the genera of the tribe
Heteromorpheae M.F. Watson & S.R. Downie (Anginon,Glia
Sond. and Heteromorpha) are n = 11 or 2n = 22.
The chromosome number of Cyclospermum leptophyllum
[=Apium leptophyllum (Pers.) F.Muell.], a widely distributed
weed of warm and adjacent temperate regions of the world, has
been determined at least 12 times. The determinations for plants
from South and North America, Oceania, Europe and China
invariably showed a count of n = 7 (or 2n = 14). This was
confirmed by the present determination, the first one for African
material.
For Dasispermum suffruticosum and Hermas villosa, our
counts confirm previous ones (n = 9 and n = 7 respec tively). In
the first case, the chromosome number distinguishes Dasisper-
mum Neck. ex Raf. from the genera Conium L., Athamanta L.,
Cicuta L., Cnidium,Selinum L., Sium L. and others, to which
the species has been referred to at various times during its long
taxonomic history. Recently, Magee et al. (2009b) have shown,
on the basis of morphological and molecular evidence, that
Dasispermum is congeneric with another South African genus
Sonderina H.Wolff, and that Stoibrax capensis is also part of
this genus. The known counts of n = 8 for Dasispermum humile
(=Sonderina humilis) and n = 9 for D. capense (=S. capensis)
(Table 1) indicate that the more broadly circumscribed
Dasispermum has n = 8 or n = 9 and that there is a reduction in
the chromosome number in this group. More counts will be
valuable and interesting, as the genus now comprises seven
species (Magee et al., 2009b, 2010a).
Hermas L. is an anomalous genus with affinities in the new
subfamily Azorelloideae, where it was moved from its
traditional position in the subfamily Hydrocotyloideae. In
general, the subfamily Hydrocotyloideae in its traditional
circumscription is not monophyletic (Nicolas and Plunkett,
2009). Within the subfamily Azorelloideae, the count of n = 7
found in Hermas, is unique. Traditionally Hermas had been
included in the tribe of Mulineae, but modern molecular
analyses (Nicolas and Plunkett, 2009; Plunkett, 2001) showed
an isolated position for the tribe on the border between
Umbelliferae and Araliaceae. Morphological analyses by Liu
(2004) and Liu et al. (2009), as well as molecular systematic
evidence (Plunkett et al., 2004) indicated that the group should
be recognized as a separate subfamily, the Azorelloideae.
Table 2
Newly determined chromosome numbers for nine genera and 12 species of South African Umbelliferae (specimens are housed in MW).
Species Origin and voucher specimen n 2n
Afrosciadium caffrum (Meisn.) P.J.D.Winter
[=Peucedanum caffrum (Meisn.) Philips]
Limpopo Province, slopes of Wolkberg, Haenertsburg, 23°56′S, 29°57′E. 05.01.2003,
Pimenov et al. 17
22
A. magalismontanum (Sond.) P.J.D.Winter
[=Peucedanum magalismontanum Sond.]
Gauteng Province, 20 km E of Pretoria, near Bronkhorstspruit, 25°46′S, 28°32′E. 03.01.2003,
Pimenov et al. 1
22
Anginon difforme (L.) B.L.Burtt Western Cape Province, Kirstenbosch, 34°03′S, 18°23′E. 14.01.2003, Pimenov et al. 73 11
A. paniculatum (Thunb.) B.L.Burtt Western Cape Province, near Citrusdal, 32°25′S, 18°57′E. 15.01.2003, Pimenov et al. 82 11
A. swellendamense (Eckl. & Zeyh.) B.L.Burtt Western Cape Province, Worcester, 33°12′S, 19°23′E. 15.01.2003, Pimenov et al. 94 11
Annesorhiza macrocarpa Eckl. & Zeyh. Western Cape Province, Noordhoek, 34°05′S, 18°21′E. 13.01.2003, Pimenov et al. 59 11, 11 + 1B
Cyclospermum leptophyllum (Pers.)
Sprague ex Britton & P.Wilson
Limpopo Province, slopes of Wolkberg, Haenertsburg, 23°56′S, 29°57′E. 05.01.2003,
Pimenov et al. 13
14
Dasispermum suffruticosum (Berg.) B.L.Burtt Western Cape Province, Betty's Bay, Stony Point, 34°21′S, 18°55′E. 12.01.2003,
Pimenov et al. 47
918
Hermas villosa (L.) Thunb. Western Cape Province, Table Mountain, 33°57′S, 18°24′E.13.01.2003, Pimenov et al. 67 7
Itasina filifolia (Thunb.) Raf. Western Cape Province, Cape Peninsula, 34°21′S, 18°22′E. 3.01.2003, Pimenov et al. 61 12
Lichtensteinia lacera Cham. & Schltdl. Western Cape Province, near Silver Mine, 34°05′S, 18°25′E. 13.01.2003, Pimenov et al. 49 11
Notobubon capense (Eckl. & Zeyh.) A.R.Magee
[=Peucedanum multiradiatum Drude]
Western Cape Province, Betty's Bay, 34°21′S, 18°55′E. 12.01.2003, Pimenov et al. 41 22
499J.V. Shner et al. / South African Journal of Botany 77 (2011) 497–502

The number of n= 11 for L. lacera is the second
determination for the genus. It corresponds to Constance and
Chuang's (1982) count for the eastern Cape form of L.
interrupta, previously known as L. kolbeana. This number is
widely distributed in Apioideae, but very rare in Saniculoideae
(only the critical South American tetraploid genus Oligocladus
Chodat & Wilczek has n = 11). This confirmation is important
because it strongly supports the idea that Lichtensteinia is more
closely related to the subfamily Apioideae than to the
Saniculoideae, despite sharing several morphological characters
with the latter. Molecular evidence indicates that Lichtensteinia
is sister to the cytologically unknown genus Marlothiella H.
Wolff and that both are sister to all other Apioideae (Calviño et
al., 2006, 2008; Magee et al., 2010b).
Finally, according to our data, three South African species
that were previously included in the broad concept of
Peucedanum L. have 2n = 22. The two herbaceous species
from the northern part of South Africa are now included in the
newly described genus Afrosciadium P.J.D.Winter, while the
shrubby species from the Cape are now placed in the genus
Notobubon B-E.Van Wyk (Magee et al., 2009a; Winter et al.,
2008). These data are unfortunately not very informative to
elucidate relationship among the species, because this count is
the most frequent in Peucedanum sensu lato as well as in related
and unrelated genera of Apioideae. The need to subdivide
African Peucedanum species was also indicated by the studies
of Ostroumova and Pimenov (1997), where the considerable
diversity in the fruit structure was highlighted for the first time.
We were unable to confirm the occurrence of tetraploidy
(n = 22), report ed in Afrosciadium platycarpum from the eastern
Cape and also Agrocharis melanantha from Sani Pass (Table 1).
4. Conclusions
Chromosome numbers of the Umbelliferae of South Africa
are insufficiently known and do not yet allow for global
comparisons of the family as a whole. We determined
chromosome numbers for 12 Umbelliferae species of the
South African flora. The genus Itasina and seven other species
were counted for the first time while the previously reported
chromosome numbers of the remaining species were confirmed
or sometimes corrected. This study and the review of available
data showed that chromosome information can be of value in
critical taxonomic revisions of the interesting and anomalous
early diverging lineages of southern African Umbelliferae.
Acknowledgements
Our cordial thanks are addressed to all organizers and
participants of the 4th International Apiales Symposium and its
excursions. The Russian authors acknowledge the RFFI; MGP
Fig. 1. Microphotographs of meiotic chromosomes of some South African Umbelliferae. (a) Anginon difforme (n= 11); (b) A. paniculatum (n = 11); (c) A.
swellendamense (n= 11); (d) Annesorhiza macrocarpa (n = 11); (e) A. macrocarpa with B-chromosomes (n = 11 + 1B) [a second set of chromosomes that did not
separate is visible to the right, with their corresponding B-chromosome]; (f) Dasispermum suffruticosum (n = 9); (g) Hermas villosa (n = 7); (h) Itasina filifolia (n = 12);
and (i) Lichtensteinia lacera (n = 11). Voucher specimens are listed in Table 2. Scale bar = 10 μm.
500 J.V. Shner et al. / South African Journal of Botany 77 (2011) 497–502

and BEVW additionally thank the University of Johannesburg
and the National Research Foundation for financial support.
References
Allison, I., Van Wyk, B.-E., 1997. A revision of the genus Anginon (Apiaceae).
Nordic Journal of Botany 17, 561–577.
Burtt, B.L., 1991. Umbelliferae of southern Africa: an introduction and
annotated checklist. Edinburgh Journal of Botany 48, 133–282.
Calviño, C.I., Downie, S.R., 2007. Circumscription and phylogeny of Apiaceae
subfamily Saniculoideae based on chloroplast DNA sequences. Molecular
Phylogenetics and Evolution 44, 175–191.
Calviño, C.I., Tilney, P.M., Van Wyk, B.-E., Downie, S.R., 2006. A molecular
phylogenetic study of southern African Apiaceae. American Journal of
Botany 95, 196–214.
Calviño, C.I., Martίnez, S.G., Downie,S.R., 2008. Morphology and biogeography
of Apiaceae subfamily Saniculoideae as inferred by phylogenetic analysis of
molecular data. American Journal of Botany 93, 1832–1833.
Cauwet, A.-M., 1971. Contribution à l'étude caryosystématique du genre
Bupleurum L. III. Bulletin de la Societé botanique de France 114, 55–68.
Cauwet-Marc, A.-M., 1979. Connaissances caryologiques actuelles sur le genre
Bupleurum L. (Umbelliferae): nombres cromosomiques ar nombres de base.
Bulletin de Musée National d'Histoire Naturelle: 4e série Botanique, vol. 1,
pp. 191–211.
Constance, L., Chuang, T.-I., 1982. Chromosome numbers of Umbelliferae
(Apiaceae) from Africa south of the Sahara. Botanical Journal of the Linnean
Society 85, 195–208.
Constance, L., Chuang, T.-I., Bell, S.R., 1976. Chromosome numbers in
Umbelliferae. IV. American Journal of Botany 63, 608–625.
Liu, M., 2004. A taxonomic evaluation of fruit structure in the family Apiaceae.
PhD Thesis, University of Johannesburg.
Liu, M., Van Wyk, B.-E., Tilney, P.M., 2007. A taxonomic revision of the genus
Choritaenia (Apiaceae). South African Journal of Botany 73, 184–189.
Liu, M., Van Wyk, B.-E., Tilney, P.M., Plunkett, G.M., Lowry II, P.P., 2009.
Evidence from fruit structure supports in general the circumscription of
Apiaceae subfamily Azorelloideae. Plant Systematics and Evolution 280,
1–13.
Magee, A.R., Van Wyk, B.-E., Tilney, P.M., Van der Bank, M., 2008a. A
taxonomic revision of the South African endemic genus Arctopus
(Apiaceae, Saniculoideae). Annals of the Missouri Botanical Garden 95,
471–486.
Magee, A.R., Tilney, P.M., Van Wyk, B.-E., Downie, S.R., 2008b.
Ezosciadium: yet another early diverging South African apioid. Plant
Systematics and Evolution 276, 167–175.
Magee, A.R., Tilney, P.M., Van Wyk, B.-E., 2009a. A revision of the woody
Cape genus Notobubon (Apiaceae). Systematic Botany 34, 220–242.
Magee, A.R., Tilney, P.M., Van Wyk, B.-E., Downie, S.R., 2009b. Generic
delimitations and relationships of the Cape genera Capnophyllum,
Dasispermum and Sonderina, the North African Krubera and Stoibrax,
and a new monotypic genus of the subfamily Apioideae (Apiaceae).
Systematic Botany 34, 580–594.
Magee, A.R., Van Wyk, B.-E., Tilney, P.M., Downie, S.R., 2010a. A taxonomic
revision of the genus Dasispermum (Apiaceae, Apioideae). South African
Journal of Botany 76, 308–323.
Magee, A.R., Calviño, C.I., Liu, M., Downie, S.R., Tilney, P.M., VanWyk, B.-E.,
2010b. New tribal delimitations for the early diverging lineages of Apiaceae
subfamily Apioideae. Taxon 59, 567–580.
Nicolas, A.N., Plunkett, G.M., 2009. The demise of subfamily Hydrocotyloi-
deae (Apiaceae) and the re-alignment of its genera across the entire order
Apiales. Molecular Phylogenetics and Evolution 53, 134–151.
Fig. 2. Microphotographs of somatic chromosomes of some South African Umbelliferae. (a) Cyclospermum leptophyllum (2n = 14); (b) Dasispermum suffruticosum
(2n = 18); (c) Afrosciadium caffrum (2n = 22); (d) A. magalismontanum (2n = 22); and (e) Notobubon capense (2n = 22). Voucher specimens are listed in Table 2. Scale
bar = 10 μm.
501J.V. Shner et al. / South African Journal of Botany 77 (2011) 497–502

Ostroumova, T.V., Pimenov, M.G., 1997. Carpological diversity of African
Peucedanum s.l. I. The species of southern Africa. Feddes Repertorium 108,
299–318.
Pimenov, M.G., Vassil'eva, M.G., Daushkevich, J.V., Leonov, M.V., 2002.
Karyotaxonomical analysis in the Umbelliferae. Science Publishers, Enfield
(USA), p. 468.
Plunkett, G.M., 2001. Relationship of the order Apiales to subclass Asteridae:
re-evaluation of morphological characters based on insights from molecular
data. Edinburgh Journal of Botany 58, 183–200.
Plunkett, G.M., Chandler, G.T., Lowry, P.P., Pinney, S., Sprenkle, T., 2004.
Recent advances in understanding Apiales with a revised classification.
South African Journal of Botany 70, 371–381.
Riley, H.P., Hoff, V.J., 1961. Chromosome studies in some South African
Dicotyledons. Canadian Journal of Genetics and Cytology 3, 260–271.
Schubert, M.T.R., Van Wyk, B.-E., 1997. A revision of Centella series
Capenses (Apiaceae). Nordic Journal of Botany 17, 301–314.
Sharma, A.K., Chatterji, A.K., 1964. Cytological study as an aid in the
interpretation of the systematic status of different genera of Araliaceae.
Cytologia 29, 1–2.
Tilney, P.M., Van Wyk, B.-E., 2002. A revision of the genus Annesorhiza
(Apiaceae). Nordic Journal of Botany 21, 615–649.
Van Wyk, B.-E., Tilney, P.M., 2003. Apiaceae.In: Germishuizen, G., Meyer, N.L.
(Eds.), Plants of southern Africa: an annotated checklist. : Strelitzia, vol. 14.
National Botanical Institute, Pretoria, pp. 123–132.
Van Wyk, B.-E., Tilney, P.M., 2004. Diversity of Apiaceae in Africa. South
African Journal of Botany 70, 433–445.
Vessio, N., 2001. The generic affinities of deciduous species of the genera
Annesorhiza Cham. & Schlechtd., Chamarea Eckl. & Zeyh. and
Peucedanum L. (Apiaceae). MSc dissertation, University of Johannesburg.
Winter, P.J.D., Van Wyk, B.-E., 1996. A revision of the genus Heteromorpha
(Apiaceae). Kew Bulletin 51, 225–265.
Winter, P.J.D., Magee, A.R., Phephu, N., Tilney, P.M., Downie, S.R., Van Wyk,
B.-E., 2008. A new generic classification for African peucedanoid species
(Apiaceae). Taxon 57, 347–364.
Yi, T., Lowry II, P.P., Plunkett, G.M., Wen, J., 2004. Chromosomal evolution in
Araliaceae and close relatives. Taxon 53, 987–1005.
Edited by TJ Edwards
502 J.V. Shner et al. / South African Journal of Botany 77 (2011) 497–502