PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Volume 52, No. 1 2, pp. 143-1 58, figs. 1-3, 1 table. December 7, 2000 
Chromosome Numbers of South African Acanthaceae 
by 
Thomas F. Daniel 
Department of Botany, California Academy of Sciences 
Golden Gate Park, San Francisco. California 94 11 8 
and 
Kevin Balkwill and Mandy-Jane Balkwil! 
C. E. Moss Herbarium 
Department of Animal. Plant, and Environmental Sciences 
University of the Witwatersrand 
Private Bag X3. WITS 2050. South Africa 
Original meiotic chromosome counts are presented for 19 species in 14 genera of 
Acanthaceae from South Africa. These counts represent the initial reports of chromosome 
numbers from southern African Acanthaceae. Chromosome numbers of 13 species repre- 
senting 1 1 genera are reported for the first time. The counts in Aulojusticia, Duvernoia, and 
Metarungia are the first for these genera. Counts for five species confirm numbers previ- 
ously reported for them based on plants from other regions. A new chromosome number is 
reported in Justicia (« = 26 in /. petiolaris). Systematic implications of these chromosome 
counts are addressed. 
The Acanthaceae are a large (ca. 4000 species in some 230 genera) pantropical family with major 
concentrations of species in southeastern mainland Asia, insular Malesia, the Indian subcontinent, 
Madagascar, tropical Africa, Brazil, Andean South America, and Mexico-Central America. Knowl- 
edge of chromosome numbers among Acanthaceae has proven useful in resolving generic positions of 
problematic species, reassessing phylogenetic relationships among subfamilial taxa, and understand- 
ing morphological variation (e.g., Ensermu Kelbessa 1990; Daniel and Chuang 1993; McDade et al. 
in press). A major problem with using chromosome number data in the study of systematic and evolu- 
tionary relationships among Acanthaceae is that numbers remain unknown for the vast majority of 
both genera and species. Daniel (2000) noted that chromosome numbers had been reported for only 
29% of the genera and less than 1 2% of the species of Acanthaceae. Similarly, chromosome numbers 
remain undetermined for five of the 12 tribes recognized by Bremekamp (1965) in the family: 
Haselhoffieae, Louteridieae, Rhombochlamydeae, Stenandriopsideae, and Whitfieldieae. Another 
problem is the lack of, or poor sampling of, species from regions rich in Acanthaceae but 
underrepresented by chromosome counts of them. One such region, as identified by Daniel (2000), is 
southern Africa. This region, comprising Namibia, Botswana, South Africa, Lesotho, and Swaziland, 
has an acanthaceous flora of some 341 species (Welman 1993). Chromosome numbers have been re- 
ported previously for 28 of these (see Appendix), but none of these reports was based on collections 
from southern Africa. This study of chromosome numbers in selected Acanthaceae from South Africa 
is our first attempt to provide this systematically useful information for taxa from this region. 
FEB 05 200 J I 
Woods i[a;c, ,w,« n-.«.,3 ) 
144 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Volume52, No. 12 
Methods 
In February and March of 1999, floral buds and herbarium vouchers of South African 
Acanthaceae were collected from their native habitats and from plants cultivated in gardens. Buds 
were fixed in absolute ethanol:glacial acetic acid (3:1) for 24 hours and subsequently washed and 
stored in ethanol (70%) until analyzed. Anthers were macerated in ferric acetocarmine ( 1 %) on a mi- 
croscope slide, squashed under a coverslip by hand, and studied under oil immersion on a phase con- 
trast microscope at a magnification of lOOOx. Counts were made from microsporocytes in various 
stages of meiosis. Preparations from which counts were obtained were recorded with camera lucida 
drawings. Voucher specimens are deposited at CAS and J. The camera lucida drawings are attached to 
the vouchers at CAS. 
In the following discussion, all previously published chromosome counts are listed as n (gametic, 
haploid, meiotic) numbers irrespective of whether they were originally reported as gametophytic or 
sporophytic numbers. Voucher specimens, if they exist, documenting previous counts by other work- 
ers have not been examined. 
Results 
Chromosome numbers determined for 19 species representing 14 genera of Acanthaceae from 
South Africa are summarized in Table 1 and illustrated in Figures 1-3. Chromosome numbers of 1 3 of 
these taxa are reported for the first time. The counts for Aulojusticia, Duvernoia, and Metarungia rep- 
resent the first reports for these genera. Counts for five species (Asystasia gangetica.. Hypoestes 
aristata, H. forskaolii, Justicia betonica, and Rhinacanthus gracilis) agree with some or all numbers 
previously reported for them from different sources. The chromosome number reported here for 
Justicia odora is the first non-approximate number reported for the species. The count of ;7 = 26 for J. 
petiolaris is the first report of this number in the genus. 
Discussion 
Asystasia Blume. Fifty or more species are recognized in this genus, which is native to the Old 
World. Eight species are known from southern Africa (Welman 1993). Daniel (2000) summarized 
previous reports of chromosome numbers in both A. gangetica and in the genus. Although a diversity 
of meiotic numbers, including 1 4, 22, 24 and 25, has been reported iox A. gangetica, counts of /? = 13 
and n = 26 are much more common. A meiotic number of 1 3 is also the most frequently reported chro- 
mosome number among other species in the genus. 
Asystasia gangetica has a broad distribution, occurring indigenously in southern and tropical Af- 
rica and in the Indian subcontinent. It has become naturalized in several other tropical regions (e.g., 
Hawaii, Java). Previous counts (including those reported for A. coromandeliana Nees; see summary 
of counts in Daniel 2000) have come from plants in cultivation (without provenance data; e.g.. Grant 
1 955), from the Indian subcontinent (e.g., Valsala Devi and Mathew 1 982), from tropical west Africa 
(e.g., Gadella 1 977), and from plants naturalized outside of the native range of the species (e.g., Daniel 
2000). Our count ofn = 1 3 for a southern African representative of the species reveals a continuity of 
chromosome number with plants from other portions of the species' range. Plants from which our 
count was determined would appear to be diploid within the species. As indicated by Daniel (2000), a 
basic number of x = 1 3 appears likely for the genus. 
Aulojusticia Lindau. Our count of a? = 40 for A. linifolia is the first chromosome count for this ge- 
nus. Aulojusticia has been variably treated by students of Acanthaceae: Dyer ( 1 975) recognized it as a 
unispecific genus endemic to northeastern South Africa; Graham ( 1 988) included it within her broad 
DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 
145 
Table 1. Meiotic chromosome counts of South African Acanthaceae. Note: Counts for Duvernoia 
aconitijJora, Justicia petiolahs, Metanmgia longistrobus, Pseitderanthemum hildehrandtii, and Ruspolia 
hypocrateriformis were obtained from plants in cultivation; * indicates first counts for taxa. 
Taxon 
Chromosome 
number (n) 
Locality 
(province) 
Collection 
number 
(Daniel et al.) 
Asystasia gangetica (L.) T. Anderson 
Aulojusticia lirtifolia Lindau 
Barleria senensis Klotzsch 
Crahbea angustifolia Nees 
Dicliptera heterostegia Presl. ex Nees 
Dicliptera magalieshergensis K. Balkwill 
Duvernoia aconitiflora A. Meeuse 
Hypoestes aristata (Vahl) Sol. ex R. & S. 
Hypoestes forskaoUi (Vahl) R.Br. 
Isoglossa hypoestiflora Lindau 
Isoglossa ovata (Nees) Lindau 
Justicia betonica L. 
Justicia betonica L. 
Justicia odora (Forssk.) Lam. 
Justicia petiolaris (Nees) T. Anderson 
Metaningia longistrobus (C. B. CI.) Baden 
Pseuderanthemum hildebrandtii Lindau 
Rhinacanthus gracilis Klotzsch 
Rhinacanthus gracilis Klotzsch 
Ruspolia hypocrateriformis (Vahl) 
Milne-Redh. 
Ruttya ovata Harv. 
circumscription oi Justicia; and Immelman (1995a) treated it as Siphonoglossa linifolia (Lindau) 
C. B. Clarke. Siphonoglossa, which is based on an American type, is conspeciflc W\i\\ Justicia (Gra- 
ham 1 988, Daniel 1 995). Molecular studies that seek to improve our understanding of Justicia and al- 
lied genera in Africa are currently underway. y4«/o/M.s//c/fl undoubtedly falls within the morphological 
circumscription of Justicia as delimited by Graham. If treated in that genus, the chromosome number 
here reported fox A. linifolia would be the highest number so far known m Justicia; the highest number 
previously reported is n = 34 (Daniel 2000). Also, if treated '
Justicia, which appears to have a basic 
number of x = 7 (see below), then this species likely would have been derived from a hexaploid ances- 
tor. 
Barleria L. Barleria is a pantropical genus of perennial herbs and shrubs comprising some 300 
species. The majority of species are African and 69 occur in southern Africa (Balkwill and Balkwill 
1997). Daniel and Chuang (1989, 1998) summarized previously reported chromosome numbers in 
Barleria. They noted the prevalence ofn = 20 among species of the genus and the likelihood of x = 20 
as a basic number in the genus. 
Our count of «= 16 for Barleria senensis is the first count for this species, which is native to 
southern and tropical Africa. Previously, this number has been reported in the genus once for B. 
aristata L. (Datta and Maiti 1 970, as "B. cristata var. dichotoma"'). There are many counts for this spe- 
cies and most of them are « = 20 (Daniel and Chuang 1 989). Because these two species are treated in 
146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
VoIume52, No. 12 
different subgenera of Barleria (Balkwill and Balkwill 1997), a common chromosome number be- 
tween them is doubtfully due to common ancestry. 
Barleria was treated in tribe Barlerieae by Lindau (1895) and in tribe Ruellieae, subtribe 
Barleriinae by Bremekamp ( 1 965) and Balkwill and Getliffe Norris ( 1 988). Based on DNA sequence 
data (Hedren et al. 1995; Scotland et al. 1995; McDade and Moody 1999), Crabbea and Lepidagathis 
would appear to be closely related to Barleria. A chromosome number of « = 2 1 has been reported in 
species from each of these three genera (Chuang et al. 1 963; Grant 1 964; De 1 966; Daniel in press, see 
below) suggesting a possible symplesiomorphic number among them. Additional counts of species in 
Barleria are needed in order to ascertain whether knowledge of chromosome numbers will help to re- 
solve infrageneric relationships. 
Crabbea Harv. Two previous chromosome counts have been reported for this Afi-ican genus of 
about 1 2 species, seven of which occur in southern Africa ( Welman 1 993). Our count of /? = 2 1 for the 
South African endemic, C. angustifolia, is the first report of a chromosome number in the species. It 
agrees with a previous count for C velutina S. Moore (Grant 1964, as C reticulata C. B. Clarke). 
Renard et al. (1983) reported n = ca. 14 for C. velutina. 
Based on the few counts for species of Crabbea, a basic number of x = 2 1 is tentatively proposed 
for the genus. Lindau ( 1 895 ) included Crabbea in his tribe Barlerieae and the genus would be included 
in Bremekamp's (1965) tribe Ruellieae, subtribe Barleriinae. As noted above under Barleria, this 
number is known in the related genera Barleria and Lepidagathis. Balkwill and Getliffe Norris ( 1 988) 
advocated the placement of Crabbea in the Ruellieae, Ruelliinae. Among genera of Ruelliinae for 
which chromosome numbers are known, n = 2\ has been reported in several species of Eranthemum 
L. (e.g., Kaur 1970; Govindarajan and Subramanian 1985; Mangenot and Mangenot 1958, 1962). 
Dicliptera Juss. Chromosome numbers have been reported for 22 of the more than 1 00 species of 
perennial herbs and shrubs in this pantropical genus. Fifteen of these species are from the New World 
and seven are native to the Old World. Counts for all New World species are n = 40 (or n = ca. 40; 
Daniel 2000, Daniel and Chuang 1993, Piovano and Bemardello 1991) whereas previous counts of 
species from the Old World comprise a? = 1 0, 1 3, 1 5, 24, and 26 (with /? = 1 3 being most frequently re- 
ported; Daniel and Chuang 1 993). The species of Dicliptera from the Old World that were studied are 
all indigenous to Asia and comprise: D. bupleuroides Nees, D. cuneata Nees, D. elegans W. W. Smith 
[Kaur (1970) identified this taxon as '"Dicliptera elegans Dalz.," and neither the source of the plant 
nor a voucher was cited; Dalziel did not publish a species with this name and whether the plant studied 
by Kaur is actually the Chinese species described by Smith or an Indian taxon is not known], D. 
leonotis Dalziel ex C. B. Clarke, D. parvibracteata Nees, D. roxhurghiana Nees, and D. verticillata 
(Forsk.) C. Chr. Although two of these species (i.e., D. leonotis andD. verticillata) also occur in tropi- 
cal Africa, chromosome numbers for all of the Old World species of Dicliptera were apparently deter- 
mined from Asian populations. Our counts would therefore appear to be the first from African plants, 
and the first for species of Dicliptera native to southern Africa. Twelve species of Dicliptera are 
known to occur in southern Africa (Balkwill et al. 1996). The count of /7=13 for D. 
magaliesbergensis agrees with the majority of previous counts for Old World species. The count of 
n= 1 5 for D. heterostegia agrees with a count by Kaur ( 1 970) for the widespread Paleofropical spe- 
cies D. verticillata. 
Daniel et al. ( 1 990) noted that the difference in ploidal level between species of Dicliptera in the 
Old and New Worlds suggests a major geographical division in the genus. Recent phylogenetic stud- 
ies of the Justicieae that included Paleotropical and Neofropical species of Dicliptera indicate that the 
genus is monophyletic and is related to Hypoestes and Peristrophe in subfribe Diclipterinae (McDade 
et al. in press). 
DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 1 47 
e 
Figure 1 . Camera-Iucida drawings of meiotic chromosome preparations, a. Justicia petiolaris (Daniel et al 9387) n = 26 
(telophase I), b. Ruttya ovata (Daniel et al 9338), n = 2\ (metaphase I), c. Dicliptera heterostegia (Daniel et al 9329) n = 1 5 
(diakmesis I), d. Duvemoia aconitiflora (Daniel et al 9361), « = 17 (metaphase I), e. Isoglossa ovata (Daniel et al 9336) 
n-n (telophase I), f. Pseuderanthemum hildebrandtii (Daniel et al. 9394), « = 21 (diakenesis I). 
Piovano and Bemardello (1991) noted that, based on chromosome numbers so far reported in 
Dicliptera, j: = 10 is likely the basic number of the genus. The occurrence of /; = 15 in species of all 
genera of Diclipterinae, however, suggests that ;c = 1 5 is symplesiomorphic for both the subtribe and 
Dicliptera (Daniel and Chuang 1 993, McDade et al. in press). 
148 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Volume 52, No. 12 
Duvernoia E. Mey. ex Nees. This genus of two species that are endemic to eastern South Africa 
and Mozambique is sometimes treated within Justicia (Graham 1 988; Brummitt 1 992). Bremekamp 
( 1 939) argued for the recognition oi Duvernoia, primarily on the basis of its pollen, and treated at least 
one species from the New World in it. Our count of /? = 1 7 for D. aconitijJora is the first report of a 
chromosome number for both the genus and the species. Manning and Getliffe Norris ( 1 995) treated 
the genus as distinct from others in the tribe Justicieae, subtribe Justiciinae on the basis of its 
pedunculate inflorescences with small bracts and campanulate calyces. It remains to be determined 
whether these characteristics are sufficient to delimit the two species sometimes treated in Duvernoia 
from other Justiciinae on a global basis. A meiotic complement of 1 7 is known, but infrequently re- 
ported, in Justicia; it has been noted in four species, J. adhatoda L. (e.g., Daniel and Chuang 1 998), J. 
hetonica L. (e.g., Daniel and Chuang 1998 and see below), J. carnea Lindl. (Daniel and Chuang 
1 989), and J. trinervia Vahl (e.g., Krishnappa and Ranganath 1 982), representing three sections of the 
genus (Graham 1 988). It is perhaps revealing that J. adhatoda is the type of one of these, section 
Vasiaca Lindau, in which Graham (1 988) placed the other species o^ Duvernoia, D. adhatodoides E. 
Mey ex Nees. 
Hypoestes Sol. ex R.Br. Hypoestes consists of about 70 species of herbs and shrubs from Africa, 
eastern Asia, Malesia, and Australia. Three species occur in southern Africa (Balkwill and Getliffe 
Norris 1 985). Our count of /? = 1 5 for H.forskaolii, a species occurring in Africa and the Arabian pen- 
insula, agrees with a previously reported chromosome number for this species (Podlech 1986) and 
with most numbers reported for other species in the genus (Daniel and Chuang 1998). Our count of 
« = 30 for H. aristata, a species native to tropical and southern Africa, agrees with a previous count for 
this species based on cultivated plants (Daniel and Chuang 1998). This is the only known species of 
Hypoestes with a chromosome number other than n= \5. If a basic number of x = 1 5 is accepted for 
Hypoestes then H aristata would represent a tetraploid species in the genus. A haploid chromosome 
number of 15 is also known in Dicliptera, Peristrophe, and Rhinacanthus (see below), all southern 
African relatives of Hypoestes in tribe Justicieae, subtribe Diclipterinae. Our collection of//, aristata 
is referable to var. alha K. Balkwill. 
Isoglossa Oerst. Isoglossa consists of about 60 species native to the Old World. Welman ( 1 993) 
listed 1 5 species as occurring in southern Africa. Our counts of /7 = 1 7 for two of these, /. hypoestifJora 
(southern and tropical Africa) and /. ovata (southern Africa), agree with the sole previous report of a 
chromosome number in the genus. Daniel and Chuang ( 1 998) reported n= 1 7 for the tropical eastern 
African species /. grandiflora C. B. Clarke. 
Isoglossa was treated by Bremekamp in tribe Justicieae, subtribe Rhytiglossinae 
(= Isoglossinae). McDade et al. (in press) demonstrated that Isoglossa is part of a "core" Isoglossinae 
that is strongly supported as monophyletic. Chromosome numbers have been determined for only two 
other genera in this subtribe; Daniel ( 1 999) reported « = 1 8 for both Stenostephanus Nees (including 
Hahracanthus Nees and Hansteinia Oerst.) and Razisea Oerst. Although a basic number ofx = 1 7 ap- 
pears likely for Isoglossa, and may represent dysploid evolution from an ancestor with a: = 1 8, a 
well-substantiated basic number for the subtribe remains to be determined. 
Justicia L. Justicia is the largest genus of Acanthaceae with estimates of about 700 species occur- 
ring worldwide (McDade et al. in press). Twenty-two species were recognized by Immelman ( 1 995b) 
in the treatment of Acanthaceae: Justicieae for the Flora of Southern Africa. Several other genera 
from the region that are recognized in the Flora of Southern Africa (i.e., Adhatoda Mill., Aulojusticia, 
Duvernoia, Siphonoglossa Oerst.) are sometimes included in Justicia (Graham 1988, Brummitt 
1 992) as well. Daniel (2000) noted the presence, frequency, and distribution of 2 1 chromosome num- 
bers reported for 93 species in the genus. He noted the prevalence of/? = 1 4 throughout JM5//c/a and a 
DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 
149 
^n a 
4 
•'•*•* 
• f 
f ^ 
4 
0% 
. ' ■*-h 
•te 
«• 
•:« 
\ 
10|im 
I 
\ 
c 
^ (^fi 
e f 
g f 
Figure 2. Camera-Iucida drawings of meiotic chromosome preparations, a. Justicia betonica {Daniel et al. 9334), n = 17 
(metaphase I), b. Isoglossa hypoestiflora {Daniel et al 9341), n=\l (diakenesis I), c. Barleria senensis {Daniel et al 9375), 
n = 16 (telophase 1). d. Asystasia gangetica {Daniel et al 9348), n = 13 (metaphase 1). e. Aulojusticia linifolia {Daniel et al 
9388), n = 40 (metaphase I), f. Justicia odora {Daniel et al. 9364), « = 14 (diakenesis I), g. Crabbea angustifolia {Daniel et al 
9370), « = 21 (early metaphase I). 
1 50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Volume52, No. 12 
probable basic number of x = 7 for the genus. Within Justicia several sections are characterized by a 
diversity of chromosome numbers whereas others are homogeneous in the counts reported (Daniel 
2000). 
Our counts for three species of southern African Justicia illustrate the diversity of numbers re- 
ported in the genus. The counts of « = 1 7 for two collections of J. betonica agree with the majority of 
previous counts for this native of Africa and the Indian subcontinent (Daniel 2000, Daniel and Chuang 
1 998). Hedren ( 1 989) reported « = ca. 1 3 from a Tanzanian collection of the African and Arabian spe- 
cies,/, odora. Our count for this species, based on a collection from South Africa, is « = 14. This num- 
ber has been reported previously in 1 of the 1 3 sections of Justicia recognized by Graham ( 1 988) in 
which at least one chromosome number has been reported. This is the first report of /? = 14 in section 
Harnieria (Solms) Benth., however. Our count of « = 26 in J. petiolaris, a species occurring in eastern 
South Africa and Swaziland, represents the first report of a chromosome number for the species. It is 
also the first report of this chromosome number in the genus. Graham ( 1 988) treated J. petiolaris in 
sect. Tyloglossa (Hochst.) Lindau. The only other chromosome number known among the species in- 
cluded in this section by Graham is « = 13, which has been reported several times (e.g., Mangenot and 
Mangenot 1962, Podlech 1 986) mJ.flava (Vahl) Vahl. Our collection of J. petiolaris pertains to sub- 
species /?e//o/flm. Chromosome numbers reported here for South African species of Justicia reflect 
the putative polyploid and dysploid evolution seen for the genus in other regions (Daniel 2000). Given 
the large number of species of Justicia, its worldwide distribution, and the diversity of chromosome 
numbers already reported within it, additional cytological studies of the genus are highly desirable. 
Metarungia Baden. This genus comprises three species occurring in eastern and southern Africa 
(Baden 1 98 1 , 1 984). Our count ofn = 1 4 for M longistrohus, a species native to South Africa, Swazi- 
land, and Mozambique, is the first report of a chromosome number in the genus. Metarungia is closely 
related to Rungia Nees in the tribe Justicieae and these genera are sister to a clade that includes Justicia 
(McDade et al. in press). Numerous counts have been reported for each of four species of Rungia. 
Counts for three of these (i.e., R. laeta C. B. Clarke, R. parviflora Nees, and R. pectinata (L.) Nees) are 
mostly n = \3 or n = 26. Most counts for Rungia repens Nees are n = ]0. Although « = 14 is not 
known in Rungia, this is the most commonly reported number in Justicia (Daniel 2000). 
Pseuderanthemum Radlk. Pseuderanthemum consists of about 60 species of perennial herbs and 
shrubs occurring in both the Old and New Worlds. Two species are known from southern Africa, P. 
subviscosum (C. B. Clarke) Stapf andP. hildebrandtii (Welman 1993; Edwards and Harrison 1998). 
Our count ofn = 21 for P. hildebrandtii, which is native to southern Africa and tropical east Africa, is 
the first report of a chromosome number for this species and for an African species of the genus. Chro- 
mosome numbers have been reported for seven other species of Pseuderanthemum native to America, 
southern Asia, and the Pacific Islands (Daniel and Chuang 1989, 1998; Daniel 2000). Six of these 
have meiotic complements of 2 1 . Kaur ( 1 969) reported « = 30 for the Fijian species, P. laxiflorum (A. 
Gray) F. T. Hubb. As noted by Daniel and Chuang ( 1 998), a basic number of x = 2 1 appears likely for 
this genus as well as for several of its relatives in the Justicieae. McDade et al. (in press) identified a 
lineage of Justicieae, including Pseuderanthemum, characterized by an androecium of two stamens 
and two staminodes and by x = 21. New World relatives include Chileranthemum Oerst., 
Odontonema Nees, and Oplonia Raf Old World relatives include Ruspolia and Ruttya. 
Rhinacatithus Nees. Rhinacanthus comprises about 20 species occurring in Africa, Madagascar, 
and Asia. Three species are known in southern Africa (Balkwill 1995). One of these, R. gracilis, is 
sometimes cultivated. Daniel and Chuang ( 1 998) reported a count ofn = 1 5 from cultivated materials 
of this species. Our count of n = 1 5 from two wild-collected plants of the same species confirms the 
previous count. All other counts reported for species of Rhinacanthus are likewise « = 1 5 (Daniel and 
DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 
151 
■ V- -v. 
'•'.■■ 
10 p,m 
<-> 
•• • • - • 
« • 
<-> 
AD 
i ♦ 
<-> 
'^' 
• 
Figure 3. Camera-lucida drawings of meiotic chromosome preparations, a. Hypoestes aristata {Daniel et al. 9351), « = 30 
(telophase II, only half of cell shown), b. Rhinacanthus gracilis {Daniel 9340), /i = 1 5 (metaphase I), c. Dicliptera 
magaliesbergensis {Daniel et al. 9357), «= 13 (telophase I), d. Ruspolia hypocrateriformis {Daniel et al 9359), n = 2\ 
(diakenesis I), e. Hypoestes forskaolii {Daniel et al. 9358), /i = 15 (diakenesis I), f Metarungia longistrobus {Daniel et al. 
9555), n= 14 (telophase 1). 
1 52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Volume 52, No. 12 
Chuang 1 998). Thus, a basic number of x = 1 5 can be advanced for the genus. Based on morphological 
characteristics Rhinacanthus would appear to be best treated in tribe Justicieae, subtribe Justiciinae 
(Balkwill and Getliffe Norris 1 988), but molecular sequence data (McDade et al. in press) place it in 
subtribe Diclipterinae with strong support. Cytological data corroborate this latter placement. Other 
Diclipterinae (i.e., Dicliptera, Hypoestes, and Peristrophe) also have probable basic chromosome 
numbers of 1 5 (see above). 
Nomenclatural and taxonomic confusion surrounding R. gracilis was discussed by Balkwill 
( 1 995), who recognized two varieties of the species in southern Africa. Plants used for our counts per- 
tain to R. gracilis var. latilabris K. Balkwill. The voucher specimen of Daniel and Chuang's (1998) 
count pertains to R. gracilis var. gracilis. 
Ruspolia Lindau. Six species of Ruspolia have been described from Africa and Madagascar and 
three of them occur in southern Africa ( Welman 1 993). Our count of /? = 21 for R. hypocraterifbrmis, 
a species native to western tropical Africa and South Africa, is the first report of a chromosome num- 
ber for the species. It agrees with previous counts of « = 21 for R. seticalyx (C. B. Clarke) 
Milne-Redh. (Daniel and Chuang 1 998). This same chromosome number has been reported in several 
relatives of Ruspolia in the Justicieae (see above under Pseuderanthemum). Plants from South Africa 
are commonly treated as R. hypocrateriformis var. australis Milne-Redh. (e.g., Heine 1963). 
Ruttya Harv. Ruttya comprises three species native to Africa. Only one of these, R. ovata, occurs 
in southern Africa where it is found in South Africa and Swaziland. Our count of/? = 2 1 for the species 
agrees with the only previous report of a chromosome number in the genus. Daniel and Chuang ( 1 998) 
reported this number for R.fruticosa Lindau, a native of eastern tropical Africa, based on cultivated 
plants. The affinities of Ruttya are with Ruspolia and other genera of tribe Justicieae with an androe- 
cium of two stamens and two staminodes andx = 21 (see above under Pseuderanthemum). Morpho- 
logical distinctions among Ruttya, Ruspolia, and Pseuderanthemum were noted by Daniel and 
Chuang (1998). 
Conclusions 
As shown by McDade et al. (in press), knowledge of chromosome numbers can be useful for sug- 
gesting or corroborating phylogenetic relationships among Acanthaceae. Even in this small sampling 
of taxa from South Africa, the importance of knowledge of chromosome numbers among 
Acanthaceae is evident. A continuity of chromosome numbers between southern Afi-ican endemics 
and congeners from other regions provides additional evidence for several previously proposed 
suprageneric relationships based on morphology and molecular markers (e.g., 
Hypoestes-Dicliptera-Peristrophe and Pseuderanthemum-Ruttya-Ruspolia). However, the chromo- 
some numbers reported for Rhinacanthus support molecular rather than morphological data in plac- 
ing this genus in the Diclipterinae rather than in the Justiciinae. Based on the treatment of Justiciinae 
in the Flora of Southern Africa (Baden et al. 1 995), Aulojusticia and Duvernoia are recognized here. 
Using morphological characteristics, both might be readily accommodated in a broadly interpreted 
Justicia. However, their respective chromosome numbers are either unknown or uncommon in that 
genus and their systematic affinities based on molecular markers are currently being studied. 
It will be useful to obtain additional chromosome numbers among Acanthaceae from southern 
Africa by sampling more taxa from South Africa as well as from other countries in the region. For ex- 
ample, chromosome numbers remain unknown for the following acanthaceous genera that occur in 
southern Africa: Acanthopsis Harv., Anisotes Nees, Chaetacanthus Nees, Chorisochora Vollesen, 
Duosperma Dayton, Glossochilus Nees, Megalochlamys Lindau, Neuracanthus Nees, and 
Ruelliopsis C. B. Clarke. Also, no chromosome counts have been determined for Acanthaceae from 
DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 1 53 
any southern African country except South Africa. An increase in the number of taxa for which chro- 
mosome numbers are known would help to identify basic numbers for additional genera and assist ef- 
forts to understand phylogenetic relationships within the family. 
Acknowledgments 
We are grateful to Fei Mei Chuang for technical assistance with the chromosome counts, R. K. 
Brummitt and M. Butterwick for assistance in the field, J. and S. Williamson for logistical assistance 
and for allowing us to collect plants in their garden, and the Witwatersrand National Botanical Garden 
and the Lowveld National Botanical Garden for permitting us to collect buds and vouchers of 
Acanthaceae. Daniel's field work in South Africa was funded by the In-House Research Fund of the 
California Academy of Sciences and the field work of K. and M.-J. Balkwill was funded by the Na- 
tional Research Foundation of South Africa and the University of the Witwatersrand 's Herbarium 
Programme. 
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Staatssamml. Miinchen 22:5-20. 
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chromosomiques de quelques angiospermes du Rwanda, du Burundi et du Kenya. Bull. Jard. Bot. Belg. 
53:343-371. 
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DANIEL, BALKWILL, and BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 1 55 
Appendix 
Species occurring in southern Africa for which chromosome numbers have been pubMshed previ- 
ously. The taxon name is followed by the author of the name, the chromosome number(s) reported 
(listed as n numbers), literature citation(s), and the country from which the plants studied came. The 
provenance of the plants is based on information provided in the original publications. Two references 
with chromosome numbers (both Ph.D. theses) have not been seen and the probable provenance is in- 
dicated with a question mark and "not seen" follows. In other publications, where the source of mate- 
rials studied is unclear or not provided, a question mark is indicated. The provenance of cultivated 
plants noted below is unknown. 
Acanthus pubescens (Oliv.) Engl., n = 28, Renard et al. 1983 (Burundi). 
Asystasia gangetica (L.) T. Anderson (including /I. coromandeliana Nees), « = 1 3: Mangenot and Mangenot 
1957 (western Africa), Mangenot and Mangenot 1 962 (western Africa), Gadella 1 977 (Cameroun), Ugborogho 
and Adetula 1988 (Nigeria); 14: Subramanian and Govindarajan 1980 (India), Govindarajan and Subramanian 
1983 (India); 22: Narayanan 1951a (India?); 24: Narayanan 1951a (India?); 25: De 1966 (India?), Sarkar et al. 
1978 (India, cultivated); 26: Narayanan 195 lb (India), Grant 1955 (U.S. A., cultivated), Ellis 1962 (India), Kaur 
1965 (India, cultivated), Valsala Devi and Mathew 1982 (India), Saggoo and Bir 1983 (India), Saggoo and Bir 
1986 (India), Daniel 2000 (U.S. A., naturalized). 
Barleria repens Nees, n = 20: Daniel and Chuang 1998 (cultivated). 
Blephahs integrifolia (L.f ) E. Mey. (including B. tnolluginifolia Pers. and B. repens (Vahl) Roth), n= \1\ 
Kaur 1966 (India, probably cultivated), Sareen and Sanjogta 1976 (India), Ranganath 1981 (India? not seen), 
Krishnappa and Ranganath 1982 (India). Ranganath and Krishnappa 1982 (India); « = 18: Govindarajan and 
Subramanian 1983 (India), Subramanian and Govindarajan 1980 (India). 
B. maderaspatensis (L.f) Heyne ex Roth (including B. boerhaaviaefolia Pers.), « = 8: Valsala Devi and 
Mathew 1 982 (India), Saggoo 1 983 (India?, not seen), Saggoo and Bir 1 982a (India), 1 983 (India), 1 986 (India), 
Govindarajan and Subramanian 1983 (India); «= 11: Ranganath 1981 (India? not seen), Krishnappa and 
Ranganath 1982 (India), Ranganath and Krishnappa 1982 (India); n= 12: Ranganath and Krishnappa 1982 (In- 
dia); «=13: Kaur 1966 (India), Sareen and Sanjogta 1976 (India); n=14: Vasudevan 1976 (India), 
Subramanian and Govindarajan 1980 (India); n= 15: Miege 1962 (Senegal), Bir and Saggoo 1979 (India), 
Saggoo 1983 (India? not seen), Bir and Saggoo 1981 (India), Saggoo and Bir 1982b (India), 1983 (India); 
2n = 23: Ranganath and Krishnappa 1982 (India). 
Crabbea velutina S. Moore (including C. reticulata C. B. Clarke), n = ca. 14: Renard et al. 1983 (Rwanda); 
« = 21, Grant 1964 (cultivated). 
Dyschoriste depressa Nees, n= 15: Govindarajan and Subramanian 1985 (India); n = 30: Bir and Saggoo 
1979 (India), 1981 (India), Saggoo 1983 (India? not seen), Saggoo and Bir 1982b (India). 
Elytraria acaulis (L.f) Lindau, n= \1: Kaur 1 969 (India?); n = 22: Govindarajan and Subramanian 1 983 (In- 
dia); n = 23: Subramanian and Govindarajan 1980 (India). 
Hygrophila auriculata (Schum.) Heine (including Asteracantha longifolia), «= 16, De 1966 (India?), 
Trivedi and Trivedi 1992 (India), Subramanyam and Kamble 1971 (India), Saggoo 1983 (India? not seen), Bir 
and Saggoo 1981 (India), Saggoo and Bir 1982b (India), 1986 (India). 
Hypoestes aristata R.Br., n = 30, Daniel and Chuang 1998 (cultivated). 
H.forskaollii Vahl, « = 15, Podlech 1986 (Yemen). 
156 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Volume 52, No. 12 
//. /n//oraRoem.andSchuIt.,« = l5:Saggoo 1983 (India? not seen), Sagoo and Bir 1 982a (India), 1983 (In- 
dia). 
Justicia anagalloides (Nees) T. Anderson, n = 9: Ensermu Kelbessa 1990 (Ethiopia); n= 18: Enseimu 
Kelbessa 1 990 (Ethiopia). 
J. betonica L., « = 1 4: Subramanian and Govindarajan 1 980 (India), Narayanan 1 95 1 b (India?); n= 17: Ellis 
1962 (India), Ranganath 1981 (India?, not seen), Bir and Saggoo 1 979 ( India), Krishnappa and Ranganath 1982 
(India), Saggoo 1983 (India?, not seen), Valsala Devi and Mathew 1 982 (India), Govindarajan and Subramanian 
1 983 (India), Bir and Saggoo 1 98 1 (India), Saggoo and Bir 1 982b (India), 1 986 (India), Daniel and Chuang 1 998 
(cultivated), Daniel 2000 (U.S. A., naturalized). 
J. exigua S. Moore, n = 7: Renard et al. 1983 (Rwanda), Ensermu Kelbessa 1990 (Ethiopia, Tanzania). 
J. flava (Forssk.) Vahl, n= 13: Mangenot and Mangenot 1957 (western Africa), 1962 (western Africa), 
Podlech 1 986 (Yemen). 
J. glabra Koenig ex Roxb., n = 27: Ranganath 1981 (India? not seen), Krishnappa and Ranganath 1982 (In- 
dia). 
J. matammensis (Schweinf ) Oliv., n = 7: Ensermu Kelbessa 1 990 (Ethiopia). 
J. odora (Forssk.) Lam., /; = ca. 13: Hedren 1989 (Tanzania). 
Mackaya bella Harv., n = 42: Daniel and Chuang 1989 (cultivated). 
Nelsonia canescens (Lam.) Spreng., n= 17: Saggoo 1983 (India? not seen), Saggoo and Bir 1983 (India); 
« = 18: Daniel and Chuang 1993 (Panama). 
Pehstrophe paniculata (Forssk.) Brummitt (including P. bicalyculata (Retz.) Nees), n= 10: Narayanan 
1951b (India); n= 15: Ahuja 1955 (India), Miege and Josserand 1972 (Senegal), Vasudevan 1976 (India), 
Saggoo and Bir 1983 (India), 1986 (India), Verma and Dhillon 1967 (India). 
Phaulopsis imbhcata (Forssk.) Sweet, « = 16: Manktelow 1996 (Ethiopia, Tanzania, Madagascar); « = 17: 
Daniel and Chuang 1998 (cultivated); n = ca. 32: Manktelow 1996 (Madagascar); 2n = 65: Manktelow 1996 
(Malawi). 
Rhinacanthus gracilis Klotz., « = 15: Daniel and Chuang 1998 (cultivated). 
Ruellia cordata Thunb., /? = 16, Rao and Mwasumbi 1981 (Tanzania, probably cultivated). 
R. patula Jacq., n= 16: Baquar et al. 1966 (Pakistan), Rao and Mwasumbi 1981 (Tanzania, probably culti- 
vated), Govindarajan and Subramanian 1983 (India), Subramanian and Govindarajan 1980 (India). 
Ruspolia seticalyx (C. B. Clarke) Milne-Redh., « = 21: Daniel and Chuang 1998 (cultivated). 
Thunbergia alata Boj., « = 9: Grant 1 955 (cultivated), Takizawa 1 957 (cultivated), Shibata 1 962 (Colombia, 
naturalized), Saggoo 1 983 (India?, not seen), Bir and Saggoo 1 98 1 (India), Saggoo and Bir 1 982b (India), 1 986 
(India), Daniel and Chuang 1989 (cultivated), Sugiura 1931 (cultivated?), Tijo 1948 (Indonesia), Narayanan 
1951b (India), Kaur 1970 (India?); /? = 16: Snoad 1952 (cultivated). 
DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 1 57 
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