262 AMERICAN JOURNAL OF BOTANY [Vol.

50

PROUT, 1'., C. HUEB8CHMAN, H. LEVEN~;, AND F. J.
RYAN. 1953. The proportions of nuclear types in
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RAPER, J. R., AND P. G. MILES. 1958. The genetics of
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- - - , AND J. P. SAN ANTONIO. 1954. Heterokaryotic
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CYTOTAXONOMY OF TWELVE SPECIES

OF HIBISCUS SECTION FURCARIA 1
MARGARET Y. MENZEL AND F. D. W ILSON2
Department of Biological Sciences, Florida State University, Tallahassee, Florida
and
Everglades Experiment Station, Belle Glade, Florida

ABSTRACT
MENZEL, MARGARET Y. (Florida State U., Tallahassee), and F. D. WILSON. Cvtotaxonomv of
twelve species of Hibiscus section Furcaria. Amer. Jour. Bot. 50(3): 262-271. IIIus. 19li3.-Meta-
phase-I chromosome numbers and pairing in 88 accessions showed that H. cannabinus, H. costatus,
and H. surattensis are diploid (n = 18); and H. acetosella, H. aculeatus, H. bijurcaius, H.furcellatus,
H. meeusei, H. radiatus, H. rostellatus and H. sabdariiJa are tetraploid (n = 36), with similar low
multivalent frequencies, hence probably allotetraploids each combining 2 well-differentiated
genomes. No intraspecific variation in ploidy was found. Fertile, vigorous F, hybrids between
H. furcellatus and H. bifurcatus showed complete chromosome pairing (n = 36), confirming a close
relationship between the parents. Two African strains of H. diversifolius were octoploid (n = 72)
with low multivalent frequency and hence probably contain 4 differentiated genomes. At least 4,
perhaps 5 or 6, differentiated genome groups are represented in tropical Africa, and at least 2 in
the American tropics.

A SHORTAGE of jute in world markets, which susceptibility to root-knot nematodes (Summers,

began during World War II and has become more
acute as jute-producing regions consume ever
larger percentages of their own output, has led
to interest in jute substitutes less demanding in
cultural requirements and more amenable to eco-
nomical production in other parts of the world
(cf. Bally, 1955a, b). Two species of the genus
Hibiscus (Malvaeeae), H. cannabinus L. ('kenaf')
and H. sabdariffa L. ('roselle'), are among the
more promising substitutes. One of the chief ob-
stacles to cultivation of kenaf in Florida is its
, Received for publication August 8, 1962.
This report is based on research conducted cooperatively
by the Crops Research Division and the Agricultural
Engineering Research Division, Agricultural Research
Service, United States Department of Agriculture, and
the University of Florida Agricultural Experiment Sta-
tion, Everglades Experiment Station, Belle Glade, Florida.
The senior author acknowledges the courtesy of the De-
partment of Biological Sciences, Florida State University,
in making available laboratory space and other facilities
for the cytological studies. Thanks are extended to the
many organizations and individuals, some of whom are
not mentioned in the paper, whose courtesy and coopera-
tion made it possible to assemble the living. materials and
information on which this study was based."
2 Geneticists, Crops Research Division, Agricultural Re-
search Service, United States Department of Agriculture.
Pate, and Wilson, 1958). Genetic resistance exists
in H. acetosella Welw. ex Hiern (Pate, Summers,
and Menzel, 1958) and H. sabdariffa L., and to a
lesser degree in H. radiatus Cav. Hibiscus sobda-
riffa has not been successfully crossed with H.
cannabinus. The tetraploid species H. acetosella
and H. radiatus cross readily with diploid H.
cannabinus (Menzel and Wilson, 1961b), but the
triploid hybrids are practically sterile and the
fertile, colchicine-induced H. cannabinus-acetosella
allohexaploid is agronomically unsatisfactory.
Hence, a survey of other relatives of H. cannabinus
and H. sabdariffa was undertaken for additional
sources of genetic variation.
Hibiscus cannabinus and H. sabdariffa belong
to section Furcaria DC. of the genus, charac-
terized by a leathery or succulent fruiting calyx
with 10 prominent nerves or ribs: each of the .5
calyx lobes has a medial nerve, and a commissural
nerve passes to each sinus and then branches to
form a prominent ridge along the margin of each
lobe. The epicalyx bracts (bracteoles) are nu-
merous, free or joined to the calyx only at the
base, and frequently divided at the tip into an
inner and an outer fork. Section Furcaria, as de-
March, 1963] MENZEL AND WILSON-CYTOTAXONOMY OF HIBISCUS
limited by Hochreutiner (1900) in the most recent
comprehensive monograph of Hibiscus, comprises
a morphologically well-defined pan-tropical group
of 40 or more species which differ from most other
species of Hibiscus in the basic chromosome
number 18 and could logically be considered a
separate genus (Skovsted, 1941). The present
paper presents cytotaxonomic data for 12 species
of Hibiscus section Furcaria,
MATERIALS AND METHODs-All accessions in-
cluded in the present study except those of H.
aculeaius were grown from seeds at the Everglades
Experiment Station farm near West Palm Beach,
Palm Beach County, Florida. Hibiscus aculeatus
was represented by wild populations in Leon,
Franklin, and Taylor Counties, Florida. Addi-
tional plants of certain accessions were also grown
in Tallahassee in garden and greenhouse.
Buds for cytological study were peeled to ex-
pose the staminal column, fixed in fresh acetic
alcohol (1 :3), transferred after 3 or more days to
fresh acetic alcohol, and stored in a food freezer
at about -15 C. Meiotic chromosomes were
studied in temporary iron-acetocarmine squashes
of pollen mother cells.
The living accessions were identified by ref-
erence to H:ochreutiner's monograph (1900), to
....
4A.
..-
~
~
.....
.....
~
~~­
~
• _~
.-.
~,..,
~
~
~
~
• 3
Fig. 1-5. Chromosomes at metaphase I in pollen mother cells of species of Hibiscus sect. Furcaria. Drawn with the
aid of a camera lueida, X 1450.-Fig.1. H. meeusei A60-176, 36 bivalents.-Fig. 2. H.furcellatus A59-86, 36 bivalents.-
Fig. 3. H. sabdariffa A59-57, 35 bivalents, 2 univalents.-Fig. 4. H. aculeatus A59-117, 36 bivalents.-Fig. 5. H. di-
versifolius A59-94, 72 bivalents,
263
various regional floras and individual species
descriptions (Masters, 1868; Hiern, 1896; Sprague,
1913; Ulbrich, 1921, 1922; Hutchinson and Dalziel,
1928; Exell, 1959, 1961; Roe, 1961), and to selected
herbarium material from the Conservatoire et
Jardin Botanique, Geneva, Switzerland; the In-
stitute of Botany, Faculty of Science, Lisbon,
Portugal; and the Royal Botanic Gardens, Kew,
Great Britain. Voucher specimens of all accessions
of wild species and many of the cultivars studied
have been deposited in the Herbarium of Florida
State University (Table 1).
OBSERVATIONS-All 12 of the species studied
(Table 1) had multiples of the basic chromosome
number 18 established previously for section
Furcaria (Skovsted, 1941). Chromosome numbers
for H. aculeaius (Fig. 4), H. bifurcatus, H. fur-
cellaius (Fig. 2), H. meeusei (Fig. 1) and H.
rostellatus apparently have not been reported
previously except in an abstract of the present
work (Menzel and Wilson, 1961a). The species dis-
cussed by Skovsted (1935, 1941) as H. bifurcatus
Cav. was later identified as H. radiatus Cav,
(Menzel and Wilson, 1961b). H. cannabinus, H.
surattensis and H. cosiaius were diploid, H. diversi-
folius was octoploid (Fig. 5), and the other 8
species were tetraploid.
~
~
~
.......,.,,~
~""'~~..-IfiI""
. ; /,........
264 AMERICAN JOURNAL OF BOTANY [Vol. 50

TABLE 1. Chromosome numbers in species of Hibiscus sect. Furcaria

Plant Chromosome
Species and Florida accession Introduction Voucher number
number or previous authority number number n 2n Source and remarks

A. OLD WORLD SPECIES

H. acetosella Welw. ex Hiern

Skovsted, 1941 72 Trinidad, as H. sp, "African red"
Menzel and Wilson, 1961b HVI 36 Old station stock, Everglades Exp.
Station; red leaves, rose flowers
"Merritt Island" HV13 36 Merritt Island, Fla., cult., from S. E.
McFadden, Jr., Dept. Ornamental
Horticulture, Fla. Agric. Exper. Sta-
tion, Gainesville; green leaves, yellow
flowers
A59-126 260164 HV65 36 Botanic Garden, Osaka City Univer-
sity, Japan; red plants, rose flowers
A59-152 260997 HV64 36 Institut National pour l'etude Agrono-
mique du Congo Belge (INEAC),
Yangambi, as H. eetoeldeanus; "P.
Ramps 649"; green, deeply lobed
leaves, yellow flowers
A60-240 265325 HV63 36 Estacion Experimental Agron6mica,
Santiago de las Vegas, Cuba, as H.
sabdarijJa; red plants, .rose flowers
H. asper Hook.'
Skovsted, 1941 72 Northern Rhodesia
H. cannabinus L.
Breslavetz et al., 1934 36
Skovsted, 1935 18 36 Trinidad
Rao, 1935 72 (India?)
Medwedewa, 1936 36 12 strains
Ford, Hl38 ca. 72 Ceylon
Skovsted, 1941 36 4 strains
Tjio, 1948 36 2 strains
BG52-1 18 Selection from Salvadorian kenaf,
A51-43\ PI207883, introduced to
Florida by.....E. Henriquez, Tropical
Fibers Inc.,' iVero Beach, Fla.
BG52-20 18 Ditto
BG52-41 HV70 18 Ditto
BG52-44 18 Ditto
BG52-71 HV69 18 Ditto
BG52-122 HV71 18 Salvadorian (PI207883) X Manchur-
ian (PI207896)
BG52-135 HV74 18 Selection from Javanese (PI207897)
BG53-43 18 Salvadorian (PI207883) X Javanese
(PI207900)
BG53-46 18 Ditto
BG53-53 18 Ditto
BG53-90 HV20 18 Egypt (PI209231) var. ECR-986, from
Ministry of Agriculture, Wad Me-
dani, Sudan
A58-7 HV8 18 J. M. Dempsey, Viet Nam
A59-80 257496 18 Nigeria, via Tropical Products Inst.,
London
A59-147 18 "Cuba 2032L," Est. Exp. Agron.,
Cuba
A60-180 263202 HV49 18 A. N. Prentice, Ministry of African
Agric., Mt. Makulu Res. Station,
Chilanga, N. Rhodesia
A6D-194 263216 HV48 18 A. N. Prentice, N. Rhodesia
March, 1963] MENZEL AND WILSON-CYTOTAXONOMY OF HIBISCUS 265

TABLE 1. Continued

Plant Chromosome
Species and Florida accession Introduction Voucher number
number or previous authority number number n 2n Source and remarks

A60-206 263964 18 D. G. Thomas, Dept. of Agriculture,

Kawanda Res. Station, Kampala,
Uganda; calyx not wooly
A307 244862 18 "Cuba 689b-2," Est. Exp. Agron.,
Cuba
H. dioersijolius Jacq.
Medwedewa, 1936 144 Kirstenbosch, 8. Africa
Ford,1938 ca. 72 S. Africa
Skovsted, 1941 ca. 180 Fiji
Skovsted, 1941 ca. 180 Colombia
A59-94 259173 HV22 72 H. B. Bycroft, National Botanic Gar-
dens, Kirstenbosch, Cape Province,
Union of S. Africa
A60-207 263965 HV87 72 D. G. Thomas, Uganda
H. meeusei Exell a
A.59-84 258739 HV44 36 A. D. G. Meeuse, from the type lo-
cality near Pretoria, Union of S.
Africa
A6D-172 263194 HV79 36 A. N. Prentice, N. Rhodesia
A60-176 263198 36 A. N. Prentice, N. Rhodesia
A60-181 263203 HV51 36 A. N. Prentice, N. Rhodesia
A60-197 263219 HVS2 36 A. N. Prentice, N. Rhodesia
H. radiatus Cay.
Skovsted, 1941 72 Trinidad (as H. bijurcatus Cav.)
Tjio, 1948 72 Java ("H. radiatus Willd.")
A214, Menzel and Wilson, 1961b 200834 HV114 36 O. E. White, Mandalay, Burma
A59-53 256035 HV21, 36 Pakistan Central Jute Committee,
HV67 Jute Research Institute, Dacca
A59-148 HVl13 36 Est. Exp. Agron., Cuba, as H. diversi-
[olius No. 19
A59-149 HV111 36 Est. Exp. Agron., Cuba, as H. dioersi-
folius No. 29
A60-231 HVll0, 36 Est. Exp. Agron., Cuba, No. 710b;
HVl18 cream flowers
A60-232 HV109, 36 Est. Exp. Agron., Cuba, No. 712a-2;
HV117 purple flowers
A60-233 HV108 36 Est. Exp. Agron., Cuba, No. 712a-3;
purple flowers
A61-384 275936 HV107 36 T. V. Ramana, Botany Dept., Andhra
Univ., Waltair, India; as H. [urcaius
Willd.; cream flowers
HV116 36 S. E. McFadden, Fla. Agr. Exp. Sta.
No. S60m39
HV11.5 36 S. E. McFadden, Fla. Agr. Exp. Sta.
No. 855m15
H. rostellatus Guill. et Per.
A59-150 260998 HVM 36 INEAC, Yangambi, Belgian Congo,
"P. Bamps 652"
A60-211 263969 HV53 36 D. G. Thomas, Uganda
H. sabdariffa L.
Skovsted, 1935 36 72 Trinidad
Rao, 1935 72
Medwedewa, 1936 72 2 strains
Skovsted, 1941 36 N. Rhodesia, 2 types
Skovsted, 1941 36 Sudan
Skovsted, 1941 72 Nyasaland
Tjio, 1948 72 Java
2GG AMERICAN JOURNAL OF BOTANY [Vol. 50

TABLE 1. Continued

Plant Chromosome
Species and Florida 'accession Introduction Voucher number
number or previous authority number number n 2n Source and remarks

A51-44 207920 HV27 36 E. Henriquez, commercial roselle

3695 36 Kilgore Seed Co., commercial roselle
A52-31 180026 36 India, "var. altissima"
A58-5 HV3 36 J. M. Dempsey, Thailand, "Thai red"
A58-6 HV25 36 J. M. Dempsey, Viet Nam
A58-31 249639 HVlO 36 New Delhi, India
A59-56 256038 HV28 36 Pakistan Central Jute Committee, Jute
Researeh Institute, Daeea
A59-57 256039 HVll 36 Pakistan Central Jute Committee
A59-58 256040 HV4 36 Pakistan Central Jute Committee
A59-59 256041 HV9 36 Pakistan Central Jute Committee
A59-68 256985 HV24 36 Joint Commission on Rural Recon-
struetion, Taipei, Taiwan, as H.
punqens
A59-92 259162 HV5 36 M. F. Rose, Gezira Research Station,
Wad Meduni, Sudan; from Kordofan
District, "kurknde"
A59-93 259163 HV2 36 M. F. Rose, Kordofan District, Sudan,
"karkade"
A60-200 263222 36 A. N. Prentice, N. Rhodesia
A60-202 263224 36 A. N. Prentiee, N. Rhodesia
A60-216 263749 36 A. N. Prentice, N. Rhodesia
A60-234 265319 36 Est. Exp. Agron., Cuba
A60-23.'5 265320 36 Est. Exp. Agron., Cuba
A60-236 265321 36 Est. Exp. Agron., Culm
A60-237 265322 36 Est. Exp. Agron., Cuba
A60-238 265323 36 Est. Exp. Agron., Cuba
A60-250 267778 36 Ministry of Agric., Sudan
A61-324 36 J. M. Dempsey, U. S. Operations Mis-
sion, Viet Nam
A61-328 36 USOM, Viet Nam
A61-329 36 USOM, Viet Nam
A61-330 36 USOM, Viet Nam
A61-332 36 USO:.YI, Viet Nam
A61-336 273388 36 C. Y. Chi, Taiwan Agric. Res. Inst.,
Taiwan
A61-338 273390 36 C. Y. Chi, Taiwan
A61-340 273392 36 C. Y. Chi, Taiwan
A61-341 273459 36 Dept. of Agric., Pretoria, Union of S.
Africa, originally from Thailand
A61-356 274246 36 Rural Ministere de l'Eeonomie, Ma-
radi, Nigeria, originally from Senegal
A61-357 274247 36 Rural Ministere de I' Eeonomie, Ma-
radi, Nigeria
A61-370 27.5413 36 Centre de Recherches Agronomique,
Bambey, Senegal
A61-371 275414 36 Centre de Rech. Agron., Senegal
Saier's roselle 36 S. E. McFadden, Fla. Agrie. Exp. Sta.
No. S60m35, originally from Saier
Seed Co., Dimondale, Miehigan
H. surottensie L.
Medwedewa, 1936 72 As "H. bifurcaius = H. eurattensis ..."
Skovsted, 1941 36 Kenya
A59-70 258429 HV56 18 D. G. Thomas, Uganda
A59-151 260999 HV55 18 INEAC, Yangambi, Belgian Congo,
"P. Bamps 648"
B. NEW WORLD SPECIES

H. aculeatus Walt.
A59-1l7 59-4 36 Taylor Co., Fla.
A59-120 59-7 36 Taylor Co., Fla.
March, 1963] MENZEL AND WILSON-CYTOTAXONOMY OF HIBISCUS
TABLE 1. Continued
Plant
Species and Florida accession Introduction
number or previous authority number
H. bijurcatus Cav.
A.59-153 260796
H. costaius Rich.
Ford,1938
A60-243 265329
1I. [urcellalus Lam.
A.59-86
A59-87
A60-224
A61-359
• The 5 tetraploid strains referred to as H. asper Hook. f. in an abstract of the present study (Menzel and Wilson,
]fl6] a) were subsequently identified as H. meeusei Exell. The 2 species appear closely related, but H. asper occurs north
and west of the Congo Basin to Senegal, whereas H. meeusei ranges from Northern Rhodesia to Transvaal.
Some accessions of H. sabdar(fJa showed a tend-
ency to asynapsis in some samples of pollen mother
cells, particularly following cold nights in De-
cember, HHj9 (Table 2). At least one such line
(A59-93) had normal pairing in a later collection.
Other lines collected on the same date were not
asynaptic, which suggests genetic differences be-
tween strains. Frequency of multivalents ap-
peared unaffected by the presence of univalents
(Table 2).
Intraspecific variation in chromosome number-
In the present study all accessions of a given
species had the same chromosome number, al-
though more than one chromosome number has
been reported by other workers for H. cannabinus,
H. surattensis, H. sabdariffa, and H. dioersifoliu«
(Table 1).
Among the 89 strains of H. cannabinus grown
at the Everglades Experiment Station since 19.18,
18 were analyzed cytologically and found to be
diploid. Results of crosses involving many other
unanalyzed strains gave no indication that any
had a different chromosome number. Hochreutiner
(1900) treated H. cannabinus as a collective species
which encompassed H. radiatus Cav. (including
H. unidens Lind!. and H. Lindleyi Wall); H. asper
Hook.; and H. acetosella Welw. ex Hiern, all of
which are tetraploid. Hibiscus radiatus frequently
is confused with H. cannabinus in herbaria and
in seed collections. It seems probable that the
reports of Rao (193.5) and Ford (1938) of tetra-
ploid H. cannabinus actually refer to one of these
other species.
Two strains of H. suraitensis L. studied by us
and one studied by Skovsted (1941) were diploid,
but Medwedewa (1936) reported "H. b~furcatus
267
Chromosome
Voucher number
number n 2n Source and remarks
HV 46 36 Botanic Garden, Rio de Janeiro, Brazil,
as 1I. kitaibelifolius
36 Cuba
HV80 18 Royal Bot. Gardens, Kew, England;
originally from Cuba, via Singa-
pore Bot. Gardens
HV76 36 Military Trail, Palm Beach Co., Fla.
HV77 36 Belvedere Road, Palm Beach Co., Fla.
HV78 36 Botanical Gardens, Honolulu, Hawaii,
via D. R. Egolf, U. S. National Ar-
boretum, as H. youngianus
HV82 36 G. van den Berghe, Colombia, Dpto.
Magdalena
Roxb. = H. suraitensis" to have 2n = 72. Hoch-
reutiner (1900) treated under H. surattensis several
taxa, including the tetraploids H. rostellatus Guill.
et Per. and H. eetveldeanus De Wild. et Durand
(= H. acetosella Welw. ex Hiern) , which more
extensive material later showed to be distinct
species. Our diploid strains were H. suratteneis
sens. strict. (H. suratiensis val'. genuinus Hochr.).
It is probable that the tetraploid strain studied by
Medwedewa belonged to one of the other forms.
Most earlier workers found H. sabdar~ffa to be
tetraploid, but Skovsted (1941) reported 2 types
from Northern Rhodesia and 1 from Sudan to he
diploid. In view of the dearth of known diploids
in section Furcaria, diploid plants of H. sabdariffa
would be of great interest. We have counted the
chromosomes of 35 of the 57 accessions of H.
sabdar(lfa available, including a wide range of
morphological types; all were tetraploid. More-
over. chromosome pairing in these strains (Fig. 3;
Table 2) indicated that, like the other tetraploid
species, they are allotetraploids.
The variahle species H. dioersifoliu« Jacq. seems
to be the only species of section Furcaria native
to both the Old and New Worlds. Two strains
examined hy Skovsted (1941), one from Fiji and
the other from Colombia, were decaploid, whereas
our 2 African strains and 2 African strains re-
ported earlier (Medwedewa, 1936; Ford, 1938)
were octoploid. Hibiscus diversifolius mav include
two different polyploid complexes, an octoploid
African one and a decaploid New World form.
Relationships of the Old World species--Section
Furcaria achieves its greatest diversity and
number of species in tropical Africa, and it is
undoubtedly in this region that both H. canna-
268 AMERICAN JOURNAL OF BOTANY [Vol. 50

TABLE 2. Mean chromosome configurations in tetraploid species and hybrids of Hibiscus sect. Furcaria

Mean number per cell of

No.
Species or hybrid II III IV I cells

A. OLD WORLD SPECIES

H. cannabinus
colchicine-induced autotetra-
plaid- 15.6 0.3 9.6 1.5 10
H. acetosella
Old Stock- 34.0 0.1 0.6 1.2 42
"Merritt Island" 35.3 0.0 0.2 0.4 9
A59-126 35.7 0.0 0.2 0.0 9
A59-152 35.2 0.0 0.4 0.1 33
A60-240 34.8 0.0 0.6 0.2 9
H. meeusei
A59-84 35.3 0.0 0.3 0.5 4
A60-172 34.7 0.0 0.7 0.0 3
A60-176 35.9 0.1 0.0 0.1 15
H. radiatus
A214- 34.2 0.0 0.9 0.1 26
A59-53 36.0 0.0 0.0 0.0 13
A59-149 34.4 0.0 0.2 2.4 5
A61-384 35.4 0.0 0.3 0.0 7
H. acetosella-radiatus F 1
Old Stock X A214- 34.2 0.1 0.1 2.6 34
H. rostellatus
A59-150 3.'5.4 0.0 0.3 0.1 17
H. sabdariffa
Saier's (1960) 35.0 0.2 0.3 0.2 6
A51-44 (1960) 34.6 0.0 0.7 0.0 12
3695 (1959) 35.8 0.0 0.1 0.0 9
3695 (1960) 34.9 0.1 0.4 0.1 9
A52-31 (1959) 35.7 0.0 0.1 0.2 24
A58-5 (1960) 35.3 0.0 0.1 0.9 7
A58-6 (1959) 31.7 0.0 0.1 8.1 16
A58-31 (1959) 33.0 0.0 0.1 5.4 7
A59-56 (1959) 28.6 0.04 0.2 14.1 26
A.59-57 (1959) 34.4 0.0 0.0 3.0 7
A59-59 (1959) '33.8 0.1 0.3 2.8 24
A59-92 (1959) 34.6 0.0 0.1 2.5 42
A59-92 (1960) 35.3 0.1 0.2 0.1 18
A59-93 (1959) 30.7 0.1 0.04 10.0 21
A59-93 (1960) 35.4 0.1 0.2 0.1 26
A60-200 (1960) 34.3 0.1 0.7 0.4 13
A60-234 (1960) 36.0 0.0 0.0 0.0 5
A60-236 (1960) 35.6 0.0 0.2 0.0 5
A60-237 (1960) 34.9 0.0 0.4 0.4 9
A60-250 (1960) 35.7 0.03 0.1 0.03 29
A61-332 (1961) 35.2 0.0 0.4 0.0 5
A61-338 (1961) 30.8 0.0 0.8 7.5 8
A61-340 (1961) 34.6 0.0 0.2 2.0 5
A61-341 (1961) 35.6 0.0 0.1 0.3 8
A61-371 (1961) 35.8 0.1 0.0 0.2 17
B. NEW WORLD SPECIES

H. aculeatus
(59-4 & 59-7) 35.3 0.1 0.3 0.1 15
H. bifurcatus
A59-153 33.7 0.0 1.1 0.2 19
March, 1963] MENZEL AND WILSON-CYTOTAXONOMY OF HIBISCUS
TABLE 2-. Continued
Mean number per cell of
Species of hybrid II
H. [urcellaius
AS9-86 3S.9
AS9-87 33.S
A60-224 3S.3
A61-3S9 3S.6
H. bi/urcatus-/urcellatu(F1
AS9-86 X AS9-1S3 3S.2
AS9-87 X AS9-1S3 3S.7
• Menzel and Wilson, 1961b.
binus and H. sabdariffa were domesticated, prob-
ably in the western Sudan (Murdock, 1959).
Hibiscus acetosella also appears to be cultivated
(as a vegetable) throughout most of its range,
although it may be wild in Angola, where its
closest wild relatives occur (Exell, 1961). Several
other ruderal species, e.g., H. diversifolius, H.
asper, and perhaps H. meeusei, are occasionally
used for fiber; we are uncertain whether they are
ever cultivated. Hibiscus surattensis and H.
rostellatus are apparently truly wild. Hibiscus
radiatus, occasionally used for fiber, probably is
native to India and Burma and grows only as an
introduced species in Africa (Ulbrich, 1921).
Hibiscus acetosella and H. radiatus are allotetra-
ploids with a relatively low multivalent frequency
compared with that in autotetraploid H. canna-
binus and in the H. cannabinus-acetosella allohexa-
ploid. They have one genome, A, similar to that
of H. cannabinus and one dissimilar genome, B
(Menzel and Wilson, 1961b). The close relation-
ship between H. acetosella and H. radiatus re-
ported earlier, evidenced by the fertility of their
F, hybrids, has been confirmed by crosses between
a number of additional accessions of the 2 species
(unpublished data). Nevertheless, all the acces-
sions can be assigned without difficulty to one
or the other species. Hibiscus acetosella and H.
radiatus remain distinct despite the existence of
fertile, widely segregating experimental F? popu-
lations. Each may have had H. cannabinus or a
near relative of it as the A-genome ancestor and
a different diploid species as the B-genome an-
cestor. The dearth of known diploids makes it im-
possible to suggest what the latter species were;
they may be extinct.
Although H. diversifolius has twice as many
chromosomes as the tetraploid species, it had no
higher frequency of multivalents. Pairing varied in
different collections from the same accession. In
a sample of 19 pollen mother cells from A59-94
collected in January, 1962, no multivalents were
found and most cells had 72 bivalents (Fig. 5).
Other samples collected in May, June, and Octo-
ber, 1960, showed mean pairing of 68.7 bivalents,
269
No.
III IV I cells
0.0 0.0 0.3 34
0.0 0.0 1.3 4
0.0 0.3 0.0 6
0.0 0.2 0.0 S
0.1 0.3 0.3 22
0.0 0.1 0.1 49
0.1 trivalent, 1.5 quadrivalents, and 0.4 univalents
in 12 cells, with a maximum of 5 quadrivalents per
cell. Since the collections were not all from the
same plant, it is uncertain whether the variation
in pairing represented differences between indi-
viduals or between seasons. Data from A60-207
were too scanty for pairing analysis, but a few
quadrivalents were present in a sample of pollen
mother cells collected in January, 1962. Associa-
tions of more than 4 chromosomes were absent
in all samples. The prevalence of bivalent pairing
indicates that H. diversifolius is an allooctoploid
combining 4 dissimilar genomes. It follows that
at least 4 differentiated groups of genomes are
represented on the African continent alone.
Meiotic chromosome behavior of hybrids between
H. diversifolius and H. cannabinus, H. acetosella,
and H. radiatus (unpublished data) suggests that
the number is actually 5 or 6.
Genome relationships of diploid H. surattensis
and tetraploid H. rostellatus, H. sabdariffa, and
H. meeusei remain unknown, since attempts to
cross them with other species have been un-
successful.
Relationships of New World species--Section
Furcaria has a second important center of vari-
ation in tropical America. The 4 species available
for study all have forked bracteoles and form a
morphologically coherent group. The several other
species (Gurke, 1892; Hochreutiner, 1900), many
of them very imperfectly known to modern
botany, are mostly shrubs with simple bracts and
appear morphologically more closely allied to H.
diversifolius and to a group of Australian species
(series Furcaria diversifolia Ulbr.?) than to other
Afro-Asian taxa.
The Cuban species H. costatus is the only known
New World diploid. Most pollen mother cells of
A60-243 had 18 bivalents at metaphase I, but
1 plant also had 2-5 chromosome fragments in
some cells, and 2 other plants exhibited an oc-
casional quadrivalent. It is uncertain whether
these aberrations characterize a wild population
or arose during the long cultivation of this line
(Table 1).
270 AMERICAN JOURNAL OF BOTANY
The species H. furcellatus and H. bifurcatus are
closely related, distinct in our material but inter-
grading to some extent elsewhere (Hochreutiner,
1900, and see the following herbarium specimens
from Geneva: Gabriel 1802, Gardner 1243, Glaziou
17466, Spruce 1688). Hibiscus furcellatus has the
wider distribution, from southern Florida to Co-
lombia and in the Hawaiian Islands; H. bifurcatus
occurs from southern Mexico to Brazil. Roe (1961)
considered the Hawaiian plants specifically dis-
tinct (H. youngianus Gaud.). According to Roe's
criteria for distinguishing H. youngianus from
H. furcellatus, all our accessions, from Florida,
Colombia, and Hawaii, are H. youngianus. In our
opinion, Horhreutiner (1900) was correct in
considering H. youngianus a synonym of H.
furcellatus.
The dose relationship between H. furcellatus
and H. bifurcatus adduced on morphological
grounds was ronfirmed by the ease with which
crosses were made. F 1 hybrids of the crosses
A.59-153 X A.59-86, A59-153 X A59-87, and
their reciprocals, grown in 1961, were vigorous,
had a pubescent indument like that of H. fur-
cellatus, leaves of intermediate shape, and capsules
and seeds like those of H. bijurcoius. Analysis of
meiotic chromosomes (Table 2) showed that
chromosome pairing was complete and similar
to that in the parents. A few capsules containing
apparently viable seeds have been harvested.
Hibiscus furcellatus and H. bifurcatus evidently
have similar genome constitutions.
Of the species of section Furcaria, H. aculeatus
has the most northerly distribution: on the coastal
plain from North Carolina (Fox, Godfrey, and
Blomquist, 19.50) south to Florida and west to the
Sabine River in Louisiana. Common in northern
Florida in palmetto thickets, particularly at the
margins of ponds and ditches, it scarcely extends
into peninsular Florida. Efforts to establish roots
and rooted cuttings in a greenhouse in Tallahassee
and in field plantings in Palm Beach County were
unsuccessful. Large quantities of fresh seeds rol-
lected from several wild Florida populations in
1959 failed to germinate. Rootstocks collected
by R. K. Godfrey in burned-over woods in
Wakulla County, Florida, in May, 1959, and
transplanted to the garden in Tallahassee have
persisted for 4 seasons, bloomed, and produced
a few seeds.
Hibiscus aculeatus appears relatively uniform
over its range, which is separated by several
hundred miles from the range of any other species
of section Furcaria. It differs from the other
species of the section studied in being strictly a
summer (long-day") plant. In northern Florida,
new shoots appear in April and May, flowers begin
to open in June, and capsules begin to ripen about
mid-August. In favorable seasons new growth and
more flowers may appear in September but the
canes die back and disappear by November,
usually well before first frost. In this respect H.
[Vol. 50
aculeatus resembles neighboring species of Hibiscus
section Trionum more than it does other fur-
carias. In our experience the latter either bloom
sporadically throughout the year with a principal
flowering season in winter, or bloom strictly
during fall and winter under short-day conditions.
The presence in the New World of 3 tetraploid
species with low multivalent frequency indicates
that at least 2 genome groups are represented.
Whether either genome group is also present in
Africa is unknown, since crosses between Old
and New World species thus far have been un-
successful.
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DEVELOPMENT AND NUTRITION OF NEW SPECIES

OF THRAUSTOCHYTRIUMI
SOLOMON GOLDSTE"IN
Department of Biology, Brooklyn College, Brooklyn, New York

ABSTRACT
GOLDSTEIN, SOLOMON. (Brooklyn Coll., Brooklyn, N.Y.) Development and nutrition of new
species of Thraustochvtrium. Amer. Jcur. Bot. 50(3): 271-27!l.Illus. 1963.-The life cycles of 2
lower Phycomycetes in sea-water-pollen culture arc described and compared with variations ob-
served in nutrient media. Some ecological implications are discussed. Both fungi are obligately
marine and stenohaline. Maximal growth occurred with 2.5 and 3.0% NaCl not replaceable by
KCI. However, omission of KCl from media resulted in reduced yields. Salinity optima were not
temperature-dependent and the organisms grew best at 12-25 C. Neither organism assimilated
nitrogen in the form of KN0 3 or (NH4),S04. Optimal harvests were obtained with glutamate as
the source of nitrogen and with carbon supplied as glucose, maltose, soluble starch, or cellobiose.

PHYCOMYCETES remain a poorly known segment
of the ocean's flora although their prevalence in
polluted littoral waters (Vishniac, 1956) and at-
tacks upon phytoplankton are well established
(Johnson and Sparrow, 1961). Investigators of
marine Phycomycetes confront "a very real
problem in endeavoring, first, to find sporangia
which have not already discharged their zoospores
in the interval between collection and examina-
tion, and secondly to provide conditions in the
laboratory favorable for the development and
discharge of these sporangia" (Sparrow, 1934).
Accordingly, the establishment of axenic cultures
has been impeded, thus restricting studies of their
morphogenetic and metabolic potentialities. The
tedious procedures required for the detection of
these bizarre forms by classical means and the
If Received for publication September 6, 1962.
This study was supported by a research grant, E1179
from the National Institute of Allergy and Infectious Dis-
eases, National Institutes of Health, Public Health
Service and by the City University of New York. The
major part of the work was done during my tenure as a
Research Associate in the Department of Microbiology,
Yale University. I wish to thank Dr. Helen S. Vishniae
for the axenic cultures, Miss Jennie Pumphrey for the
excellent technical assistance, and Prof. W. H. Stahl of
the Department of Classics and Comparative Literature,
Brooklyn College, for the Latin diagnoses. •. _ ~
fortuitous nature of the results limited our knowl-
edge of their diversity and distribution. More
recently, methods utilizing antibiotics as ad-
juvants to plating techniques have been described
for the isolation of marine Thallophytes (Pro-
vasoli, Pintner, and Packer, 1951; Vishniac,
1955a, b ; Watson and Ordal, 1957). The applica-
tion of these simple microbiological measures
heralds a systematic approach to the investigation
of the marine Mycota.
In her exploration of fungus ecology, Vishniac
(1956) employed semiquantitative plating pro-
cedures that markedly decreased the role of chance
in the collection of evanescent organisms. She
demonstrated that polluted coastal waters sustain
an unexpectedly large population of non-fila-
mentous Phycomycetes that may exist as free-
living saprophytes competitive with marine bac-
teria. These fungi constitute a well-defined
ecological group specifically adapted to growth
in the ocean (Vishniac, 1960) and some are useful
as assay organisms for cobalamin and thiamine
in sea water (Adair and Vishniac, 1958; Vishniac,
1961; Vishniac and Riley, 1961). Except for
these investigations involving Thraustochutrium.
globosum and unidentified forms, the literature is
barren of experimental studies of chytridiaceous
marine fungi in axenic culture. '