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Genet Resour Crop Evol (2011) 58:831–835
DOI 10.1007/s10722-010-9620-2
Manihot fortalezensis Nassar, Ribeiro, Bomfim et Gomes
a new species of Manihot from Ceará, Brazil
Nagib M. A. Nassar • Dalva G. Ribeiro •
Nayra N. Bomfim • Pollyanna T. C. Gomes
Received: 11 May 2010 / Accepted: 13 September 2010 / Published online: 17 October 2010
Ó Springer Science+Business Media B.V. 2010
Abstract A new species of Manihot has been
collected from Fortaleza, Ceará state, Brazil. It was
grown at the living collection in the experimental
station, University of Brasilia since 1980 and left for
natural reproduction. It bred true during all these
years giving rise to identical typical plants of the
ancestor one. The closest species to it is M. glaziovii.
Apparently this new species was formed by sporadic
hybridization between M. glaziovii and cassava itself
in its natural habitat, Ceará. It possesses morphological marker gene of the latter one which is ribbed fruit
that is found only in cassava. Interspecific hybrids
produced experimentally between M. glaziovii and
cassava showed similar morphological markers.
Exam of its ovules revealed formation of multiple
embryos, an indicator of apomixis. Grafting it with
common cassava was compatible, giving increased
root size up to seven times.
Keywords Apomixis Caatinga vegetation Grafting Interspecific hybridization Manihot
glaziovii Multiembryonic ovule
N. M. A. Nassar (&) D. G. Ribeiro N. N. Bomfim P. T. C. Gomes
Universidade de Brası́lia, Brası́lia, Brazil
e-mail: [email protected]
The genus Manihot was initially authored by Adanson
(1763). Miller in the fourth edition of the Gardner
Gardner0 s dictionary in 1974, provided the first valid
post-Linean description of the genus Manihot (Rogers
and Appan 1973). Pohl (1827) in Plantarum Brasiliae
Icones et Discriptions provided the first monograph
study of the genus Manihot. Miller and Webster
(1962) revised it focusing on its ressemblance with
the genus Cnidoscolus because both have a single
floral envelope, and share a base chromosome number
(x = 9). The last full monographic description was
presented in 1973 by Rogers and Appan. They
established 98 species, and separated from it
Manihotoides giving it status of a second genus.
Nassar added in 1986 the species M. neusana
(Nassar 1986).
Wild Manihot species are precursors of cassava, an
important food staple crop for more than 800 million
people in tropics and subtropics. They have received
attention from research institutions in the decade
1970s as a source of useful genes for improving the
crop. It is aspirated that this may contribute to increase
world production of food and eradicate hunger in poor
countries. For this reason, an extensive living collected was grown and maintained at the University of
Brasilia, with the help of IDRC, since the decade
1970s up to this date (Nassar 1978a, 1999, 2003).
One of these grown introductions is the subject of
this article. Its seeds were collected in 1976 from
plants growing on the road between Fortaleza and
Itapipoca, Km 32, in the State of Ceará, Northeast of
The referred seeds were propagated at Experimental Station of Biology at University of Brasilia.
These plants were examined morphologically, anatomically and embriologically. Stalks of both
Manihot glaziovii and the new introduction were
grafted to cassava stalks, and then planted in
October 2006.
Latin diagnosis
Manihot fortalezensis a M. glaziovii Muell. habitu
gracili, caule 10–15 cm diameter. (versus caule
20–30 cm diameter), folio obovato, 10–12 cm lato
(versus folio ovato, 10 cm lato), lobo mediano 15 cm
longo (versus 25 cm longo), fructu coerulescenti
(versus virescenti), leviter costato (versus fructu haud
costato), sacco embryonali singuli embryone munito
(versus multis embryonibus munito) abunde differt.
Holotype 109906 Herbario UnB, Brasilia Brasil,
Ceará, 32 km from Fortaleza to Itapipoca, collected
by N. Nassar.
Description of the new species; Manihot
fortalezensis Nassar, Ribeiro, Bomfim et Gomes
Shrub ca 4 m high. Trunk at the base 10–15 cm in
diameter. Stem slender weak, branches by 3 m
height. Young and mature stems glabrous. Leaves
alternate, stipules caduceous, petioles ca 35–40 cm
long, glabrous. Petiole attachment to lamina slightly
peltate, width between basal edge of lamina and
petiole junction 0.5–1.0 cm. Leaves coriaceous.
Abaxial surface white greenish. Venation camptodromous, veins glabrous, palmately 5 lobed. Lobes
obovate with acute apex. Median lobes ca 15 cm
long, 12 cm width. Apex extremely acute never
broad. Base of lobes 3–3.5 cm wide. Lamina at sinus
never overlapping.
Inflorescence monoecious, terminal, branched ca
20 cm long, glabrous reddish bloom. Pistillate flowers
restricted to upper ‘ of the inflorescence. Pedicels ca
0.1 cm long. Flower tepal 1.5 cm long, reddish.
Genet Resour Crop Evol (2011) 58:831–835
Bracteoles setaceous ca 0.5 cm long. Margin slightly
Disk prominent reddish, ovario subglobose. Staminate buds obvoid, flower tepal 1 cm long.
Disk prominent reddish. Stamens 10 into whorls 5
each. Capsules 2.5 cm diameter slightly winged.
Seed 1–1/2 cm long caruncule prominent.
Anatomical sections of primary root showed
presence of sub epidermal layer. Exame of ovules
revealed presence of multiembryonic sacs in about
28% of ovules (Fig. 1).
Holotype 109906 Herbario UnB, Brasilia Brasil,
Ceará, 32 km from Fortaleza to Itapipoca, collected
by N. Nassar.
Results and discussion
Screening results
The collected seeds were propagated every year
since 1977 giving further generations. The offspring
bred true without segregation during the whole
subsequent 30 years. This triggered our attention to
examine it for apomixis. Results of investigation
were positive, and more than 28% of examined
ovules showed polyembryonic confirming apomictic
nature (Fig. 1).
Apparently apomixis is responsible to perpetuate
this type and evolving this new species in its
natural habitat. It seems that apomixis has played
Fig. 1 Manihot fortalezensis. Multiembryonic sac
Genet Resour Crop Evol (2011) 58:831–835
Fig. 2 a Stem of M. fortalezensis, b M. glaziovii, c M. esculenta
Fig. 3 At left, leaf of M. esculenta. In the middle, leaf of M.
fortalezensis. At right, leaf of M. glaziovii
Fig. 4 At left, fruit of cassava. In the middle, fruit of Manihot
fortalezensis. At right, fruit of M. glaziovii
an important role in evolution of species of this
genus. Nassar (1985) has reported the new species
Manihot neusana which showed later to be
highly apomictic. In the decade 2000 the same
author (Nassar 2006) reported evolving of new
species by chromosome doubling accompanied by
Grafting stalks of the new material native to
Fortaleza with cassava was compatible far more than
grafts made between cassava and original Manihot
glaziovii Muell. It was success in 100% out of 200
grafts, while in case of original M. glaziovii Muell.
percent of success was 30% within the same number
of grafts made. Successful grafts of the new material
grew vigorously too. This new approach of examining relationship between species by grafting proved
successful. In addition, it showed very much promising for improving cassava production nationally
Genet Resour Crop Evol (2011) 58:831–835
since it enlarged root size up to sevenfolds. We
designed a project to introduce it to small farmers to
enable them adopt technique of grafting and use it in
their properties.
Grafting it with cassava was highly compatible,
giving enlarged roots up to seven fold. When
propagated by cuttings it germinates easily contrary
to M. glaziovii Muell.
Interspecific hybrids between cassava and M.
glaziovii were produced experimentally and compared morphologically with this new material. It was
highly similar, having stem slender 3–3.5 cm diameter (Fig. 2), median lobe 15 cm long, obovate
10–12 cm width (Fig. 3), fruit exhibited the morphological marker of ribbed fruit (Fig. 4).
The features found in M. fortalezensis are intermediate between M. glaziovii and M. esculenta, as noted
in the Table 1. Therefore the new material collected
from Fortaleza, Ceará is defined as a new species and
called Manihot fortalezensis Nassar.
Ecology and distribution
The region where the seeds of Manihot fortalezensis
Nassar were collected is characteristic of caatinga
vegetation composed principally from dry scrubs,
latitude 4.8, occurring along road side. It flourishes in
limestones soil and in dry vegetation.
Morphological relationships
The new species, M. fortalezensis Nassar, is distinguished from M. esculenta Nassar and its closest
relative M. glaziovii Muell. according to the table
Position in section Glaziovianae
The modified Key to the species of sect. Glaziovianae
according to Manihot sect. Glaziovianae Pax emend.
Rogers and Appan is as follows:
1. Inflorescence a panicle.
2. Bracts and bracteoles setaceous, less than 0,25 cm wide.
3. Petiole attachment peltate.
4. Inflorescence usually more than 10.0 cm long, many flowered.
5. Leaves usually 3 lobed; median lobes obovate, usually more than 3.0 cm wide less than 7.0 cm;
lobes usually overlapping at the sinus; basal lobes recurved
M. glaziovii Muell.- Arg.
5. Leaves usually 5 lobed; median lobes very obovate with acute apex, more than 10.0 cm; lobes
not overlapping at the sinus; basal lobes, basal lobes curved up.
M. fortalezensis Nassar
4. Inflorescence less than 10.0 cm long, few flowered; leaves 5 lobed; median lobes oblong usually
less than 3.0 cm wide; lobes separated at sinus; basal lobes straight. M. pseudoglaziovii Pax et K. Hoffmann
3. Petiole attachment basal.
6. Inflorescence totally glabrous (with the rare exception of the interior surface of tepal)
M. epruinosa Pax et K. Hoffmann
6. Peduncles, pedicels, bracteoles, bractlets and filaments pubescent.
M. brachyandra Pax et K. Hoffmann
2. Bracts and bracteoles foliaceous, more than 0.5 cm wide.
M. maracasensis Ule
1. Inflorescence a raceme.
7. Staminate buds ovoid-ellipsoid; leaves usually 3 lobed; lobes never pandurate.
M. catingae Ule
7. Staminate buds conical; leaves 5 lobed; lobes frequently pandurate.
M.dichotoma Ule
Genet Resour Crop Evol (2011) 58:831–835
Table 1 Differences between the new species, and its closest relative M. glaziovii Muell.
M. glaziovii
M. esculenta
M. fortalezensis
Erect 20–30 cm diameter
Slender 1–1.5 cm
Slender 3–3.5 cm
Leaf node
Slightly prominent
Very prominent
Median lobe 25 cm long
Ovate 10 cm width
Usually 14–17 cm
2.6–5 cm
Median lobe 15 cm
Obovate 10–12 cm
Fruit hybrid
Globose, greenish
Embryo sac
One sexual embryo
Grafting with cassava
Weak compatibility
Cuttings germinate
Very difficult, almost impossible
Acknowledgments This work was carried out with the
assistance of the Conselho Nacional de Desenvolvimento
Cientifico—CNPq, Brasilia, Brazil. The new material and the
above-mentioned living collection were established at the
University of Brasilia with support from the Canadian
International Development Research Centre, Ottawa in the
1970s. Thanks are due to Tarciso filgueiras for latin diagnosis
and M. Nayar, and M. Jain for reviewing the manuscript.
Adanson M (1763) Families dês Plantaes II, Partie Paris, p 356
Cruz ND (1967) Nova espécie do gênero Manihot Adans. do
Estado de Minas Gerais. Bragantia 26 (n.unico)
Gratapaglia DE, Nassar NMA, Dianese JC (1986) Biossistemática de espécies brasileiras do gênero Manihot
baseada em padrões de proteı́na da semente. Cienc. Cult.
Miller KI, Webster GI (1962) Systematic position of Cnidoscolus and Jatropha. Brittonia 14:174–180
Nassar NMA (1978a) Conservation of the genetic resouces of
cassava (Manihot esculenta): Determination of wild
Species localites with emphasis on probable origin. Econ
Bot 32:311–320
Bluish, slightly
One sexual embryo
Multiembryo sac
High compatibility
Easily germinate
Easily germinate
Nassar NMA (1978b) Some further species of Manihot with
potential value to cassava breeding. Can. J. Pl. Sci.
Nassar NMA (1978c) Wild Manihot species of Central Brazil
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native to Paraná, Brazil. Can. J. Plant Sci. 65:1097–1100
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native to Brazil and its potential for cassava improvement.
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