InTech-Amaranthaceae As A Bioindicator of Neotropical Savannah Diversity

Chapter 10
Amaranthaceae as a Bioindicator of
Neotropical Savannah Diversity
Suzane M. Fank-de-Carvalho, Sônia N. Báo and Maria Salete Marchioretto
Additional information is available at the end of the chapter
http://dx.doi.org/10.5772/48455
1. Introduction
Brazil is the first in a ranking of 17 countries in megadiversity of plants, having 17,630
endemic species among a total of 31,162 Angiosperm species [1], distributed in five Biomes.
One of them is the Cerrado, which is recognized as a World Priority Hotspot for
Conservation because it has around 4,400 endemic plants – almost 50% of the total number
of species – and consists largely of savannah, woodland/savannah and dry forest ecosystems
[2,3]. It is estimated that Brazil has over 60,000 plant species and, due to the climate and
other environmental conditions, some tropical representatives of families which also occur
in the temperate zone are very different in appearance [4].
The Cerrado Biome is a tropical ecosystem that occupies about 2 million km² (from 3-24° Lat. S
and from 41-43° Long. W), located mainly on the central Brazilian Plateau, which has a hot,
semi-humid and markedly seasonal climate, varying from a dry winter season (from April to
September) to a rainy summer (from October to March) [5-7]. The variety of landscape – from
tall savannah woodland to low open grassland with no woody plants - supports the richest
flora among the world’s savannahs (more than 7,000 native species of vascular plants) and a
high degree of endemism [6,8]. This Biome is the most extensive savannah region in South
America (the Neotropical Savannah) and it includes a mosaic of vegetation types, varying
from a closed canopy forest (“cerradão”) to areas with few grasses and more scrub and trees
(“cerrado sensu stricto”), grassland with scattered scrub and few trees (“campo sujo”) and
grassland with little scrub and no trees (“campo limpo”) [3,9]. Among the grassland areas
there are some flat areas with rocky soil, called “campos rochosos”, which are considered
Cerrado areas because of their flora, especially when located in Chapada Diamantina (Bahia
State), a transition area between Cerrado and Caatinga Biomes.
Although the Cerrado is considered a Hotspot for the conservation of global biodiversity,
with plant species completely adapted to survive adverse conditions of soil and climate,
© 2012 Fank-de-Carvalho et al., licensee InTech. This is an open access chapter distributed under the terms
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236 Biodiversity Enrichment in a Diverse World
only 30% of this Neotropical Savannah biodiversity is reasonably well known [8,10].
Coutinho [11] believes that the frequent occurrence of fire is one of the most important
factors to determine this Biome´s vegetation, acting as a renewal element that selects
structural and physiological characteristics. Nowadays it is believed that more than 40% of
the original vegetation has already been converted into human-disturbed areas, due to the
expansion of crops [12,13]. This process has accelerated the fragmentation of natural habitat,
increasing the pressure on local biodiversity extinction and introducing exotic species, also
amplifying soil erosion, water pollution and alterations in vegetation and hydrologic
conditions [2,8,14].
The Amaranthaceae family is composed of 2,360 species and here will be listed those that
occur in Brazil, emphasizing the Cerrado species and including information on endemism,
endangerment and economic or potential use. We also provide a list of the most important
bibliographical references for those who are interested in studying the species of this
family. Some aspects of morphology, leaf anatomy and ultrastructure will be shown for
six species found in the Neotropical Savannah core area (Chapada dos Veadeiros) and
some of these aspects, as well as taxonomy and ecology, will be discussed in order to
propose the use of this plant family as an indicator of the diversity in open areas of this
Biome.
2. Methodology
2.1. List of the Brazilian Amaranthaceae species and those in RPPN Cara Preta
The Brazilian Amaranthaceae list (Table 1) was based on the research by the Brazilian
taxonomists Marchioretto [15-21] and Siqueira [22-25] and on the most important taxonomic
references to this Family both from the literature (Table 2) and Brazilian Herbaria (Table 3).
All cited Herbaria are listed according to the Index Herbariorum [26,27].
The species to be detailed were collected in a Conservation Unit named Reserva Particular
do Patrimônio Natural Cara Preta (RPPN Cara Preta), located in Alto Paraíso, Goiás State,
Brazil. After obtaining authorization from the NGO Oca Brasil, random walks were done in
order to locate, photograph and mark species with a Global Positioning System device, and
to collect and make exsiccates for Herbaria deposits, from September 2006 until March 2009.
Although some plant leaves were collected during the vegetative stage, these specimens
were visited until flowering to identify them correctly. All exsiccates were deposited in
Brazilian Herbaria as standard control material (prioritizing PACA, UnB and IBGE
Herbaria) and these species are included in Table 1.
2.2. Leaf anatomy and ultrastructure

Completely expanded leaves, from 3rd to 5th node from the apex, of two to six specimens of
each species were collected and sectioned. Part of the leaf medial region was fixed in
ethanol, acetic acid and formaldehyde [28] for 24 hours and preserved in ethanol 70% until
analysis to describe the anatomy and identify starch and crystal composition [28].
Amaranthaceae as a Bioindicator of Neotropical Savannah Diversity 237
Some pieces of the leaf medial region were immediately submerged in a Karnovsky solution
[29] of glutaraldehyde 2%, paraformaldehyde 2% and sucrose 3% in sodium cacodylate 0.05
M buffer for 12 to 24 hours and preserved in sodium cacodylate 0.05 M until processing for
analysis under an electron microscope. For the latter analysis, these pieces were post-fixed
in 2% osmium tetroxide and 1.6% potassium ferricyanide (1:1 v/v), followed by in-block
staining with 0.5% uranyl acetate solution (overnight). These samples were then dehydrated
in an acetone ascending series and slowly embedded in Spurr´s epoxy resin. Semi-thin and
ultra-thin sections were obtained in ultramicrotome with glass and diamond knives. Semi-
thin sections were stained with toluidine blue and analysed under the optical Zeiss
Axiophot, and ultra-thin sections were analysed under the transmission electron microscope
TEM JEOL JEM 1011.
3. Results and discussion

The Amaranthaceae family is composed of 2,360 species and 146 of them are found in Brazil
(Table 1). Ninety-eight species within the family are found in the Cerrado and 73 spp. are
endemic to Brazil, of which 13 are endemic to the Cerrado Biome (Table 1). Twenty
Amaranthaceae species are exclusive to the Cerrado (Table 1).
At least 22 Amaranthaceae species are referred to as being used in folk medicine (Table 1).
In Brazil, only two of these species are already used as commercial drugs, as capsules
containing their powdered roots, with studies to support their medicinal activity: Hebanthe
eriantha (Poir.) Pedersen and Pfaffia glomerata (Spreng.) Pedersen (Table 1), both known as
“Brazilian-ginseng”. However, there is neither registered success in isolating or synthesizing
their components nor any economic studies about the viability of this kind of
pharmaceutical procedure.
Although the species Gomphrena macrocephala St.-Hil. is not cited as medicinal (Table 1), the
fructan content in its roots has been determined [30] because this species was considered
synonymous with G. officinalis Mart. [31]. Later, it was determined that G. officinalis was
synonymous with G. arborescens L.f. and not with G. macrocephala [22]. Studying G.
arborescens, fructan was also determined as the principal carbohydrate in its subterranean
system [32]. This species is used in popular medicine to heal respiratory diseases (asthma
and bronchitis), to reduce fever and as a tonic [33-35]. An in vivo study (in cats) with the use
of fructans isolated from Arctium lappa L. (Asteraceae) reported a cough-suppressing activity
[36], and the presence of fructan in G. arborescens roots can partially justify the use of this
species as a medicinal plant.
Most members of Brazilian Amaranthaceae are only known by taxonomists and 42 species
are in danger of extinction according to Brazilian regional lists; 14 of them are recognized as
endangered by the Brazilian Ministry of the Environment (MMA – “Ministério do Meio
Ambiente”) (Table 1). Most of the endangered species are classified according to the IUCN
Red List of vulnerability categories, some even with the same criteria, and there is a wide
range of research still to be done.
Species
Bioma and level of Habit, popular name and species
Species Threat
endemism knowledge
Level
Herb; plant used as indigenous
Achyranthes aspera L. Cerrado medicine in Ethiopia with chemistry
study [37]
Achyranthes indica (L.) Mill. Cerrado Herb
Alternanthera adscendens
Cerrado exclusive Shrub
Suess.
Alternanthera albida (Moq.) Subshrub; C4 photosynthesis
Griseb. physiology [38]
Alternanthera aquatica
Herb
(D.Parodi) Chodat
Alternanthera bahiensis Cerrado, endemic to
Herb or subshrub
Pedersen Brazil
Herb; popularly named "anador"; folk
Alternanthera bettzichiana
medicinal plant, used as analgesic and
(Regel) G.Nicholson
antipyretic [39]
Herb; called "perpétua-do-mato,
periquito-gigante, penicilina" or
Brazilian joyweed; folk medicinal
Alternanthera brasiliana (L.) Cerrado, endemic to
plant, used as diuretic, digestive,
Kuntze Brazil
depurative, bequic, astringent and
antidiarrhoeal; ornamental plant; C3
photosynthesis structure [40-42]
Brazilian Cerrado
Alternanthera decurrens J. C.
endemic (Januária - CR [43] Subshrub
Siqueira
MG)
Alternanthera dendrotricha Cerrado, endemic to
Shrub
C.C.Towns. Brazil
Alternanthera flavida Suess. Subshrub
Alternanthera hirtula (Mart.)
EN [44] Herb
R.E.Fr.
Brazilian Cerrado
Alternanthera januariensi J. C.
endemic (Januária - CR [43] Subshrub
Siqueira
MG)
Alternanthera kurtzii Schinz Herb
Alternanthera littoralis
Herb; called "periquito-da-praia" [45]
P.Beauv.
Alternanthera malmeana
EN [44] Herb
R.E.Fr.
Brazilian Cerrado
Alternanthera markgrafii endemic (Serra de
Herb
Suess. Grão Mogol - MG)
Alternanthera martii (Moq.) Cerrado, endemic to

Subshrub
R.E. Fries Brazil
Alternanthera micrantha
Endemic to Brazil VU [44] Herb; called "periquito-da-serra" [45]
R.E.Fr.
Alternanthera minutiflora
Endemic to Brazil Herb
Suess.
Alternanthera multicaulis
Endemic to Brazil Herb
Kuntze
Herb; called "periquito-roseta,
periquito"; C3-C4 intermediary
Alternanthera paronychioides VU
Cerrado photosynthesis structure; C4
A.St.-Hil. [44,46]
photosynthesis physiology;
ornamental plant [38,41,42,45,47]
Herb; called "perna-de-saracura,
Alternanthera philoxeroides
Cerrado carrapicho-de-brejo" and alligatorweed
(Mart.) Griseb.
[45]
Alternanthera pilosa Moq. Herb
Alternanthera praelonga
CR [44] Herb
A.St.-Hil.
Alternanthera puberula
Cerrado exclusive Herb
D.Dietr.
Alternanthera pulchella Herb; C4 photosynthesis physiology
Kunth [38]
Herb; called "erva-de-pinto"; folk
medicinal plant, used to treat syphilis
Alternanthera pungens Kunth Cerrado
and skin diseases; C4 photosynthesis
physiology [38,39]
Alternanthera ramosissima
Cerrado Herb
(Mart.) Chodat
Alternanthera regelii (Seub.) Cerrado exclusive,
Herb
Schinz endemic to Brazil
Herb; called "periquito-de-reineck"
Alternanthera reineckii Briq. Cerrado VU [44]
[45]
Alternanthera rufa (Mart.) Cerrado, endemic to
Herb
D.Dietr. Brazil
Alternanthera sessilis (L.)
Cerrado LC [48] Herb
R.Br.
Herb; called "apaga-fogo,
carrapichinho, corrente, folha-de-
papagaio, periquito, periquito-figueira,
perpétua-do-mato, sempre-viva"and
Alternanthera tenella Colla Cerrado VU [44] joyweed; folk medicinal plant, used as
diuretic; this species is naturally
infected by a potyvirus; C3-C4
photosynthesis physiology and
structure [38,39,40,45,47,49,50]
Alternanthera tetramera
Herb
R.E.Fr.
Herb; called "caruru"; folk medicinal
Amaranthus blitum L. plant, used to fight anemia; C4
photosynthesis physiology [38,39,51]
Herb; called "rabo-de-gato, cauda-de-
raposa, disciplina-de-freira, rabo-de-
raposa"; folk medicinal plant, used to
Amaranthus caudatus L. Cerrado treat pulmonary diseases and as
emoliente; C4 photosynthesis
physiology; ornamental plant
[33,38,39,41]
Herb; called "caruru-vermelho, veludo,
bredo-de-jardim, crista-de-galo"; folk
Amaranthus cruentus L. Cerrado medicinal plant, used as emollient and
laxative; C4 photosynthesis physiology
[33,38,39,51]
Herb; called "caruru-rasteiro"; C4
Amaranthus deflexus L.
photosynthesis physiology [38,51]
Herb; called "caruru, bredo" and
Amaranthus hybridus L. Cerrado smooth pigweed; C4 photosynthesis
physiology [38,51]
Amaranthus muricatus Herb;, C4 photosynthesis physiology
(Moq.) Hieron. [38]
Herb; C4 photosynthesis physiology
Amaranthus retroflexus L. Cerrado
[38]
Amaranthus rosengurtii
EN [44] Herb
Hunz.
Herb; called "caruru-bravo, caruru-de-
espinho, bredo-de-espinho, caruru-de-
porco”; folk medicinal plant, used to
Amaranthus spinosus L. Cerrado
combat eczema and as emollient,
laxative and antiblenorragic; C4
photosynthesis physiology [33,38,39]
Herb; called "caruru-bravo, caruru-
verdadeiro, cururu, caruru-de-
soldado, caruru-de-folha-miúda,
Amaranthus viridis L. Cerrado amaranto-verde"; folk medicinal plant,
used as emollient and diuretic
desobstruente; C4 photosynthesis
physiology [33,38-40,51]
Herb; called “capotiraguá"; folk
Blutaparon portulacoides medicinal plant, used to combat
Cerrado VU [44]
(A.St.-Hil.) Mears leukorrhea; C4 photosynthesis
physiology [38,39]
Blutaparon vermiculare (L.) Herb; C4 photosynthesis physiology
Cerrado
Mears [38]
Herb; called "celosia-branca, celósia-

plumosa, crista-de-galo, crista-de-galo-
plumosa, suspiro, veludo-branco"; folk
Celosia argentea L. Cerrado
medicinal plant, used to combat
diarrhea and as anthelmintic and
astringent; ornamental plant [39,41,52]
Celosia corymbifera Didr. Endemic to Brazil Subshrub
Herb, subshrub; called "bredo-do-
Celosia grandifolia Moq. EN [44]
mato" [45]
Subshrub; called "mofungo-rabudo"
Chamissoa acuminata Mart. VU [44]
[45]
Chamissoa altissima (Jacq.) Subshrub; called "mofungo-gigante"
Cerrado VU [44]
Kunth [45]
Chenopodium album L. Cerrado Herb
Herb; called "erva-de-santa-maria,
Chenopodium ambrosioides L. Cerrado
erva-santa, quenopódio" [40]
Chenopodium murale L. Cerrado Herb
Cyathula achyranthoides
Herb
(Kunth) Moq.
Cyathula prostrata Blume Cerrado Herb
Froelichia humboldtiana Herb; C4 photosynthesis physiology
Cerrado
(Roem. & Schult.) Seub. [38]
Froelichia interrupta (L.) Herb; C4 photosynthesis physiology
Moq. [38]
Froelichia procera (Seub.) Herb; called "ervaço"; C4
Cerrado
Pedersen photosynthesis physiology [38,41]
Froelichia sericea (Roem. &
Herb
Schult.) Moq.
Froelichia tomentosa (Mart.) Herb; C4 photosynthesis physiology
Cerrado
Moq. [38]
Braziliann Cerrado
Froelichiella grisea R.E. Fries endemic (Chapada VU [43] Herb; C3 photosynthesis structure [42]
dos Veadeiros - GO)
Cerrado, endemic to
Gomphrena agrestis Mart. EN [46] Herb
Brazil
Herb, subshrub; called "perpétua,
perpétua-do-campo, perpétua-do-
mato, paratudo-do-campo, paratudo-
erva, raiz-do-padre"; folk medicinal
plant, used as tonic, to reduce fever
Gomphrena arborescens L.f. Cerrado exclusive
and against respiratory deseases;
potential use as ornamentalplant; roots
are fructan-rich; C4 photosynthesis
physiology/structure
[32,35,38,40,42,49,53,54]
Subshrub; C4 photosynthesis
Gomphrena basilanata Suess. Endemic to Brazil
physiology [38]
Gomphrena celosoides Mart. Cerrado
physiology [38]
Gomphrena centrota E.Holzh. Endemic to Brazil VU [43]
physiology [38]
Brazilian Cerrado
Gomphrena chrestoides
endemic (Chapada VU [43] Subshrub
C.C.Towns.
Diamantina - BA)
Cerrado, endemic to
Gomphrena claussenii Moq. Subshrub
Brazil
Gomphrena debilis Mart. Endemic to Brazil
physiology [38]
Subshrub; folk medicinal plant, used
Cerrado, endemic to
Gomphrena demissa Mart. to combat the flu; C4 photosynthesis
Brazil
physiology [38,49]
Cerrado, endemic to Subshrub; C4 photosynthesis
Gomphrena desertorum Mart.
Brazil physiology [38]
Gomphrena duriuscula Moq. Endemic to Brazil EN [43]
physiology [38]
Cerrado, endemic to
Gomphrena elegans Mart. VU [46] Subshrub
Brazil
Gomphrena gardnerii Moq.
Subshrub; called "gonfrena, perpétua,
perpétua-roxa, sempre-viva, suspiro,
suspiro-roxo"; folk medicinal plant
Gomphrena globosa L. Cerrado
used to fight respiratory diseases; C4
photosynthesis physiology;
ornamental plant [38,40,45,55,56]
Subshrub; called "perpétua-gramínea";
Gomphrena graminea Moq. Cerrado VU [44]
C4 photosynthesis physiology [38,45]
Gomphrena hatschbachiana Cerrado, endemic to
VU [43] Subshrub
Pedersen Brazil
Brazilian Cerrado Subshrub; C3 photosynthesis
Gomphrena hermogenesii J.C.
endemic (Chapada physiology; C4 photosynthesis
Siqueira
dos Veadeiros - GO) structure [38,42]
Brazilian Cerrado
Gomphrena hillii Suess. endemic (Paraíso do
physiology [38]
Norte - TO)
Cerrado exclusive, Subshrub; C4 photosynthesis
Gomphrena incana Mart.
endemic to Brazil physiology [38]
Gomphrena lanigera Pohl ex Cerrado exclusive, Subshrub; C4 photosynthesis
Moq. endemic to Brazil physiology and structure [38,42]
Gomphrena leucocephala Subshrub; C4 photosynthesis
Endemic to Brazil
Mart. physiology [38]
Gomphrena macrocephala Cerrado exclusive, Subshrub; roots are fructan-rich; C4
A.St.-Hil. endemic to Brazil photosynthesis physiology [30,38]
Brazilian Cerrado
Gomphrena marginata Seub. endemic Subshrub
(Diamantina - MG)
Gomphrena matogrossensis Cerrado exclusive,
Subshrub
Suess. endemic to Brazil
Gomphrena microcephala
endemic to Brazil Subshrub
Moq.
Subshrub; called "erva-mole, erva-
Cerrado, endemic to rosa"; folk medicinal plant, used as
Gomphrena mollis Mart.
Brazil tonic and carminative [39]
Brazilian Cerrado
Gomphrena moquini Seub. endemic (Serra do Subshrub
Cipó - MG)
Cerrado, endemic to
Gomphrena nigricans Mart. VU [43] Subshrub
Brazil
Cerrado exclusive, Subshrub, C4 photosynthesis
Gomphrena paranensis R.E.Fr.
Subshrub; called "perpétua-
Gomphrena perennis L. VU [44] sempreviva"; C4 photosynthesis
physiology [38,45]
Subshrub; called "infalível, paratudo,
paratudinho, paratudo-amarelinho";
Gomphrena pohlii Moq. Cerrado exclusive roots are used in folk medicine against
respiratory deseases; C4 photosynthesis
physiology and structure [38,39,42,49]
Gomphrena prostrata Mart.
Brazil physiology and structure [38,42]
Gomphrena pulchella Mart. EN [44]
physiology [38]
Gomphrena pulvinata Suess. Endemic to Brazil
physiology [38]
Gomphrena regeliana Seub.
Gomphrena riparia Pedersen Endemic to Brazil CR [43]
physiology [38]
Gomphrena rudis Moq.
Cerrado, endemic to
Gomphrena rupestris Nees Subshrub
Brazil
Gomphrena scandens (R.E.Fr.)
endemic to Brazil VU [43] Subshrub
J.C.Siqueira
Gomphrena scapigera Mart.
Subshrub; called "perpétua-
Gomphrena schlechtendaliana
EN [44] schlechtendal"; C4 photosynthesis
Mart.
physiology [38,45]
Gomphrena sellowiana Mart. Endemic to Brazil VU [44] Subshrub

Gomphrena serturneroides Subshrub; C4 photosynthesis
Endemic to Brazil
Suess. physiology [38]
Cerrado, endemic to Subshrub; called "thoronoé"; folk
Gomphrena vaga Mart. VU [44]
Brazil medicinal plant, used as analgesic [57]
Subshrub; called "cangussú-branco,
Cerrado, endemic to vergateza"; folk medicinal plant,
Gomphrena virgata Mart.
Brazil antiletargic; C4 photosynthesis
physiology and structure [33,38,42]
Subshrub, shrub; called "corango-açu,
ginseng-brasileiro, picão-de-
tropeiro,solidonia, suma"; folk
EN
Hebanthe eriantha (Poir.) medicinal plant, used to combat colic
Cerrado [44],
Pedersen and enteritis; most of its chemical
VU [58]
constituents are known and roots of
this plant are already used by
pharmaceutical companies [40,59]
Hebanthe grandiflora (Hook.)
Cerrado Bush scandentia
Borsch & Pedersen
Hebanthe occidentallis
Cerrado Subshrub scandentia
(R.E.Fr.) Borsch & Pedersen
Subshrub scandentia; called "corango-
Hebanthe pulverulenta Mart. Cerrado VU [58]
veludo" [45]
Hebanthe reticulata (Seub.)
Subshrub, shrub scandentia
Borsch & Pedersen
Hebanthe spicata Mart. Shrub erect or scadentia
Herbstia brasiliana (Moq.)
EX [46] Subshrub
Sohmer
Iresine diffusa Humb. & Subshrub; called "bredinho-difuso"
Cerrado
Bonpl. ex Willd. [45]
Iresine poeppigiana Klotzsch Subshrub
Lecosia formicarum Pedersen Endemic to Brazil Subshrub
Lecosia oppositifolia Pedersen Endemic to Brazil CR [43] Herb or subshrub

Pedersenia argentata (Mart.)
Herb
Holub
Pfaffia acutifolia (Moq.)
Cerrado Herb or subshrub
O.Stützer
Brazilian Cerrado
Pfaffia aphylla Suess. endemic (Gouveia - Subshrub
MG)
Cerrado exclusive,
Pfaffia argyrea Pedersen VU [43] Herb or subshrub
endemic to Brazil
Brazilian Cerrado
Pfaffia cipoana Marchior. et
endemic (Itambé do Subshrub
al.
Mato Dentro - MG)
Pfaffia denudata (Moq.) Cerrado exclusive,

Herb, subshrub, shrub
Kuntze endemic to Brazil
Cerrado exclusive,
Pfaffia elata R.E.Fr. Subshrub
endemic to Brazil
Herb, subshrub; called "corango-
Pfaffia glabrata Mart. Cerrado exclusive
sempreviva" [45]
Herb, subshrub; called "anador,
canela-velha, ginseng-brasileiro,
finseng, páfia, paratudo, corango-
sempreviva"; folk medicinal plant,
most of its chemical constituents are
Pfaffia glomerata (Spreng.)
Cerrado VU [44] known and roots of this plant are
Pedersen
already used by pharmaceutical
companies; butanolic extract showed
antihyperglycemic potential in vivo; C3
photosynthesis physiology and
structure [38,40,42,45,60]
Herb, subshrub, called "corango-de-
Pfaffia gnaphaloides (L.f.)
Cerrado VU [44] seda", C3 photosynthesis physiology
Mart.
and structure [38,42,45]
Cerrado exclusive,
Pfaffia hirtula Mart. Herb, subshrub
endemic to Brazil
Herb, subshrub; called "marcela-
branca, marcela-do-campo, marcela-
Cerrado, endemic to
Pfaffia jubata Mart. do-cerrado"and kytertenim; roots are
Brazil
used in folk medicine against intestinal
problems [39,49]
Cerrado exclusive,
Pfaffia minarum Pedersen VU [43] Subshrub
endemic to Brazil
Brazilian Cerrado
Pfaffia rupestris Marchior. et
endemic (Rio Pardo Subshrub
al.
de Minas - MG)
Subshrub; nickel hyperaccumulator, it
Brazilian Cerrado
is one of the first species to recolonize
Pfaffia sarcophylla Pedersen endemic
the ground with high concentrations of
(Niquelândia - GO)
total Ni in the soil (>1%) [61]
Pfaffia sericantha (Mart.) Cerrado, endemic to
Pedersen Brazil
Pfaffia siqueiriana Marchior. Cerrado, endemic to
Subshrub
& Miotto Brazil
Pfaffia townsendii Pedersen VU [43]
Brazil physiology and structure [38,42]
Cerrado, endemic to
Pfaffia tuberculosa Pedersen Herb, subshrub
Brazil
Pfaffia tuberosa (Spreng.) Herb, subshrub; called "corango-de-
Cerrado
Hicken batata" [45]
Cerrado exclusive,
Pfaffia velutina Mart. Subshrub
endemic to Brazil
Pseudoplantago friesii Suess PE [44] Popular name is "caruru-açu" [45]

Cerrado exclusive,
Quaternella confusa Pedersen Shrub
endemic to Brazil
Quaternella ephedroides Cerrado, endemic to
Shrub
Pedersen Brazil
Quaternella glabratoides
Endemic to Brazil EN [44] Subshrub; called "corangão" [45]
(Suess.) Pedersen
Xerosiphon angustiflorus Cerrado, endemic to
Subshrub
(Mart.) Pedersen Brazil
Xerosiphon aphyllus (Pohl ex Cerrado, endemic to
Subshrub
Moq.) Pedersen Brazil
Notes: The species threat level category is the same as that used in the IUCN Red List: CR (critically endangered), EN
(endangered), EX (extinct), LC (Least Concern) and VU (vulnerable).
Table 1. Amaranthaceae species found in Brazil, identifying those endemics to Brazil and the ones
found in the Neotropical Savannah (Cerrado), level of threat, habit, popular name (mostly in
Portuguese) and some of the knowledge about the species.
3.1. Morphology, taxonomy challenge and species list of Brazilian

Amaranthaceae
Taxonomy is the science that aims to identify and characterize species. It includes the study
of the plant´s behaviour in nature and is based on plant morphology. The use of other data,
such as anatomy studies, genetic characters, ecology and geographic pattern, aims to
include and define affinities and parental relations among plant groups. Only by knowing
the species is it possible for Botany to contribute to other scientific areas, including to the
conservation of species in situ, not only of plants but also of animals.
It is not easy to correctly identify Brazilian Amaranthaceae species. Different species can be very
alike in habit and vegetative morphology. The correct identification depends almost exclusively
on some flower details, whose small dimensions make it especially difficult to work in the field,
demanding a highly specialized work, only partially carried out for this family (15-22).
Brazilian species of this family are predominantly herbs, shrubs or climbing plants. They can
be annual or perennial, with erect, prostrate, decumbent or scandent stem. In species from the
Neotropical Savannah or from rocky fields, the underground organ is thickened and
composed of roots and a xylopodium – a portion of the subterranean system which is
responsible for the re-sprouting after a fire or other environmental stress [62]. The leaf
arrangement can be opposite, alternate or with a basal aggregation of leaves. Leaves are
exstipulate, glabrous or pubescent, with entire lamina and margins. Inflorescences can be
cymoses, in spikes, in heads, corymboses or paniculates, axillary or axial. Flowers are bisexual
or monoecious and small. The perianth is undifferentiated, actinomorphic, with five distinct or
partially connated sepals. Flowers are associated with dry and papery bracts. Fruits are dry,
usually a single-seeded achene or capsules with few seeds [15-22]. A short list of the most
important Brazilian Herbaria to visit in order to study Amaranthaceae taxonomy is presented
on Table 2 and the literature used to identify the species of this family is presented in Table 3.
Index Herbarium Name Institution and municipality

UFBA/Campus de Ondina, Salvador, Bahia,
ALCB Herbário da Universidade Federal da Bahia
Brazil
Herbário da Universidade Federal de Minas
BHCB UFMG, Belo Horizonte, Minas Gerais, Brazil
Gerais
BOTU Herbário da Universidade Estadual Paulista UNESP, Botucatu, São Paulo, Brazil
Herbário da EMBRAPA Recursos Genéticos EMBRAPA/CENARGEN, Brasília, Distrito
CEN
e Biotecnologia Federal, Brazil
CEPEC Herbário do Centro de Pesquisas do Cacau CEPEC, Itabuna, Bahia, Brazil
Herbário da Universidade Federal de Juiz de
CESJ UFJF, Juiz de Fora, Minas Gerais, Brazil
Fora
Herbário do Centro de Pesquisas
CPAP CPAP, Corumbá, Mato Grosso do Sul, Brazil
Agropecuárias do Pantanal
ESA Herbário da Universidade de São Paulo ESALQ/USP, Piracicaba, São Paulo, Brazil
FEEMA/INEA, Rio de Janeiro, Rio de Janeiro,
GUA Herbário Alberto Castellanos
Brazil
Herbário da Universidade Federal do
HTO UFTO, Porto Nacional, Tocantins, Brazil
Tocantins
Herbário da Universidade Estadual de Feira
HUEFS UFES, Feira de Santana, Bahia, Brazil
de Santana
Herbário do Instituto Agronômico de
IAC IAC, Campinas, São Paulo, Brazil
Campinas
IBGE Herbário da Reserva Ecológica do IBGE IBGE/RECOR, Brasília, Distrito Federal, Brazil
Herbário da Universidade Federal da UFPB, Cidade Universitária, João Pessoa,
JPB
Paraíba Paraíba, Brazil
Prefeitura Municipal/SMA, Curitiba, Paraná,
MBM Herbário do Museu Botânico Municipal
Brazil
Instituto Anchietano de Pesquisas/UNISINOS,
PACA Herbarium Anchieta
São Leopoldo, Rio Grande do Sul, Brazil
Herbário do Jardim Botânico do Rio de
RB JBRJ, Rio de Janeiro, Rio de Janeiro, Brazil
Janeiro
Secretaria de Meio Ambiente, São Paulo, São
SP Herbário do Instituto de Botânica
Paulo, Brazil
SPF Herbário da Universidade de São Paulo USP, São Paulo, São Paulo, Brazil
UB Herbário da Universidade de Brasília UnB, Brasília, Distrito Federal, Brazil

Herbário da Universidade Estadual de
UEC UNICAMP, Campinas, São Paulo, Brazil
Campinas
UFG Herbário da Universidade Federal de Goiás UFG, Goiânia, Goiás, Brazil
VIC Herbário da Universidade Federal de Viçosa UFV, Viçosa, Minas Gerais, Brazil
Notes: Herbaria are cited according to the Index Herbariorum and all of them have good collections of Amaranthacae.
The Institution/city where the Herbaria are located is also referred.
Table 2. List of the most important Herbaria references for researchers interested in studying the
Brazilian Amaranthaceae
[15-21] Revisions of Brazilian Froelichia, Froelichiella, Hebanthe and Pfaffia; species list and
phytogeography
[22-25] Revision of Brazilian Gomphrena; species list na phytogeography
[45] Amaranthaceae from Santa Catarina State, Brazil
[63] Restoring the Hebanthe genera
[64,65] Brazilian Amaranthaceae species and the Family in the World
[66] Revision of Amaranthaceae in the World
[67-71] Studies in South American Amaranthaceae
[72] Amaranthaceae in Flora Brasiliensis
[73] Studies of Pfaffia and Alternanthera genera
[74,75] Amaranthaceae in Central and South America
[51,52,56,76] Amaranthaceae from Rio Grande do Sul State, Brazil
Note: These references are ordereb by author and should be consulted in order to identify Brazilian species correctly.
Table 3. List of the most important bibliographical references for researchers interested in studying the
Brazilian Amaranthaceae
The Reserva Particular do Patrimônio Natural (RPPN) Cara Preta, in Alto Paraíso, Goiás
State, is a good representative of Neotropical Savannah vegetation, at about 1,500 meters of
altitude and showing rocky slopes with Cerrado sensu stricto (Figure 1), grassland with
scattered scrubs and few trees and grassland with few scrubby plants and no trees (Figure
2). The Pfaffia genus was restricted to a rocky slope and the other species were found in a
level field of sandy soils, usually covered by Poaceae and Cyperaceae. It was very difficult
to find all the species. It was only possible because of frequent visits to RPPN Cara Preta,
using GPS to mark the local after finding any probable member of the family in order to be
able to accompany them until the flowering stage. The area was monitored for one and a
half year and only Gomphrena hermogenesii J.C. Siqueira and Pfaffia townsendii Pedersen
(Figure 3) were localized, the first one always in vegetative stage. A key event to help
finding all six species was a fire that burned out the vegetation in August of the year 2008:
without the competition of the grasses, the Amaranthaceae species regrew and flowered
rapidly, in order to spread their seeds before the grasses could fully recover (Figures 4-8).
Pfaffia townsendii is a shrub species with persistent aerial portions that flowers throughout
the year (Figure 3). The herb G. hermogenesii is endemic to Chapada dos Veadeiros and also
has permanent aerial portions (about 10-20 cm high), but it was commonly found in
vegetative stage under the grass leaves; its flowering stage was stimulated by fire (Figure
4). Froelichiella grisea R.E.Fr. (Figure 5), G. lanigera Pohl. ex Moq. (Figure 6), G. prostrata
Mart. (Figure 7) and P. gnaphaloides (L.f.) Mart. (Figure 8) species were recorded in the
flowering stage at RPPN Cara Preta around 20 days after a fire that burned out all the
vegetation in the area, which is evidence of the pirophytic behaviour of most Neotropical
Savannah Amaranthaceae. Five of these species had never been recorded in this RPPN
before and one of them was last recorded in 1966 (F. grisea), according to Herbaria data.
Figures 1-8 are reproduced [77] with the authorization of the Biota Neotropica Editor, Dr.
Carlos Joly.
Figure 1. (Figures 1-8) Photographs of the environment and of the studied species at Reserva Particular
do Patrimônio Natural (RPPN) Cara Preta, Alto Paraíso, Goiás State, Brazil. Fig. 1. Rocky slope where
were found the species Pfaffia townsendii Pedersen and P. gnaphaloides (L. f.) Mart. Fig. 2. Humid rocky
grassland where were found the species Froelichiella grisea R.E.Fr., Gomphrena hermogenesii J.C. Siqueira,
G. lanigera Pohl. ex Moq. and G. prostrata Mart. Fig. 3. P. townsendii. Fig. 4. G. hermogenesii. Fig. 5. F.
grisea. Fig. 6. G. lanigera. Fig. 7. G. prostrata. Fig. 8. P. gnaphaloides.
Five species are herb to subshrub, and only P. townsendii is a shrub (Figure 3). Well-
developed tuberous subterranean systems were found in F. grisea (Figure 5) and G.
hermogenesii, while in G. lanigera and P. gnaphaloides the underground organ was less
developed, also tuberous. G. prostrata and P. townsendii presented a well-developed and
lignified underground organ. Leaves of F. grisea and G. hermogenesii are opposite and
alternate in the other studied species. F. grisea and G. lanigera can present a basal
aggregation of leaves. Leaves are always tomentose, with exception of the adaxial face of F.
grisea, which can be glabrous. Inflorescence is axial in all these species, spikes in F. grisea and
G. lanigera and heads in the other studied species. Flowers are yellowish in F. grisea, G.
hermogenesii and G. lanigera, with a tendency to turn red in the first and last one. In G.
prostrata, P. gnaphaloides and P.townsendii flowers are white, turning beige in the last species.
All the species have flowers associated with dry and papery bracts that persist alongside
their dry fruits, usually a single-seeded achene, favouring anemocoric dispersion.
The fastest lifespan was observed in G. lanigera, which took around 20 days to regrowth and
finish the flowering phase. In Figure 6, G. lanigera was about 20 days old and fruits were
almost mature, indicating proximity to the seed dispersal phase. Pfaffia townsendii alone
showed behaviour that was independent of fire, since even G. hermogenesii only flowered
after being burned to the ground and regrowing from its xylopodium. The other four
species were found after the occurrence of fire, all of them in the flowering stage.
In the Taxonomy and Morphology areas, studies of the genera Achyranthes, Alternanthera,
Amaranthus, Blutaparon, Celosia, Chamissoa, Chenopodium, Cyathula, Iresine, Lecosia,
Pedersenia, Pseudoplantago, Quaternella and Xerosiphon still need to be done, not only
covering the revision of the Brazilian species, biogeography and morphological evolution,
but also molecular biology to establish synonyms and to delimit variations among
individuals of each species.
3.2. Leaf anatomy of Amaranthaceae species

Leaves of the six studied species have anatomical variation among the genera and are more
similar between species of the same genus. Transverse sections show that G. hermogenesii
(Figure 9), G. lanigera (Figure 10) and G. prostrata (Figure 11) have large nonglandular
trichomes covering the single layered epidermis, dorsiventral mesophyll with upper
palisade parenchyma and spongy parenchyma near the lower epidermis. All these three
species are amphistomatic, and a complete well-developed parenchymatous sheath with
thicker cell walls surrounds the vascular bundles (Kranz cells), in which starch accumulates.
Calcium oxalate druses were found in the mesophyll. The leaf anatomy of the three
Gomphrena spp. is compatible with the C4 photosynthesis pathway.
Pfaffia gnaphaloides (Figure 12) and P. townsendii (Figure 13) have more undulating
surfaces and a thinner leaf blade in relation to the Gomphrena species. Trichomes are also
more frequent and thinner and the mesophyll is dorsiventral. The parenchymatous
sheath has thinner walls than the neighbouring cells in Pfaffia species. Both species had
elevated stomata on the lower epidermis and only P. gnaphaloides had few stomata on the
upper epidermis. Starch was distributed in all mesophyll cells and calcium oxalate
druses were rare. The anatomy of Pfaffia spp. leaves is compatible with C3 photosynthesis
metabolism.
Froelichiella grisea (Figure 14) has the only isobilateral mesophyll among the studied
species, with palisade parenchyma near both upper and lower epidermis. Palisade cells
are shorter near the lower epidermis. The parenchymatic vascular bundle is not
conspicuous and organelles in these cells are positioned towards the outer cell walls, in
the same way as they are found in the other mesophyll cells. Calcium oxalate druses were
more common near the midrib, and the reaction to starch was similar to that of all the
mesophyll cells. Its leaf anatomy is compatible with C3 photosynthesis metabolism.
Figures 9-14 [77] were reproduced with the authorization of the Biota Neotropica Editor,
Dr. Carlos Joly.
Gomphrena trichomes are similar to the ones described for G. arborescens [32,54,78]. Although
it is expected that stomata are reduced on the upper surface of land plants, the Cerrado
Gomphrena species G. arborescens, G. pohlii and G. virgata have a similar number of stomata
on both surfaces [78], subjecting them to a greater water loss, which is compensated by the
well-developed subterranean systems that guarantee water supply during the lifespan of
their leaves. The size and number of stomata on both leaf surfaces of G. hermogenesii, G.
lanigera and G. prostrata is still to be verified, but simple observation indicates that it should
be similar to the phenomena observed in the first cited species, since they also have a
relatively well-developed subterranean system.
Figure 2. (Figures 9-14) Micrographies of the middle leaf transversal sections of the studied
Amaranthaceae species. Fig. 9. Gomphrena hermogenesii - leaf blade thickness from medium to thick,
dorsiventral mesophyll and complete parenchymatous bundle sheath, with thick cell walls and
collateral vascular bundles. Fig. 10. G. lanigera - medium leaf blade, dorsiventral mesophyll and
complete parenchymatous bundle sheath, with thick cell walls and collateral vascular bundles. Fig. 11.
G. prostrata – thin to medium leaf blade, dorsiventral mesophyll and complete parenchymatous bundle
sheath, with thick cell walls and collateral vascular bundles. Fig. 12. Pfaffia gnaphaloides – thin leaf blade,
dorsiventral mesophyll, less defined parenchymatous bundle sheath and collateral vascular bundles.
Fig. 13. P. townsendii – only hypostomatous leaf species, thin leaf blade, dorsiventral mesophyll, less
defined parenchymatous bundle sheath and collateral vascular bundles. Fig. 14. Froelichiella grisea -
thick leaf blade, isobilateral mesophyll with elongated palisade parenchyma under the adaxial
epidermis, less defined bundle sheath and collateral vascular bundles. Legend: eab = abaxial epidermis;
ead = adaxial epidermis; pl = spongy parenchyma; pp = palisade parenchyma; arrowhead = stoma;
circle = druse; arrow = parenchymatous bundle sheath. Bar = 100 µm.
The leaf anatomy of the three Gomphrena spp. is similar to that of G. arborescens L.f. [54], G.
cespitosa, G. dispersa, G. nitida, G. sonorae [79] and G. conica, G. flaccida [80] among others,
most of them arranged in a Gomphrena atriplicoid-type of Kranz anatomy [81,82]. As
expected, there was no significant variation in these species’ leaf anatomy due to the life
cycle stage, although older leaves collected during the vegetative stage have a thicker cuticle
covering both epidermis surfaces, especially in G. hermogenesii species. The leaf anatomy
observed in the two Pfaffia spp. is similar to that described in P. jubata [83], which also lacks
the Kranz anatomy. There is no previous study about the anatomy of F. grisea leaves and its
genus is monoespecif.
There are still a number of studies to be done in the field of anatomy and histology of
Brazilian Amaranthaceae plants. Most of the medicinal species need to be analyzed
and validated for their use as drugs, including anatomic description and an
investigation of the secondary compounds of the used organs, by histology and by
chromatography. Due to the difficulties in correctly identifying the species in the field,
anatomical and morphological markers should be defined to guarantee these species’
identity even during the vegetative stage. Besides that, anatomical studies can improve
the taxonomy data and explain some morphological characters of this plant family, like
the anatomical variations in the leaf that are connected to photosynthesis, or the
secondary thickening and xylopodium development in underground organs, which is a
character for the Cerrado species. The anatomy of few Brazilian Amaranthaceae species
has been described, with the exception of some from the Gomphrena and Pfaffia genera
[83-86].
3.3. Leaf ultrastructure of Amaranthaceae species

Leaves of the six studied species have less ultrastructural variation among the species of the
same genus. Froelichiella grisea organelles are equally distributed among chlorenchyma
tissues, usually near the cell walls. The chloroplasts of this species are always granal (Figure
15), even in the vascular cells, with large starch granules (usually one or two per organelle)
in all tissues. Plastoglobuli are small and less numerous in mesophyll chloroplasts (Figure
15), but guard cell chloroplasts usually have just one large plastoglobulus and less
conspicuous grana. Mitochondria and peroxisomes (Figure 15) were found in mesophyll
and bundle sheath cells. Leaf ultrastructure is compatible with C3 photosynthesis
metabolism.
Mesophyll cell chloroplasts of Gomphrena species have conspicuous grana, rare starch
granules and variable size of plastoglobuli: G. hermogenesii has larger ones in relation to G.
lanigera and G. prostrata. Bundle sheath chloroplasts are completely devoid of grana or
have few stacked thylakoids (Figure 16) in all studied Gomphrena species, but always have
large starch granules and plastoglobuli. The larger the starch granules, the more deformed
the chloroplasts’ typical lens shape, as shown in G. hermogenesii (Figure 16). Mitochondria
are usually numerous in bundle sheath cells and are always near chloroplasts, grouped
next to the inner cell wall (towards the vascular bundle). Peroxisomes are rare, and a few
were observed near chloroplasts in palisade and spongy parenchyma cells, but not in the
bundle sheath cells. Phloem companion cells are mitochondria-rich in all Gomphrena
species, as shown in G. prostrata (Figure 17). The presence of dimorphic chloroplasts,
disposition of the organelles and the occurrence of Kranz syndrome seen in the leaf
anatomy indicate that the C4 photosynthesis pathway operates in the three studied
Gomphrena spp.
Pfaffia species organelles are equally distributed among chlorenchyma tissues, usually near
the cell walls. Pfaffia chloroplasts are granal even in the vascular cells, showing large starch
granules and a similar size in all mesophyll cells, as can be observed in the palisade
parenchyma of P. townsendii (Figure 18). Mitochondria and peroxisomes are common near
chloroplasts (Figure 18). Phloem companion cells are mitochondria-rich and chloroplasts are
smaller and granal, as in the other species of this study. Along with the aspects of Pfaffia
anatomy described previously, their ultrastructure is compatible with the C3 photosynthesis
pathway.
Pfaffia gnaphaloides (Figure 19) and G. hermogenesii (Figure 20) leaves, collected during the
flowering stage, were colonized by two distinct forms of microorganisms: (i) a smaller
organism was found in the intercellular spaces (ics) of the spongy parenchyma (Figure 19);
(ii) a larger and distinctly eukaryotic organism was found within distinct cells, with some
morphological alterations suggesting an infectious process (Figures 19-20).
The external envelopae membranae system of the chloroplasts is disrupted in infected cells
(Figure 20) and a size reduction was observed in the chloroplast plastoglobuli. All
morphological characteristics observed in the intracellular microorganism suggest that it
should be an obligate biotroph endophytic fungus belonging to the Ascomycete division
(Figure 20). The invading fungus may be using the plastoglobuli lipids as its primary source
of carbon and energy; the reduction of the plastoglobuli could also be due to its mobilization
by the host plants in response to the stress caused by these biotic interactions. The complete
identification of the fungus and its effect on the plants depends on its isolation from the
environment/hosts and complementary studies.
The rare peroxisomes in Gomphrena spp. leaf cells and their presence among all
chlorenchyma tissues of the Pfaffia spp. leaf cells is compatible with their possible
photosynthesis metabolisms. Along with the presence of Kranz syndrome and dimorphic
chloroplasts, the absence of peroxisome indicates that Gomphrena spp. perform
photosynthesis via the C4 pathway. In Gomphrena species, CO2 concentration in the bundle
sheath cells must be efficient, leading to a significant reduction in the oxygenase function of
its RuBisCO enzyme. This leaves the species virtually free of the photorespiration process,
aided by the large walls of the bundle sheath cells. Although a carbon isotope ratio study
[38] indicates that G. hermogenesii is not a C4 species, this species also has Kranz anatomy
and ultrastructure compatible with C4 metabolism, as do all the other studied Gomphrena
spp. [42]. The distribution of its key photosynthetic enzymes will be carried out using
immuno-cytochemistry, in our laboratory, in order to complete these data.
Figure 3. (Figures 15-20) Citological aspects of Amaranthaceae species as seen through a Transmission
Electron Microscope. Fig.15. Froelichiella grisea palisade parenchyma cell. Fig. 16. Gomphrena hermogenesii
bundle sheath cell. Fig. 17. G. prostrata phloem companion cell and bundle sheath cell on top. Fig. 18.
Pfaffia townsendii palisade parenchyma cells.Fig. 19. P. gnaphaloides spongy parenchyma cells and
invading microorganisms (black arrows). Fig. 20. G. hermogenesii bundle sheath cell and the invading
Ascomycete fungus (black arrow) and the disrupted chloroplasts with smaller plastoglubuli. Legend:
black arrow = invading organism; white arrow = mitochondria; ellipsis = septum with a simple pore; bsc
= bundle sheath cell; cw = cell wall; ics = intercellular space; n = nucleus of the microorganism;
N = nucleus of the plant species; p = peroxisome; pc = palisade parenchyma cell; pg = plastoglobulus in
a chloroplast; s = starch granule in a chloroplast; sc = spongy parenchyma cell.
4. Conclusions and perspectives

This chapter presents data on Amaranthaceae species, with no pretension to explain the full
potential of this plant family for scientific studies, but rather to provide a basic tool for those
interested in amplifying studies on the species of this family. Based on our results, we are
convinced of the importance of studying this family further, not only as a tool in the better
preservation of endemic species, but also to explore its undoubted economic importance
more fully. Basic research is still needed, with the aim of applying knowledge on these
species to technological advances, especially in growing crops - since C4 species have a faster
metabolism and growing capacity, as observed in the species found in the RPPN Cara Preta
- and to explore medicinal molecules of these plants. C4 species are also important to balance
CO2 in the atmosphere because of their efficiency in the transformation of carbon into
biomass; in Cerrado Amaranthaceae species, this storage is basically underground in their

well-developed subterranean roots and xylopodium.
The number of medicinal plants among the Brazilian Amaranthaceae species may well be
higher than already reported (Table 1), because Cerrado inhabitants are particularly
interested in the highly developed subterranean systems of some medicinal species
[34,49,58] which can be collected at any time of the year, even from species whose aerial
portions are not persistent. Due to the morphological similarity among Amaranthaceae
species in the Neotropical Savannah, their collectors can easily mistake one species for
another during the vegetative stage, which confirms the need for further and more complete
studies of the known medicinal and endangered species, at least.
Preparation of plant samples for transmission electron microscopy also proved to be useful
in studying the morphology of fungi inside plant cells, as well as aspects of host-parasite
interaction. This kind of study could be recommended for plants considered toxic to
herbivores and to any medicinal plant consumed by humans, in order to give more
information about the real source of poisoning or medicinal effect and for fine quality
control. In both studied species (G. hermogenesii and P. gnaphaloides) the external macro
aspects of the plants did not indicate the presence of the endophytic fungus.
RPPN Cara Preta is a small Private Conservation Unit (only 1.5% of the area of the Chapada
dos Veadeiros National Park, a government-preserved area of 65,038 hectares). Both
Conservation Units are separated only by a road, in Alto Paraíso municipality of Goiás
State, Brazil. The latter site is registered by UNESCO as a natural protected Cerrado zone.
RPPN Cara Preta has 245 species representing 47 family plants [75,86], which is 9.2% of the
2,661 plant species of Chapada dos Veadeiros [87], a good diversity of plants in relation to
the occupied area. There are six Amaranthaceae species in the RPPN – 25% of the 27 species
found in the National Park [87]. Considering that the RPPN Cara Preta Utilization Planning
Report [86] indicated the presence of three endemic species, plus two Amaranthaceae
species not reported initially [75], this Conservation Unit has 2% of endemic species – which
is more than expected. According to [2,12], the Cerrado Biome is one of the priority hotspots
for conservation because it has, among others, 4,400 endemic plants (1.5% of the Earth’s
300,000 species). The Amaranthaceae family in RPPN Cara Preta can be considered a taxon
indicator of the good diversity of the Neotropical Savannah. This taxon could be considered
a plant diversity indicator in other works on flora in open areas of the Cerrado Biome.
Because of the predominant habit (herbs and shrubs) and survival strategies, the presence of
species from this family among the collected species clearly indicates a well performed
collection effort.
There are a number of important factors indicating that this plant family deserves more
studies for a greater understanding by researchers working in Brazil, and we recap them as
follows: the Cerrado Biome holds 98 of the 146 Brazilian Amaranthaceae species (almost
70% of the total species) (Table 1); their pirophytic behavior and survival strategies (fast
regrowth and seed dispersal before the complete recovery of grasses after fire) are coherent
with the Biome’s characteristics; their morphology shows exceptional adaptation to the
seasonal climate and open areas (hairy aerial portions, partial or total loss of the aerial
portions during the dry season, well-developed underground system with xylopodium, dry
fruit dispersal by wind); their metabolism (evolution of C4 and intermediary C3-C4
photosynthesis) may have importance for biomass conversion and CO2 balance; and, finally,
many of these plants are already used in medicines by Cerrado inhabitants and there may
be much wider medicinal potential in other species of this family.
Author details
Suzane M. Fank-de-Carvalho*
National Council for Scientific and Technological Development - CNPq, Brasília,
Distrito Federal, Brazil
Universidade de Brasília – UnB, Biological Institute, Electron Microscopy Laboratory, Brasília,
Distrito Federal, Brazil
Sônia N. Báo
Universidade de Brasília – UnB, Biological Institute, Electron Microscopy Laboratory,
Brasília, Distrito Federal, Brazil
Maria Salete Marchioretto

Instituto Anchietano de Pesquisas/UNISINOS, PACA Herbarium, São Leopoldo,
Rio Grande do Sul, Brazil
Acknowledgement
We would like to thank CAPES, CNPq and FINEP for financial support; NGO Oca Brasil
and Herbaria IBGE, UB and PACA for access authorization and research infrastructure; the
aditional collectors for help in searching for and collecting the species at RPPN Cara Preta;
and Susan Casement Moreira for the English review.
5. References
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Bicudo, C.; Zappi, D.; Costa, D.P.; Lleras, E.; Martinelli, G.; Lima, H.C.; Prado, J.;
Stehmann, J.R.; Baumgratz, J.F.A.; Pirani, J.R.; Sylvestre, L.; Maia, L.C., Lohmann, L.G.;
Queiroz, L.P.; Silveira, M.; Coelho, M.N.; Mamede, M.C.; Bastos, M.N.C.; Morim, M.P.;
Barbosa, M.R.; Menezes, M.; Hopkins, M.; Secco, R.; Cavalcanti, T.B. & Souza, V.C.
(2010). Catálogo de Plantas e Fungos do Brasil – Vol. 1. Rio de Janeiro: Andrea
Jakobsson Estúdio/Instituto de Pesquisa Jardim Botânico do Rio de Janeiro. 870 p.
[2] Myers, N.; Mittermeier, R.A.; Mittermeier, C.G.; Fonseca, G.A.B. & Kent, J. (2000).
Biodiversity hotspots for conservation priorities. Nature, 403: 853-858.
* Corresponding Author
[3] Conservation International (2012). Cerrado. In: Biodiversity Hotspots. Available:

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Chapter 10

Suzane M. Fank-de-Carvalho, Sônia N. Báo and Maria Salete Marchioretto

Additional information is available at the end of the chapter

2.2. Leaf anatomy and ultrastructure

3. Results and discussion

Alternanthera martii (Moq.) Cerrado, endemic to

Herb; called "celosia-branca, celósia-

Gomphrena sellowiana Mart. Endemic to Brazil VU [44] Subshrub

Lecosia formicarum Pedersen Endemic to Brazil Subshrub

Lecosia oppositifolia Pedersen Endemic to Brazil CR [43] Herb or subshrub

Pfaffia denudata (Moq.) Cerrado exclusive,

Pseudoplantago friesii Suess PE [44] Popular name is "caruru-açu" [45]

3.1. Morphology, taxonomy challenge and species list of Brazilian

Index Herbarium Name Institution and municipality

UB Herbário da Universidade de Brasília UnB, Brasília, Distrito Federal, Brazil

3.2. Leaf anatomy of Amaranthaceae species

3.3. Leaf ultrastructure of Amaranthaceae species

4. Conclusions and perspectives

biomass; in Cerrado Amaranthaceae species, this storage is basically underground in their

Maria Salete Marchioretto

[3] Conservation International (2012). Cerrado. In: Biodiversity Hotspots. Available:

[86] Handro, W. (1967). Contribuição ao estudo da venação e anatomia foliar das

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