KEW BULLETIN VOL.

65: 691Y709 (201 0)

Amazon vegetation: how much don_t we know and how much does
it matter?

William Milliken1, Daniela Zappi1, Denise Sasaki2, Mike Hopkins3 & R. Toby Pennington4

Summary. In spite of the existence of a vast body of information on the plant diversity of the Amazon, there
remain significant obstacles to informed decision-making and management for conservation. Species distributions
are poorly understood and the relationships between diversity and composition of vegetation, ecosystem services
and resilience to climatic fluctuations are insufficiently clear. The geographic distribution of phylogenetic diversity
in relation to current protected areas is unexplored and very little is known about intraspecific genetic variability
and its practical significance. Interpretation of vegetation differentiation and distribution remains relatively sim-
plistic; there are still large parts of the basin for which few or no botanical data exist, and many rare and sparsely
distributed species remain undiscovered. Improved understanding of the ecological roles, dynamics and associat-
ions of the species of greatest importance for the maintenance of sustainable livelihoods and ecosystem services,
habitat restoration and adaptation to climate change is a high priority. In many cases these are common and
widespread species. Some of these issues are explored by looking at the Cristalino region in northern Mato Grosso
as a case-study. Effective integration, quality assessment, synthesis and application of existing data on the plant
diversity of the Amazon will help to address these issues. However, more targeted information is needed from the
ground. Future prioritisation of research effort will require a careful and pragmatic balance between the ‘traditi-
onal’ focus on rare and endemic species and species-rich communities, and the growing need to understand the
key ‘framework’ elements that will determine the future of the Amazon environment. Similar situations are faced
elsewhere in the tropics: for botanical research institutes in the 21st century this demands an urgent re-evaluation
of core activities and concerted engagement with the issues and challenges facing conservation in a context of
rampant population growth, climate change and environmental destruction.
Resumo. Apesar da existência de um grande volume de informação a respeito da diversidade vegetal da
Amazônia, uma série de obstáculos ainda dificulta a tomada de decisões devidamente informadas sobre con-
servação e manejo sustentável da região. Falta conhecimento com relação à distribuição das espécies, e às
interações entre diversidade e composição da vegetação, aos serviços do ecossistema e sua adaptabilidade e
resistência às flutuações climáticas. A distribuição da diversidade filogenética das espécies em relação às áreas
atualmente protegidas ainda é desconhecida, e muito pouco é sabido sobre o significado prático da variabil-
idade intraespecífica. A interpretação dos diferentes tipos de vegetação permanece extremamente simplificada,
com grandes áreas da bacia para as quais não há dados botânicos disponíveis ou os mesmos são insuficientes, e
muitas espécies raras ou esparsamente distribuídas ainda aguardam descobrimento. Uma melhor compreensão
dos papéis ecológicos, da dinâmica e das associações das espécies mais importantes para a manutenção de
estilos de vida sustentáveis e dos serviços do ecossistema, recuperação de áreas degradadas e adaptabilidade às
mudanças climáticas, são as grandes prioridades. Em muitos casos essas espécies são comuns e amplamente
distribuídas. Alguns desses temas são explorados utilizando a região do Cristalino, no norte do Mato Grosso,
como um estudo de caso. A integração efetiva, o controle da qualidade, a síntese e a aplicação dos dados
existentes da diversidade vegetal da Amazônia serão fundamentais para a solução dessas questões. No entanto,
ainda é necessário gerar a informação relevante a partir do estudo da área. A priorização das pesquisas futuras
requer um equilíbrio cuidadoso e prático entre o foco ‘tradicional’, interessado em espécies raras e endêmicas
e nas comunidades ricas em termos de espécies e a necessidade crescente de compreender os elementos da
‘estrutura’ que irá determinar o futuro do bioma amazônico. Situações semelhantes estão sendo enfrentadas
nos trópicos como um todo: os institutos botânicos do século 21 precisam reavaliar urgentemente as suas
principais atividades e engajar-se de modo coordenado para incluir os tópicos e desafios causados pelo cres-
cimento populacional desordenado, mudanças climáticas e destruição ambiental.

Accepted for publication November 2010.

1
HLAA, Royal Botanic Gardens, Kew, London, UK. e-mail: w.milliken@kew.org, d.zappi@kew.org
2
Fundação Ecológica Cristalino, Alta Floresta, Mato Grosso, Brazil.
3
Instituto de Pesquisas da Amazônia—INPA, Manaus, Brazil.
4
Royal Botanic Garden, Edinburgh, UK.

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011

692
Key Words. Alpha, Amazon, Amazonia, beta and gamma diversity, botanical survey, climate change, conservation
challenges, Mato Grosso, Parque Estadual Cristalino, vegetation.
Introduction
Biodiversity and conservation in the Amazon
With increasing recognition of the vital ecosystem
services provided by the world’s tropical forests,
public, scientific and political attention has returned
to the Amazon. The basin supports around 40% of the
world’s remaining rainforest, accounts for some 10%
of global terrestrial primary productivity, provides
approximately 16% of the world’s freshwater and
harbours 20 – 30% of the world’s plant species, of
which over half are endemic to the region (Melillo et
al. 1993; Muller-Karger et al. 1988; Salati & Santos
1998). Based on current trends in agricultural expan-
sion, 40% of the Amazon forests are predicted to be
eliminated by 2050 (Soares-Filho et al. 2006).
Biodiversity and ecosystem services are inextricably
linked and a thorough understanding of the former is
vital to successful management of the latter. Conserva-
tion and management of the Amazon basin has
proven challenging in a relatively stable climate; in
an increasingly dynamic one it is likely to become
substantially more so. Although modelling may help to
understand future trends both in climate and vegeta-
tion, uncertainties surrounding the influences of each
on the other, coupled with limited understanding of
the roles and significance of floristic diversity and
composition in this context, makes meaningful pre-
diction — and hence the development of appropriate
pre-emptive conservation strategies — difficult.
Informed action is critical and research resources
are limited. It is vital that efforts be directed towards
the most pressing data requirements. This invites a re-
evaluation of the types of information necessary for
effective decision-making, and of the impact of
current and recent research findings on conservation.
In this paper we provide a broad overview of the
state of knowledge of Amazonian vegetation and
botanical diversity, drawing on examples from recent
studies in the Cristalino region, in the Brazilian State
of Mato Grosso. We discuss the implications of specific
knowledge and likewise of current data lacunae for
conservation prioritisation and management, high-
lighting areas of potential future research.
Programa Flora Cristalino
Programa Flora Cristalino (PFC) was a collaborative
project developed between 2006 and 2009 by the Royal
Botanic Gardens Kew, the Cristalino Ecological Founda-
tion (FEC), the University of São Paulo (USP), the
University of the State of Mato Grosso (UNEMAT) and
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
the State Secretariat for the Environment of Mato
Grosso (SEMA). Its objectives were to support conserva-
tion in northern Mato Grosso with a particular focus on
management planning for the Cristalino State Park
(CSP), to assist the development of new private reserves
in the area, and to raise scientific knowledge and public
awareness of the region’s biodiversity. This was achieved
through a programme of collaborative botanical
research (vegetation mapping and inventory), training
and capacity building, public communication and the
development of educational materials.
The CSP lies in the basin of the Teles Pires River on the
northern boundary of Mato Grosso (Map 1), spanning
the municipalities of Novo Mundo and Alta Floresta. It is
situated within the Southern Amazonian Marginal
Depression, an area of considerable geological and
geomorphological complexity, characterised by gentle,
rounded hills and outcrops (principally sandstone and
granite) between 100 m and 400 m altitude (Brasil &
Alvarenga 1989; Ross 2003; SEPLAN/MT 1997). Soils
range from quartzite sands to red-yellow podzolics and
dark red dystrophic lithosols (SEPLAN/MT 2001b). The
climate is hot and humid with average temperatures
above 24°C, annual rainfall around 2,400 mm, and a
well-defined dry season lasting 3 – 5 months (Nimer
1989; Köppen 1948; SEPLAN/MT 2001a).
Established in 2000 and augmented in 2001, the Park
covers an area of 184,900 ha (Fig. 1). The adjacent
private reserves (Reservas Particular do Patrimônio
Natural — RPPN) Cristalino I, II e III (6,476 ha) and
RPPN Lote Cristalino (670 ha), managed by FEC, and
the Brazilian Air Force (FAB) Reserve, in the state of
Pará, form a contiguous unit of protected land that lies
directly on the frontier of the Amazon’s southern ‘arc
of deforestation’: the northwards expansion of agricul-
ture, ranching and logging that has devastated much of
the state’s natural vegetation.1
The state of knowledge of Amazonian
botanical diversity
Taxonomy and diversity
The question of how many plant species exist in
Amazonia arises repeatedly in the botanical literature,
1
In the first quarter of 2009 Mato Grosso was listed as the state
with the highest deforestation rate in the Brazilian Amazon
(INPE-DETER — http://www.amazonia.org.br/noticias/print.
cfm?id=296443; Imazon — http://www.imazon.org.br/
novo2008/publicacoes_ler.php?idpub=672).
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER? 693

Map 1. Location of the Cristalino State Park in Brazil.

Fig. 1. The Cristalino State Park in 2001, the year of its establishment, showing the complexity of the vegetation and the spread of
deforestation from the south. Adapted from LANDSAT TM 2001.

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011

694
with estimates ranging from 50,000 (e.g. Hubbell et al.
2008) to substantially lower (Silman 2007). These
figures, however, are necessarily based on educated
guesswork. The problems associated with such esti-
mates arise partly from the low density of collections
and their highly skewed distribution towards a very few
geographical centres (Nelson et al. 1990; Hopkins
2007; Schulmann et al. 2007). The overall density of
botanical collections in the Brazilian Amazon is about
0.1 per square kilometre and, although this is mas-
sively higher close to major research foci at Manaus,
Iquitos and Belém, elsewhere vast areas remain totally
unexplored botanically.
The accuracy of our knowledge of the plant species
of Amazonia should improve through databasing of
collections and comparison and compilation of data
from these and other sources. In Brazil, for example, a
list of all 33,885 currently known terrestrial plant
species has recently been published (Forzza et al.
2010), involving over 400 botanists coordinated by the
Rio de Janeiro Botanical Garden. In addition there are
efforts to database and repatriate herbarium data from
crucially important herbaria, such as the Mellon
funded Latin American Plants Initiative (LAPI) focus-
ing on type specimens, the National Science Founda-
tion (NSF) project digitising the New York Botanical
Garden herbarium collections, and forthcoming proj-
ects from the Brazilian National Research Council
(CNPq) to digitise Brazilian collections from impor-
tant European herbaria. In a few years it should be
possible to provide better estimates for the number of
known species in the region.
Fig. 2. Percentage of Sapotaceae specimens with x specimens in the IAN Herbarium; over half are represented by three collections
or fewer.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
A rapid query of the Instituto Nacional de Pesquisas
da Amazônia (INPA) herbarium database in Manaus,
which has been largely cleansed of inconsistencies in
spelling, names and synonymy that otherwise result in
high levels of data redundancy, indicates a total of
about 11,500 species recorded from the Brazilian
Amazon states of Acre, Amapá, Amazonas, Mato Grosso,
Pará and Roraima. Eventually it will be possible to bring
together — and likewise standardise — data from all
herbaria housing collections from the Amazon Basin,
and we might perhaps expect the final number of known
plant species registered in herbaria to be double or triple
this figure.
Though not specifically the subject of this paper,
there is compelling evidence that with more collecting
intensity the number of species known for the region
is likely to increase considerably. This is supported by
analyses of collection frequency in Amazonian herba-
ria which indicate that most described species have
been collected very rarely (Fig. 2), that in large study
plots most species are represented by very few
individuals (Fig. 3; Hubbell 2001), and that monogra-
phic treatments of principally Amazonian taxa indi-
cate no sign of an asymptote in the discovery of
sparsely distributed species (Fig. 4). Given that many
Amazonian species are locally rare and have limited
distributions, the low density and patchiness of
botanical collecting activity suggest that there remain
a large number of undescribed and uncollected
species in the region. Lack of botanical collection
may therefore reconcile the estimates of Amazonian
tree species made by Hubbell et al. (2008), derived
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER? 695

Fig. 3. Number of species with x individuals in study plots in Reserva Ducke (Manaus, Brazil). Data from 40,142 trees courtesy of
Carol Castilha.

from a neutral biodiversity theory that emphasises the
long ‘tail’ of rare taxa in the abundance curve for rain
forest trees (Figs 2 and 3).
The Flora of Reserva Ducke project (INPA/
Department for International Development 1992 –
1999) described 50 new species and almost doubled
the total known for this relatively small biological
reserve on the outskirts of Manaus, which was
previously thought to have been comprehensively
catalogued. This helped promote a revitalisation of
botanical research at INPA but does not reflect a
wider trend. It would be pleasing to report that plant
collection and taxonomy in the region are increas-
ing, but collecting effort in the Brazilian Amazon has

Fig. 4. Species discovery of Sapotaceae since 1700 (based on description dates from Flora Neotropica), split into widely distributed
species (dark blue), sparsely distributed (light blue) and intermediates. Whilst discovery of widespread species has reached an
asymptote, that of restricted species is still climbing steeply.

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011

696
been in steady decline over the last three decades.
Although activity remains higher in some of the
country’s other biomes, this is symptomatic of an
overall need for increased knowledge of the coun-
try’s flora (Giulietti et al. 2005; Sobral & Stehmann
2009). This phenomenon is by no means restricted to
Brazil or the Amazon: analyses of the GBIF (Global
Biodiversity Information Facility) and the Missouri
Botanical Garden VAST databases by Cayuela et al.
(2009) indicate trends of decreasing collecting activ-
ities across a range of tropical countries.
The evolving nature of our understanding of the
region’s flora is also reflected in ongoing taxonomic
accounts. The Meliaceae for example — important
components of the Amazonian tree flora — were
revised for Flora Neotropica in 1981 with a total of 120
species (Pennington & Styles 1981). A new revision,
yet to be published, will have raised this number by
47.5% (to 177) over a 30 year period, and juxtaposi-
tion of specimen- and species-distribution data from
the two studies (Fig. 5) shows significant advances in
the geographical spread and depth of the data set.
Much progress has been made over recent years in
the study of the distribution patterns of alpha-diversity
through large-scale syntheses of tree data from (prin-
cipally terra firme) forest plots. There are intrinsic
consistency problems with some of the underlying
data, ranging from plot shape and size to site-selection
criteria (habitat diversity versus homogeneity) and
species identification reliability. The first of these can
to some degree be overcome by the use of Fisher’s α
(Fisher et al. 1943); the last is more challenging. There
is a clear pattern of tree diversity across the basin with
highest levels in an east-west band to the south of the
equator (Silman 2007; Stropp et al. 2009; ter Steege et al.
2003), peaking in the upper (western) Amazon where
one-hectare plots with over 300 species of trees ≥ 10 cm
dbh (diameter at breast height) have been recorded
(on the northern and southern Amazonian fringes
there may be 100 – 150 or fewer). Similar patterns of
east-west diversity increase have been recorded by
Wittmann et al. (2006) for white-water flooded forests
(várzea).
These figures have been the focus of much
discussion and analysis in the literature in the contexts
of understanding the drivers of Amazonian alpha
diversity and predicting rates of species loss (Feeley
& Silman 2008; Hubbell et al. 2008; Pitman et al. 2002;
ter Steege et al. 2000, 2003, 2010). All of these analyses
suffer to some degree from weaknesses or inconsisten-
cies in the underlying data, generally recognised by
the authors in their own or each others’ work,
including oversimplification of vegetation classifica-
tion, lack of taxonomic specificity (some do not go
beyond the level of genus), incompleteness (for
example, reliance on GBIF data in one case) and
varying degrees of taxonomic rigour. Conclusions vary,
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
both in terms of extinction rates and causation: ter
Steege et al. (2003) and Pitman et al. (2002) suggest
that length of dry season is a critical factor, but
acknowledge the potential contribution of others,
including stem density and meta-population (as
reflected by vegetation area). More recent and
detailed analyses by ter Steege et al. (2010) suggest
that palaeoclimate, and its influence on speciation
and extinction, may prove more significant than
current factors in influencing alpha diversity in the
Amazon.
Data from two one-hectare terra firme forest plots at
Cristalino, inventoried in collaboration with the RAIN-
FOR programme,2 indicate diversity levels of between 103
and 122 species of trees ≥ 10 cm dbh per hectare. These
figures fall within the expected range for the peripheral
zone of the Amazon, as discussed above. Interpreting the
overall diversity of the flora in the Amazonian context is
more of a challenge. A total of 1,377 species have so far
been recorded at Cristalino, representing 627 genera and
151 families. The most species-diverse families and genera
were, respectively, Leguminosae (128), Rubiaceae (93),
Melastomataceae (52), Moraceae (43), Malvaceae3 (38),
Euphorbiaceae (36), Apocynaceae (35)3and Annonaceae
(34); and Psychotria (Rubiaceae — 25), Miconia
(Melastomataceae — 22), Ficus (Moraceae — 18), Inga
(Leguminosae — 16) and Pouteria (Sapotaceae — 13).
It is difficult to make valid comparisons of gamma
diversity between different parts of the Amazon as there
are few inventories that can be considered complete.
The inventory conducted at Cristalino is based on about
3,500 collections and the species/sampling curve was
still climbing steeply at the end of the study. It is likely
that further collecting would raise the species total to at
least 2,000. In spite of these difficulties, it is worth
comparing the data with other studies to gain a better
understanding of diversity patterns across the Amazon.
One of the most important attempts at this was made by
Clarke et al. (2001), comparing data from Iwokrama
Forest (Guyana) with four areas of Amazonia sensu lato
(two from Guyana, one from French Guiana and one
from Brazil). According to the authors the data are
strongly influenced by various factors including collect-
ing effort and the size of the study area.
At Iwokrama 3,887 collections were made represent-
ing 1,251 species (579 genera) in an area of 360,000 ha.
These results are similar to those from Cristalino, where
the area studied was substantially smaller (192,000 ha).
Clarke et al. (2001) concluded that the fact that over
50% of their species were represented by a single
2
RAINFOR is an international project bringing together a
network of researchers and permanent forest plots in Ama-
zonia to improve understanding of ecosystem dynamics and
relationships with climate.
3
Malvaceae and Apocynaceae are here used in their broad sense
following Souza & Lorenzi (2005).
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER? 697

Fig. 5. Collection and species abundance maps for the Meliaceae family from Flora Neotropica (1981 — left hand column) and its
unpublished revision (2009 data — right hand column). Data courtesy of T. D. Pennington.

specimen indicated an incomplete survey. The same is
true of the collections from Cristalino.
Regions with more diverse floras cited by Clarke et
al. (2001) included the Reserva Ducke in central
Brazilian Amazonia (Ribeiro et al. 1999), and Central
French Guiana (Mori et al. 1997). Species richness
reported from Ducke, with 2,079 species of vascular
plants, is high considering its relatively small area
(10,000 ha). Although this may reflect a greater
collecting effort over a longer period of time, with
over 11,000 fertile specimens made, it also reflects its
position within one of the proposed Tertiary and
Quaternary refuges (Prance 1990; Oliveira & Daly
1999). The survey of Central French Guiana (12,847
specimens representing 2,069 species in 140,000 ha)
also suggests high diversity, although the collecting
effort was again much greater than at Cristalino.
The inclusion of several different vegetation types
in the Cristalino study (i.e. the high beta diversity of
the region) contributed much to the total number of
species found in the area, as emphasised by the fact
that 59% of species were recorded in one habitat only.
Of these exclusive species, 31% (299) were from terra
firme forest and 19% from open habitats (open
campinarana and campo rupestre da Amazônia).
Understanding patterns of biogeography:
strengths and limitations of the data set
On the whole, plant biogeographic patterns remain
poorly understood within Brazil with few significant
studies made and little incorporation of phylogenetic
perspectives (Fiaschi & Pirani 2009). More resolved

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011

698
phylogenies containing more exhaustive sampling of
species are needed for Amazonian plants, both to
understand biogeographic history and to enable
identification of areas that contain most lineage
diversity (e.g. Forest et al. 2007), which might be
considered priorities for conservation. Nevertheless, in
the Amazon recent syntheses have improved our
understanding of patterns at higher taxonomic levels.
The major families in the terra firme tree flora, for
example, show distinct geographical centres of abun-
dance across the basin: Chrysobalanaceae and Lecy-
thidaceae in the east and the Guianas; Myristicaceae
and Urticaceae in the west; Burseraceae and Laur-
aceae in the south; Moraceae in the south-west; and
Sapotaceae in the centre (ter Steege et al. 2003, 2010).
Leguminosae tend to dominate throughout but are
particularly abundant on poor soils, for example in
the Guianas. At the generic level two gradients across
the basin, apparently related to soil fertility and dry
season length, have been identified, resulting in major
geographical differences in composition across large
distances (ter Steege et al. 2010).
The capacity of vegetation to adapt to climate
change, and the nature of the changes that it is likely
to undergo as a consequence, will doubtless be
influenced by its heterogeneity. Detailed mapping
and interpretation of the vegetation of the Amazon is
restricted to specific study sites, where data commonly
indicate higher levels of complexity and variability than
large-scale maps based on remote-sensing interpretation
Fig. 6. Phytosociological data from evergreen terra firme forest at Cristalino (point-centred quarter transects of trees≥10 cm dbh),
showing strong dominance of Burseraceae.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
would suggest. This emphasises the importance of
ground truthing and vegetation studies in the field.
During the present work certain similarities were
found between the flora of Cristalino and Acre which
will be discussed in more detail elsewhere (Milliken et
al. 2011). Meaningful analysis of species-level biogeo-
graphic patterns, however, is limited by problems with
species distribution data, as discussed above. Quanti-
tative sampling of the evergreen terra firme forest
(Fig. 6) at family level showed a strong dominance of
Burseraceae (principally due to abundance of Tetra-
gastris altissima (Aubl.) Swart), followed by Moraceae
and Leguminosae. In contrast, semi-deciduous forest
on nutrient-poor, sandy soils was heavily dominated by
Leguminosae (with Dialium guianense (Aubl.) Sand-
with particularly abundant), whilst Rubiaceae and
Bignoniaceae dominated the deciduous forest (nota-
bly Dialypetalanthus fuscescens Kuhlm., Tabebuia spp.).
The relatively high diversity of Moraceae in the
Cristalino region, both in tree plots and the overall
species inventory, coincides with the patterns shown by
ter Steege et al. (2000) for woody plant families, and
likewise the dominance of Burseraceae and Legumi-
nosae in the terra firme forests.
Within the semi-deciduous and deciduous forest
on sandy soils and granite substrates, a few tree
species characteristic of the semi-arid region of
Eastern Brazil — Tabebuia aurea (Silva Manso) Benth.
& Hook. f. ex S. Moore, Cordia alliodora (Ruiz & Pav.)
Oken, Coutarea hexandra (Jacq.) K. Schum. and
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER?
Sterculia striata A. St.-Hil & Naud. — were found that
corroborate the suggestions made by Pennington et
al. (2000) regarding the former expansion of dry-
forests during the Pleistocene. These authors sug-
gested that in drier episodes of the Pleistocene,
species currently characteristic of now isolated semi-
deciduous or deciduous forests may have been more
widespread.
As well as discovering species new to science,
records from Cristalino have extended the known
distribution of species previously recorded only from
Peru and Bolivia (for example, Psychotria ownbeyi
Standl. ex C. M. Taylor, Passiflora miniata Vanderpl.,
Hydrocotyle bonplandii A. Rich.) and likewise Amazonian
species not formerly known from the state of Mato
Grosso (for example, Hymenocallis tubiflora Salisb.,
Cymbopetalum longipes Benth. ex Diels, Murdannia
gardneri (Seub.) G. Brückn., Alchorneopsis floribunda
(Benth.) Müll. Arg., Manihot tristis Müll. Arg., Stigma-
phyllon palmatum (Cav.) A. Juss., Poecilanthe effusa
(Ducke) Huber, Strychnos cogens Benth., Calathea
capitata (Ruiz & Pav.) Lindl. and others). Amazonian
forests in Mato Grosso have received little attention
and the register at Cristalino of 485 species and 77
genera not previously included in the state checklist
(Dubs 1998) reflects this fact. As discussed, the extent
to which one can interpret these range extensions and
apparent disjunctions is to some degree limited by the
paucity of the data set. Data from the GBIF database,
for example (Fig. 7), can potentially be useful for
indicating broad patterns and associations in the flora,
but must be treated with a strong measure of caution
in the Amazonian context.
According to the Instituto Brasileiro de Geografia e
Estatística — IBGE (2004), the vegetation types found
in the Cristalino region are transitional between
evergreen forest, dry forest and savanna. Studies in
and around the Park have revealed a somewhat more
complex mosaic of habitats heavily influenced by
underlying edaphic, geological and hydrological fac-
tors, and in some cases by human influence. Vegeta-
tion types include evergreen terra firme forest
(generally on clayey soils), semi-deciduous forest
(sandy soils), deciduous forests (granite or sandstone
hills), campo rupestre da Amazônia (rocky scrub forma-
tions on sandstone and granite outcrops), open forest
formations with bamboo and palms, seasonally
flooded forest, campinarana (low forests on sandy soils
generally associated with hydrological stress) and
various riverine/lacustrine habitats, as well as secon-
dary forest, scrub and pasture.
To exemplify the lack of clarity around some
Amazon vegetation types we will briefly examine campo
rupestre da Amazônia. According to Giulietti & Pirani
(1988), campo rupestre vegetation is largely restricted to
the Serra de Espinhaço, a range of mountains running
from the Brazilian state of Bahia to Minas Gerais with
699
a disjunction in Goiás. It is associated with quartzitic/
sandstone outcrops generally around or above
1,000 m and is typically characterised by the diversifi-
cation of a suite of families including Velloziaceae,
Eriocaulaceae, Asteraceae and Melastomataceae, and
a high proportion of endemic species. Campo rupestre
da Amazônia is a term coined by Pires & Prance (1985)
to describe the discontinuous vegetation associated
with rock outcrops within the Amazon (Fig. 8), and
bears little relation floristically or geologically to the
campo rupestre of eastern Brazil (Daly & Mitchell 2000).
The examples found within the Cristalino region
fall into two different types depending on the local
substrate (granite or sandstone). In the sandstone
areas, depending on the topography and erosion of
the rocks (forming pockets of soil), this vegetation
may acquire a more woody form bearing some
physical resemblance to cerrado savanna. The compo-
sition affiliations of its tree species, however, are
stronger with campinarana vegetation.4 Overall, several
of its herbaceous and rupicolous species have proven
to be endemic to this type of vegetation (for example,
Encyclia caximboensis L. C. Menezes, Parkia cachimboensis
H. C. Hopkins, Aspilia paraensis (Huber) Santos,
Retiniphyllum parvifolium Steyerm., Piper cachimboense
Yunck., Thrasya auricoma Burman, Psittacanthus dentatus
Kuijt), with links to similar outcrops to the north of
Cristalino (Serra do Cachimbo, state of Pará). On
flatter low-lying sandstone outcrops another vegeta-
tion type occurs which, whilst closest to campinarana
gramíneo-lenhosa in the IBGE vegetation classification
(Veloso et al. 1991), shows some physiognomic sim-
ilarity with campo rupestre, and indeed shares some of
the genera commonly found in Eastern Brazil (Vellozia,
Xyris, Cephalostemon, Microlicia, Hyptis and Perama).
However, the species are either widespread in the
Amazonian campinaranas or occur also in the Guianan
and Venezuelan highlands and savannas. This vegeta-
tion type, elsewhere described from Amapá as Amazon
rock savanna (Pires-O’Brien 1992), is also poorly
researched and understood.
The granite outcrops found within the region fall
within the concept of inselberg (Sarthou et al. 2003),
and, although the species composition overlaps to a
certain extent with the sandstone campo rupestre areas,
a number of species were found exclusively on this
substrate (for example, Hymenocallis tubiflora “Salisb.”
4
Analysing the range of 131 species found in this vegetation
type, 23% (31% of the trees) were linked to campinarana
vegetation, 23% (25% of the trees) to cerrado and 12% (8% of
the trees) were widely distributed Amazonian species. The
remaining species were classified as widely distributed and,
although many of these do occur in campo/cerrado, they are
not exclusive to these habitats, occurring scattered throughout
Brazil and other countries e.g. Bolivia, Guiana, Venezuela and
Colombia.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
700 KEW BULLETIN VOL. 65(4)

Fig. 7. GBIF distribution data (November 2009) for six species from Cristalino (red circle): 1 Aspidosperma carapanauba Pichon
(Apocynaceae); 2 Metrodorea flavida K. Krause (Rutaceae); 3 Iriartea deltoidea Ruiz & Pav. (Arecaceae); 4 Celtis schippii Standl.
(Ulmaceae); 5 Raputiarana subsigmoidea (Ducke) Emmerich (Rutaceae), 6 Heliconia aemygdiana Burle-Marx (Heliconiaceae).
Background © 2009 Google, Image © 2009 Europa Technologies, 2009 DMapas, 2009 Maplink/TeleAtlas.

Philodendron acutatum “Schott" Ichthyothere rufa “Gard-
ner” Bromelia balansae “Mez” Bredemeyera floribunda
“Willd.”, Marsdenia sp. nov.), some of them endemic
to the region. As a general trend, the species found in
such inselbergs are more drought tolerant than the
Amazon forest species, and tend to be found else-
where in open habitats (rock outcrops associated with
rivers, open vegetation on sandy soils), similar to what
was found by Sarthou et al. (2003).
Genetic diversity of widespread species
and ecosystem resilience
Common and widespread tree species are the funda-
mental building blocks of Amazonian vegetation
(Pitman et al. 1999, 2001). These plants are in many
cases able to survive a wide range of edaphic, climatic
and hydrological conditions, some crossing the forest-
cerrado divide. Many of these common species play
important roles in human livelihoods, supporting the

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011

AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER? 701

Fig. 8. Campo rupestre da Amazônia vegetation on the Serra de Rochedo, Cristalino State Park.

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011

702
capacity of forest communities for sustainable
resource management. Although isolated studies of
selected species been conducted on regional scales,
for example, the Dendrogene Project (Martins-da-
Silva et al. 2001), there have been very few if any
basin-wide studies of the genetic variability of these
species (Pennington & Dick 2010). This hampers the
ability to answer key questions in relation to historical
biogeography and biome history (ter Steege et al.
2010), and to ecosystem resilience in the face of
climatic change. Many of the dominant tree species at
Cristalino, for example, are widespread across the
Amazon basin. Of the 15 most abundant species in
four quantitative studies of terra firme forest (two plots
and two plotless surveys ranging from evergreen to
semi-deciduous), all but two present large geograph-
ical ranges spanning varying ecological and climatic
conditions (Table 1).
Have the widespread species that dominate the
transitional Amazonian forest at Cristalino recently
expanded their range from the central Amazon
(which would predict low genetic diversity that is a
subset of central populations), or have they persisted
in some form of drought-adapted transitional forest
over millions of years (which would predict higher
genetic diversity)? Future molecular studies may help
to answer such questions.
The demonstrated importance of adapted local
genotypes in temperate forests (for example, Ennos et
al. 2000) is leading to an emphasis on the use of
locally sourced germplasm in temperate zone forest
rehabilitation. We are far from such a sophisticated
approach in Amazonia, and it is clearly imperative to
fill the vacuum with studies of local adaptation in
Amazonian trees. In the context of predicted climate
change, with shifting environmental conditions across
the Amazon basin, an understanding of the relation-
ships between genotype and resilience could play an
important role in conservation management, habitat
restoration and assisted migration.
Table 1. Abundant terra firme forest tree species at Cristalino and their geographical distributions.
Brosimum lactescens (S. Moore) C. C. Berg (Moraceae)
Cheiloclinium cognatum (Miers) A. C. Sm. (Celastraceae)
Dialium guianense (Aubl.) Sandwith (Leguminosae)
Micropholis guyanensis (A. DC.) Pierre (Sapotaceae)
Protium tenuifolium (Engl.) Engl. (Burseraceae)
Trichilia quadrijuga Kunth (Meliacaee)
Protium sagotianum Marchand (Burseraceae)
Tetragastris altissima (Aubl.) Swart (Burseraceae)
Abuta grandifolia (Mart.) Sandwith (Menispermaceae)
Pseudolmedia laevis (Ruiz & Pav.) J. F. Macbr. (Moraceae)
Rinoreocarpus ulei (Melch.) Ducke (Violaceae)
Celtis schippii Standl. (Ulmaceae)
Theobroma speciosum Willd. ex Spreng. (Malvaceae)
Iriartea deltoidea Ruiz & Pav. (Arecaceae)
Qualea homosepala Ducke (Vochysiaceae)
Protium cf. nitidifolium (Cuatrec.) Daly (Burseraceae)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
Challenges for meaningful modelling
of vegetation trends in the Amazon
Most studies of the effect of climate change on the
Amazon region have understandably focused on the fate
of the Amazon rain forest, but have neglected the type of
vegetation that might replace it under potentially
warmer and drier climates. For example, in response to
the often discussed scenario of ‘Amazon dieback’ (e.g.
Sitch et al. 2008; White et al. 1999), the crucial question
of what vegetation the Amazonian rain forest might ‘die
back’ to has been neglected. To address this question, it
is important to understand the distinctions between
dry-adapted biomes in the Neotropics such as savannas
(for example, the cerrados of central Brazil, Ratter et al.
2003), seasonally dry tropical forests (sensu Murphy &
Lugo 1986; Pennington et al. 2006, including the
caatingas of north eastern Brazil), and the chaco
woodlands of Argentina, Paraguay and Bolivia (e.g.
Prado 1993a, b). These biomes are not adequately
represented in some recent Neotropical vegetation
maps (e.g. Eva et al. 2004) which have subsequently
been used by earth system science for predictions of
vegetation response to a drier climate (e.g. Malhi et al.
2009). Eva et al.’s (2004) map, for example, confounds
chaco and seasonally dry tropical forest which, though
physiognomically similar, share virtually no woody
species in common (Prado 1993a, b). A key ecological
factor driving the distinction of some of these dry area
biomes is soil type — for example, cerrado savanna on
acid, dystrophic soil will grow alongside seasonally dry
tropical forest on base-rich soils under exactly the same
rainfall regime.
Within Amazonia itself, a crucial region to consider
under changing climates is the margin, which is
already under climatic tension. Palaeopalynological
research (e.g. Mayle et al. 2000; Mayle & Beerling
2004) demonstrates that Amazonian forest-savanna
and Amazonian forest-seasonally dry tropical forest
boundaries have moved remarkably rapidly since the
last glacial maximum (LGM), and ecological studies
Widespread through northern South America and Central America
Widespread through northern South America and Central America
Widespread through northern South America and Central America
Widespread through northern South America and Central America
Widespread through northern South America and Central America
Widespread through northern South America and Central America
Widespread through northern South America
Widespread through northern South America
Widespread through Amazonia and the Guianas
Widespread through Amazonia and the Guianas
Widespread through Amazonia and the Guianas
Widespread through western Amazonia, Guianas and Central America
Widespread around southern, eastern and western Amazonia
Western South America and Central America
Few records from Rondônia and Acre (W Amazon)
Not known
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER?
(Marimon et al. 2006; Ratter 1992) demonstrate that
Amazonian forest has continued to expand into
savanna areas in recent decades. This most likely
represents the continuation of a long-term process,
driven by increased precipitation, which has been
underway for the last three millennia and has yet to be
reversed by anthropogenic climate change (Mayle et
al. 2007).
Conservation planning and implementation:
identifying the priorities
Traditional conservation planning in the Amazon has
placed a heavy emphasis on sites with high levels of
species diversity and endemism. Nevertheless Kress et
al. (1998), in their evaluation of taxonomic data
distribution across a range of life forms in the
Amazon, found little correspondence between the
centres of biological diversity indicated by the data
and those recommended for conservation in earlier
quantitative studies. According to Silva et al. (2005),
the proportion of principal areas of endemism within
strict protected areas in the Brazilian Amazon ranges
from 0.3 – 11.7% (mean 4.8), with between 17.7 and
62.5% in protected areas sensu lato (including indige-
nous territories and sustainable use zones). The
authors argue for a conservation strategy that integra-
tes the protection of these areas with the establish-
ment of strategic conservation corridors at biome
level. Ter Steege et al. (2010) predict that the majority
of rare tree species (among an estimated 12,500 total
for the Amazon) occur in the western part of the
basin, where the climate was wetter during the
Pleistocene, making this a conservation priority from
the biodiversity perspective (Malhi et al. 2008).
Levels of species diversity are decoupled from
phylogenetic lineage diversity in species-rich biomes
such as the Cape Floristic region (Forest et al. 2007).
Better understanding of phylogenetic diversity across
the Amazon would therefore be useful in conserva-
tion planning. Estimates of phylogenetic diversity
(e.g. Faith 1992) might be generated from a genus-
level phylogeny for all Amazonian plants, or at a finer
scale from multiple phylogenies for key genera of
Amazonian plants that are ecologically dominant and
species-rich such as Inga, Protium, Eschweilera. Phylo-
genetic studies of such genera are under way (e.g.
Kursar et al. 2009; Erkens et al. 2007), and a genus-
level phylogeny of trees may be achievable if DNA
‘barcode’ markers start to be sequenced routinely in
inventory studies.
Analyses of conservation strategies for the Amazon
are becoming increasingly focused on managing the
predicted impacts of climate change (e.g. Malhi et al.
2008; Killeen & Solórzano 2008). Opinions vary as to
the likely changes that the Amazon will experience but
it is clear that successful management of existing
703
protected areas and strategic allocation of new ones
will be major factors in limiting these effects. Walker et
al. (2009) predict that effective conservation of
current protected areas in Brazil will provide a
sufficient buffer to avoid the ‘tipping-point’ precipitat-
ing change from forest to semi-arid vegetation.
Proposed approaches include conservation corridors
for species migration, minimising fragmentation, fire
control, watershed and river management, and the
protection of large-scale biological refuges. For corri-
dors to be effective as mitigators of the effects of climate
change they will need to incorporate environmental
gradients, taking into account ecotypes, altitude, soils
and precipitation (Killeen & Solórzano 2008).
As discussed, there has been much work on
measuring patterns of alpha diversity across the
Amazon but it is questionable how significant this is
in conservation terms. At a simplistic level, in terms of
species preservation it is the overall (gamma) diversity
of an area that is important. However, due to
ecological resilience afforded by high levels of beta
diversity, the ecotonal areas marginal to the Amazon
basin will possibly play an important role in the
adaptation to climate change and must be included
as a focus of specific conservation strategies (Killeen &
Solórzano 2008), in spite of their lower levels of
species diversity. Both the ‘optimistic’ and ‘non-optimis-
tic’ scenarios for the Amazon as proposed by Laurance et
al. (2001), taking into account land-use trends and
planned development projects (such as Avança Brasil),
predict greatest levels of forest loss along the southern
and eastern parts of the basin, and this has to some
degree been borne out by the last decade.
Implementation of strategic conservation approaches,
as we have shown, requires a better and more
integrated understanding of the flora of the region.
Cayuela et al. (2009), for example, evaluated the role
of species distribution modelling in conservation
decision-making and found that deficiencies in the
data set were a significant obstacle to the success of
this approach. They conclude that, when based on
adequate data, these techniques have considerable
potential to inform conservation strategies and plan-
ning and to assist the evaluation of climate change
impacts, although the rarest and most threatened
species are also those for which this process is least
likely to prove useful.
It is in the nature of many taxonomists, as innate
‘collectors’ of rarity and novelty, to undervalue or
overlook common and abundant species. Yet gaining a
better understanding of the ecological roles and
adaptations, population genetics and distributions of
these framework species, will greatly benefit applied
conservation and management. As proposed, the
complex nature of vegetation mosaics on the Amazon
periphery may provide a measure of buffering against
the effects of climate change, but it also raises concern
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
704
over the future of species restricted to specialised
vegetation types associated with localised soil/sub-
strate conditions occurring in isolated patches such
as white sand or rock outcrops. The ability of such
species to migrate will be influenced by their repro-
ductive biology and genetic variability, again high-
lighting the importance of gaining a better
understanding of these factors for successful adaptive
management.
It is widely recognised that successful conservation
requires effective stakeholder engagement and this is
particularly relevant in the Amazon. In Brazil, indige-
nous reserves cover about 20% of the basin and are
officially regarded as conservation units, boosting the
‘protected’ area of the Amazon by about 100%.
Unsurprisingly, there is clear evidence that these areas
offer better protection against deforestation and fire
than unprotected areas (Nepstad et al. 2006), and
although analysis shows no significant difference from
the level of protection afforded by unpopulated
protected areas, this may be due to their differing
contexts. The establishment of indigenous areas has
often been precipitated by contact with an advancing
development frontier, placing them directly in the
path of deforestation (Fig. 9). As such, several authors
have considered indigenous reserves to be the most
Fig. 9. The northern section of the Parakana Indigenous Reserve (Pará, Brazil) in 2009; the border of the reserve coincides clearly
with the limits of deforestation. Adapted from © Google 2009, Image © 2009 DigitalGlobe/2010 TerraMetrics, 2010 Maplink/
TeleAtlas.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
important barrier to forest loss in the region (Nepstad
et al. 2006; Peres 1994).
Supporting the capacity of indigenous and other
forest communities to remain as custodians of their
land is therefore an imperative. From the botanical
perspective this shifts the emphasis towards resource
inventory and management, development of sustain-
able economic alternatives (including through finan-
cial mechanisms such as REDD — Reduced
Emissions from Deforestation and Degradation),
adaptation to climate change and management of
genetic resources.
The CSP is a classic example of the challenges and
realities facing conservation in the Amazon. When
established in 2001 it already included substantial
areas under pasture. However, over subsequent years
further land was converted to cattle ranching and
smallholdings through illegal land invasion and ques-
tionable financial transactions. Unauthorised ‘ecotour-
ism’ infrastructure was established within the Park and
privately funded hydroelectric schemes initiated.
Some of the more accessible areas have also been
subjected to selective logging, and in 2006 there was a
concerted, albeit unsuccessful, attempt by Mato Gros-
so’s Vice-Governor to reduce its area by 27,500 ha,
including much of its most fertile land.
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER? 705

Fig. 10. Secondary vegetation and pasture in the Cristalino State Park. The striped areas were already cleared when the park was
established in 2001 and the solid areas were converted subsequently.

The park is of some considerable conservation
significance, lying in a strategic position in the southern
Amazon conservation corridor proposed by the WWF
(Morton et al. 2007). This has been identified as an
important buffer to northwards spread of the agricul-
tural frontier into the Amazon. The region is identified
by Maury (2004) as a priority conservation area by the
Brazilian Ministry of the Environment. Deforestation
models predict major loss of forest for the region,
including 76% of Mato Grosso’s 420,000 km2 of dry
forest (Soares-Filho et al. 2006).
Soil quality and habitat destruction in the Park are
intimately connected. The largest areas cleared for
cattle ranching coincide with the former occurrence
of open-forest formations (Fig. 10). Whilst such areas
are not the most productive in terms of biomass, they
are relatively easy to clear and their soils are locally
reputed to be well suited for agriculture and pasture.
This is not altogether surprising, as it has been
postulated that these open-forest formations, which
occur in a large band across the south of the Amazon,
may themselves be the consequence of former anthro-
pogenic activity, having been selected and cleared by
indigenous peoples that have since disappeared (Balée
& Campbell 1990).
Conversely the sandy soils that underlie much of
the park’s vegetation (specifically semi-deciduous
forest and campinarana) are poorly suited for agricul-
ture and pasture, and these areas have generally been
left intact by farmers. Nevertheless INCRA, the Brazil-
ian rural development agency, has recently established
‘assentamentos’ (blocks of smallholdings commonly
occupied by poor economic migrants from other parts
of the country) on extremely nutrient-poor, sandy soils
immediately to the north/northeast of the Park. The
people who have been settled in this area will, without
doubt, exhaust their meagre resources within a very
short time and be forced either to move on or to clear
further areas of forest. Meanwhile their activities have
introduced a significant fire hazard, principally
through burning cleared land outside the permitted
season, which has already resulted in wildfires within

Fig. 11. Sciadocephala sp. from the Cristalino State Park: a conservation priority or an interesting oddity? Photo G. Henicka and
map data for distribution of the genus from GBIF 2009.

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011

706
the park. The leakage of fire from agricultural areas
into adjacent forests is described by Betts et al. (2008)
as “the most pervasive and threatening synergy
between deforestation and climate change”.
In the development of the Park’s management
plan, Programa Flora Cristalino provided much of the
key data and recommendations for zoning and
prioritisation. The level of information required for
this is perhaps worth analysis for the benefit of
research prioritisation in similar situations. The most
significant data provided by the project were vegeta-
tion maps and the accompanying characterisations,
conservation evaluations and threat assessment for
each one of the component habitats. Prioritisation
and zoning were not based on species criteria
(diversity, rarity and so on), but rather on the
composition, diversity, importance, quality and con-
nectivity of vegetation types and habitats.
This is not to say that accurate species identification
was not important for this process: indeed it is vital for
meaningful analysis of vegetation. However, compar-
ison of vegetation composition tends to focus on the
most abundant or dominant species, which in turn
tend to be widespread and abundant in the Amazo-
nian context. So what was (or might have been) the
role of less common species in conservation planning
and management?
It is difficult to estimate the significance of rare or
new species encountered at Cristalino, partly because
of the level of uncertainty over whether their
restricted distribution is genuine or an artefact of
the incomplete Amazonian data set. In certain cases
there is evidence to suggest that their rarity or
novelty is associated with relatively high levels of
endemism in the nearby Serra de Cachimbo, whose
flora remains relatively poorly studied. For example
a new species of Ichthyothere (Asteraceae), found
among Amazon campo rupestre on sandstone hills
(Frisby & Hind 2011), almost certainly occurs in
similar vegetation types in the Cachimbo, of which
these hills are effectively outliers. Its discovery and
that of others in similar habitats (such as Hyptis sp.,
Dichorisandra sp.) highlight the need for additional
research in these habitats, and their inclusion as
conservation priorities.
Other cases are harder to interpret: a new species
of Sciadocephala (Asteraceae) has been found growing
in a single, highly restricted population, approximately
100 m from the edge of a farm within the Park (Fig. 11;
Biggs et al. 2011). This variegated forest herb — an
unusual habitat for its family — is only encountered
during the rainy season and is technically Critically
Endangered under the IUCN Red List criteria. This
might indeed be the case, although further research in
the region may identify additional populations. Cer-
tainly its occurrence in Mato Grosso appears to
represent a surprising disjunction in the genus, the
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
nearest record of which (Sciadocephala schultze-rhonhofiae
Mattf. in Diels) appears to be over 2,000 km away in
Peru (Fig. 11; GBIF 2010).
New species such as this are valuable for raising
public interest, and likewise for highlighting how
much we still have to discover yet may unwittingly be
losing. This new Sciadocephala, for example, is unlikely
to have a major impact on the way that conservation is
managed in the region, but it does represent a small
step forward in our knowledge of the flora of a
strategically important area. Whether it and the other
new and rare species discovered through the Cristalino
project prove to be of significance, or merely curious
footnotes in the wider and more pressing catalogue of
large-scale change in the vegetation, biodiversity, ecology
and livelihoods of the southern Amazon, remains to be
determined.
Acknowledgements
The authors would like to thank Terry Pennington for
sharing his unpublished Meliaceae data, Justin Moat
of Kew’s GIS Unit for help with maps, two anonymous
reviewers for their valuable comments, and the
organisers of the Kew 250th anniversary conference,
particularly Mike Fay and Rhian Smith, for arranging a
stimulating and diverse meeting.
Regarding Projeto Flora Cristalino, we would like
to thank CNPq for the collecting permit (EXC
13/07), Renato de Mello-Silva and staff at Universi-
dade de São Paulo, Renato A. de Farias, Gracieli S.
Henicka, José H. Piva and all staff at Fundação
Ecológica Cristalino, Cristalino Jungle Lodge and Hotel
Floresta Amazônica, José F. Ramos, Carlos Franciscon and
José Eduardo L. S. Ribeiro from INPA, Célia Soares, staff
and students from Universidade do Estado de Mato
Grosso - AF, staff from SEMA (Eliani Fachim, Eliani Pena,
Elton Silveira, Martinho Philippsen, Marcos Bessa and
Ênio Beltrante), Jovita Yesilyurt, Nicky Biggs, Sue Frisby
and many colleagues and students who participated in the
fieldwork.
References
Balée, W. & Campbell D. G. (1990). Evidence for the
successional status of liana forest (Xingu River
Basin, Amazonian Brazil). Biotropica 22: 36 – 47.
Betts, R. A., Malhi, Y. & Roberts, J. T. (2008). The
future of the Amazon: new perspectives from
climate, ecosystem and social sciences. Philos.
Trans., Ser. B 363: 1729 – 1735.
Biggs, N. & Hind, D. J. N. (2011). A new species of
Sciadocephala (Composiate) from Mato Grosso. Kew
Bull.
Brasil, A. E. & Alvarenga, S. M. (1989). Relevo. In: A.
C. Duarte (ed.), Geografia do Brasil. Vol. 1. Região
Centro-Oeste, pp. 53 – 72. IBGE, Rio d e Janeiro.
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER?
Cayuela, L., Golicher, D. J., Newton, A. C., Kolb, M., de
Albuquerque, F. S., Arets, E. J. M. M., Alkemade, J.
R. M. & Pérez, A. M. (2009). Species distribution
modelling in the tropics: problems, potentialities,
and the role of biological data for effective species
conservation. Trop. Conserv. Sci. 2: 319 – 352.
Clarke, H. D., Funk, V. A. & Hollowell, T. (2001). Plant
Diversity of the Iwokrama Forest, Guyana. Sida, Bot.
Misc. 21.
Daly, D. C. & Mitchell, J. D. (2000). Lowland vegetation
of Tropical South America — an overview. In: D.
Lentz (ed.), Imperfect balance: landscape transformations
in the pre-Columbian Americas, pp. 391 – 454. Colum-
bia University Press, New York.
Dubs, B. (1998). Prodromus Florae Matogrossensis, Part 1.
Betrona-Verlag, Switzerland.
Ennos, R., Worrell, R., Arkle, P. & Malcom, D. (2000).
Genetic variation and conservation of British native trees
and shrubs. Forestry Commission Technical Paper
31. Forestry Commission, Edinburgh.
Erkens, R. H. J., Chatrou, L. W., Maas, J. W., van der
Niet, T. & Savolainen, V. (2007). A rapid diversifi-
cation of rainforest trees (Guatteria; Annonaceae)
following dispersal from Central into South America.
Molec. Phylogenet. Evol. 44: 399 – 411.
Eva, H. D., Belward, A. S., De Miranda, E. E., Di Bella,
C. M., Gond, V., Huber, O., Jones, S., Sgrenzaroli,
M. & Fritz, S. (2004). A land cover map of South
America. Global Change Biol. 10: 731 – 744.
Faith, D. P. (1992). Conservation evaluation and
phylogenetic diversity. Biol. Conserv. 61: 1 – 10.
Feeley, K. J. & Silman, M. R. (2008). Unrealistic
assumptions invalidate extinction estimates. Proc.
Natl. Acad. Sci. U.S.A. 105: E121.
Fiaschi, P. & Pirani, J. R. (2009). Review of plant biogeo-
graphic studies in Brazil. J. Syst. Evol. 47: 477 – 496.
Fisher, R. A., Corbert, A. S. & Williams, C. B. (1943).
The relationship between the number of individu-
als in a random sample and an animal population.
J. Animal Ecol. 12: 42 – 58.
Forest, F., Grenyer, R., Rouget, M. T., Davies, J.,
Cowling, R. M., Faith, D. P., Balmford, A. Manning,
J. C., Proches, S., van der Bank, M., Reeves, G.,
Hedderson, T. A. J. & Savolainen, V. (2007).
Preserving the evolutionary potential of floras in
biodiversity hotspots. Nature 445: 757 – 760.
Forzza, R. C., Baumgratz, J. F. A., Bicudo, C. E. M.,
Carvalho Jr., A. A., Costa, A., Costa, D. P., Hopkins,
M., Leitman, P. M., Lohmann, L. G., Costa Maia, L.
Martinelli, G., Menezes, M., Morim, M. P., Nadruz
Coelho, M. A., Peixoto, A. L., Pirani, J. R., Prado, J.,
Queiroz, L. P., Souza, V. C., Stehmann, J. R.,
Sylvestre, L. S., Walter, B. M. T. & Zappi, D.
(2010). Lista de Espécies da Flora do Brasil, 2 vols.
Jardim Botânico do Rio de Janeiro.
Frisby, S. & Hind, D. J. N. (2011). A new species of
Ichthyothere (Compositae) from Mato Grosso. Kew Bull.
707
GBIF (2010). Global Biodiversity Information Forum
(http://www.gbif.org).
Giulietti, A. M. & Pirani, J. R. (1988). Patterns of
geographic distribution of some plant species from
the Espinhaço range, Minas Gerais and Bahia. In:
W. R. Heyer & P. E. Vanzolini (eds), Proceedings of a
Workshop on Neotropical Distribution Patterns, pp. 39 –
69. Academia Brasileira de Ciências, Rio de Janeiro.
____, Harley, R. M., Queiroz, L. P. de, Wanderley, M.
das G. L. & Van den Berg, C. (2005). Biodiversity
and conservation of plants in Brazil. Conserv. Biol.
19: 632 – 639.
Hopkins, M. J. G. (2007). Modelling the known and
unknown plant biodiversity of the Amazon Basin. J.
Biogeogr. 34: 1400 – 1411.
Hubbell, S. P. (2001). The unified neutral theory of
biodiversity and biogeography. Princeton University
Press, Princeton.
____, Fangliang, H., Condit, R., Borda-de-Água, L.,
Kellner, J. & ter Steege, H. (2008). How many tree
species are there in the Amazon and how many of
them will go extinct? Proc. Natl. Acad. Sci. U.S.A.
105: 11498 – 11504.
IBGE (Instituto Brasileiro de Geografia e Estatística)
(2004). Mapa da vegetação brasileira. 3a edição.
Ministério do Planejamento, Orçamento e Gestão.
Killeen, T. J. & Solórzano, L. A. (2008). Conservation
strategies to mitigate impacts from climate change
in Amazonia. Philos. Trans., Ser. B. 363: 1881 – 1888.
Köppen, W. P. (1948). Climatologia. Fondo de Cultura
Economica, México.
Kress, W. J., Heyer, W. R., Acevedo, P., Coddington, J.,
Cole, D., Erwin, T., Meggers, B. J., Pogue, M.,
Thorington, R. W., Vari, R. P., Weitzman, M. J. &
Weitzman, S. H. (1998). Amazonian biodiversity:
assessing conservation priorities with taxonomic
data. Biodivers. & Conserv. 7: 1577 – 87.
Kursar, T. A., Dexter, K. G., Lokvam, J., Pennington R. T.,
Richardson, J. E., Weber, M. G., Murakami, E., Drake,
C., McGregor, R. & Coley, P. D. (2009). The
importance of plant-herbivore interactions for diver-
sification and coexistence in the tropical tree genus
Inga. Proc. Natl. Acad. Sci. U.S.A. 106: 18073 – 18078.
Laurance, W. F., Cochrane, M. A., Bergen, S., Fearnside,
P. M., Delamônica, P., Barber, C., D’Angelo, S. &
Fernandes, T. (2001). The future of the Brazilian
Amazon. Science 291: 438 – 439.
Malhi, Y., Roberts, J. T., Betts, R. A., Killeen, T. J., Li, W.
& Nobre, C. A. (2008). Climate change, deforesta-
tion, and the fate of the Amazon. Science 319: 169 –
172.
____, Aragão, L. E. O. C., Galbraith, D., Huntingford,
C., Fisher, R., Zelazowski, P., Sitche, S., McSweeney,
C. & Meir, P. (2009). Exploring the likelihood and
mechanism of a climate-change-induced dieback of
the Amazon rainforest. Proc. Natl. Acad. Sci. U.S.A.
106: 20610 – 20615.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
708
Marimon, B. S., de S. Lima, E., Duarte, T. G., Chier-
egatto, L. C. & Ratter, J. A. (2006). Observations on
the vegetation of northern Mato Grosso, Brazil. IC.
An analysis of the cerrado-amazonian forest ecotone.
Edinburgh J. Bot. 63: 323 – 341.
Martins-da-Silva, R. C. V., Degen, B., & Amaral, W.
(2001). Improving conservation values of managed
forests: The Dendrogene Project in the Brazilian
Amazon. Unasylva 209: 25 – 33.
Maury, C. M. (org.) (2004). Biodiversidade Brasileira:
Avaliação e identificação de áreas e ações prioritárias
para conservação, utilização sustentável e repartição dos
benefícios da biodiversidade nos biomas brasileiros. Min-
istério do Meio Ambiente, Secretaria de Biodiversi-
dade e Florestas, PROBIO. Brasília, DF.
Mayle, F. E., Burbridge, R. & Killeen, T. J. (2000).
Millenial-Scale Dynamics of Southern Amazonian
Rain Forests. Science 290: 2291 – 2294.
____ & Beerling, D. J. (2004). Late quaternary changes
in Amazonian ecosystems and their implications for
global carbon cycling. Palaeogeogr. Palaeoclimatol.
Palaeoecol. 214: 11 – 25.
____, Langstroth, R. P., Fischer, R. A. & Meir, P.
(2007). Long-term forest-savannah dynamics in the
Bolivian Amazon: implications for conservation.
Philos. Trans., Ser. B 362: 291 – 307.
Melillo, J. M., McGuire, A. D., Kicklighter, D. W.,
Moore, B., Vörösmarty, C. J. & Schloss, A. L. (1993).
Global climate change and terrestrial net primary
production. Nature 363: 234 – 240.
Milliken, W. et al. (2011). Cristalino: new insights
into the vegetation of the Southern Amazonian
fringe. Biotropica.
Mori, S., Cremers, G., Gracie, C., Granville, J. J., Hoff,
M. & Mitchell, J. D. (1997). Guide to the vascular
plants of Central French Guiana, Part 1. Pterido-
phytes, Gymnosperms, and Monocotyledons. Mem.
New York Bot. Gard. 76: 1 – 422.
Morton, D. C., Shimabukuro, Y. E., Rudorff, B. F. T.,
Lima, A., Freitas, R. M. & Derfries, R. S. (2007).
Conservation Challenge at the Agricultural Fron-
tier: deforestation, fire, and land use dynamics in
Mato Grosso. Revista Ambiente e Água — An Inter-
disciplinary J. Appl. Sci. 2: 5 – 20.
Muller-Karger, F. E., McCain, C. R. & Richardson, P. L.
(1988). The dispersal of the Amazon’s water. Nature
333: 56 – 59.
Murphy, P. & Lugo, A. E. (1986). Ecology of tropical
dry forest. Ann. Rev. Ecol. Syst. 17: 67 – 88.
Nelson, B. W., Fereira, A. C., da Silva, M. F. &
Kawasaki, M. L. (1990). Endemism centres, refugia
and botanical collection density in Brazilian Ama-
zonia. Nature 345: 714 – 716.
Nepstad, D., Schwartzmann, S., Bamberger, B.,
Santilli, M., Ray, D., Schlesinger, P., Lefebvre,
P., Alencar, A., Prinz, E., Fiske, G. & Rolla, A.
(2006). Inhibition of Amazon deforestation and
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
fire by parks and indigenous lands. Conserv. Biol.
20: 65 – 73.
Nimer, E. (1989). Clima. In: A. C. Duarte (ed.),
Geografia do Brasil, Vol. 1. Região Centro-Oeste, pp.
23 – 34. IBGE, Rio de Janeiro.
Oliveira, A. A. & Daly, D. C. (1999). Geographic
distribution of tree species occurring in the region
of Manaus, Brazil: implications for regional diversity
and conservation. Biodivers. & Conserv. 8: 1245 –
1259.
Pennington, R. T. & Dick, C. W. (2010). Diversification
of the Amazonian flora and its relation to key
geological and environmental events: a molecular
perspective. In: C. Hoorn, H. Vonhof & F. Wesselingh
(eds), Amazonia, Landscape and Species Evolution: a
Look into the Past, pp. 373 – 385. Blackwell, Oxford.
____, Prado, D. E. & Pendry, C. A. (2000). Neotropical
Seasonally Dry Forest and Quaternary Vegetation
Changes. J. Biogeogr. 27: 261 – 273.
____, Ratter, J. A. & Lewis, G. P. (2006). An overview of
the plant diversity, biogeography and conservation of
neotropical savannas and seasonally dry forests. In: R.
T. Pennington, G. P. Lewis & J. A. Ratter (eds),
Neotropical savannas and seasonally dry forests: plant
biodiversity, biogeography and conservation, pp. 1 – 29.
CRC Press, Boca Raton, Florida.
Pennington, T. D. & Styles, B. T. (1981). Meliaceae.
Flora Neotrop. Monogr. 28.
Peres, C. A. (1994). Indigenous reserves and nature
conservation in Amazonian Forests. Conserv. Biol. 8:
586 – 588.
Pires, J. M. & Prance, G. T. (1985). The Vegetation
types of the Brazilian Amazon. In: G. T. Prance & T.
E. Lovejoy (eds), Amazonia, pp. 109 – 145. Perga-
mon Press, Oxford.
Pires-O’Brien, M. J. (1992). Report on a remote
swampy rock savanna at the mid-Kari river basin,
Lower Amazon. Bot. J. Linn. Soc. 108: 21 – 33.
Pitman, N. C. A., Terborgh, J., Silman, M. R. & Nuñez,
P. (1999). Tree species distributions in an upper
Amazonian forest. Ecology 80: 2651 – 2661.
____, ____, ____, ____, Neill, D. A., Cerón, C. E.,
Palacios, W. A. & Aulestia, M. (2001). Dominance
and distribution of tree species in upper Amazo-
nian terra firme forests. Ecology 82: 2101 – 2117.
____, ____, ____, ____, ____, ____, ____ & ____
(2002). A comparison of tree species diversity in
two upper Amazonian forests. Ecology 83: 3210 –
3224.
Prado, D. E. (1993a). What is the Gran Chaco
vegetation in South America? I. A review. Contri-
bution to the study of flora and vegetation of the
Chaco. V. Candollea 48: 145 – 172.
____ (1993b). What is the Gran Chaco vegetation in
South America? II. A redefinition. Contribution to
the study of flora and vegetation of the Chaco. VII.
Candollea 48: 615 – 629.
AMAZON VEGETATION: HOW MUCH DON’T WE KNOW AND HOW MUCH DOES IT MATTER?
Prance, G. T. (1990). The floristic composition of
Central Amazonian Brazil. In: A. H. Gentry (ed.),
Four Neotropical Forests, pp. 112 – 140. Yale University
Press.
Ratter, J. A. (1992). Transitions between cerrado and
forest vegetation in Brazil. In: P. A. Furley, J. Proctor
& J. A. Ratter (eds), Nature and dynamics of forest–
savannah boundaries, pp. 417 – 429. Chapman &
Hall, London.
____, Bridgewater, S. & Ribeiro, J. F. (2003). Analysis
of the floristic composition of the Brazilian cerrado
vegetation III: comparison of the woody vegetation
of 376 areas. Edinburgh J. Bot. 60: 57 – 109.
Ribeiro, J. E. L. S., Hopkins, M. J. G., Vicentini, A.,
Sothers, C. A., Costa, M. A. S., Brito, J. M., Souza,
M. A. D., Martins, L. H. P., Lohmann, L. G.,
Assunção, P. A. C. L., Pereira, E. C., Silva, C. F.,
Mesquita, M. R. & Procópio, L. C. (1999). Flora da
Reserva Ducke, Guia de Identificação. DfID & INPA,
Manaus.
Ross, J. L. S. (2003). Os fundamentos da geografia da
natureza. In: J. L. S. Ross (ed.), Geografia do Brasil,
pp. 13 – 65. Edusp, São Paulo.
Salati, E. & Santos, A. A. (1998). The Amazon and
Global Issues. In: M. L. D. Freitas (ed.), Amazonia,
Heaven of a New World, pp. 7 – 22. Editora Campus,
Rio de Janeiro.
Sarthou, C., Villiers, J. F. & Ponge, J. F. (2003). Shrub
Vegetation on Tropical Granitic Inselbergs in
French Guiana. J. Vegetation Sci. 14: 645 – 652.
Schulmann, L., Toivonen, T. & Ruokolainen, K.
(2007). Analysing botanical collecting effort in
Amazonia and correcting for it in species range
estimation. J. Biogeogr. 34: 1388 – 1399.
SEPLAN/MT (1997). Geomorfologia (texto). Zonea-
mento Sócio-econômico Ecológico. PRODEAGRO. Min-
istério de Integração Nacional.
____ (2001a). Distribuição da Pluviosidade Média Anual
(1983 – 1994). Zoneamento Sócio-econômico Ecológico.
PRODEAGRO. Ministério de Integração Nacional.
http://www.seplan.mt.gov.br/.
____ (2001b). Mapa de solos. Zoneamento Sócio-econômico
Ecológico. PRODEAGRO. Ministério de Integração
Nacional. http://www.seplan.mt.gov.br/.
Silman, M. R. (2007). Plant species diversity in
Amazonian forests. In: M. B. Bush & J. R. Flenley
(eds), Tropical Rainforest Responses to Climatic Change,
pp. 269 – 294. Springer, Berlin.
Silva, J. M. da, Rylands, A. B. & Fonseca, G. A. B. da
(2005). O destino das áreas de endemismo da
Amazônia. Megadiversidade 1: 124 – 131.
Sitch, S., Huntingford, C., Gedney, N., Levyz, P. E.,
Lomas, M., Piao, S. L., Betts, R., Ciais, P., Cox, P.,
Friedlingstein, P., Jones, C. D., Prentice, I. C. &
Woodward, F. I. (2008). Evaluation of the terrestrial
carbon cycle, future plant geography and climate-
carbon cycle feedbacks using five Dynamic Global
709
Vegetation Models (DGVMs). Global Change Biol. 14:
2015 – 2039.
Soares-Filho, B. S., Nepstad, D. C., Curran, L. M.,
Coutinho, G. C., Garcia, R. A., Ramos, C. A., Voll,
E., McDonald, A., Lefebvre, P. & Schlesinger, P.
(2006). Modelling conservation in the Amazon
basin. Nature 440: 520 – 523.
Sobral, M. & Stehmann, J. M. (2009). An analysis of
new angiosperm species discoveries in Brazil
(1990 – 2006). Taxon 58: 227 – 232.
Souza, V. C. & Lorenzi, H. (2005). Botânica Sistemática.
Instituto Plantarum de Estudos da Flora Ltda.,
Nova Odessa, SP.
Stropp, J., ter Steege, H., Malhi, Y., ATDN and
RAINFOR. (2009). Disentangling regional and local
tree diversity in the Amazon. Ecography 32: 46 – 54.
ter Steege, H., Sabatier, D., Castellanos, H., van Andel,
T., Duivenvoorden, J., Oliveira, A. A., Ek, R. C.,
Lilwah, R., Maas, P. & Mori, S. (2000). A regional
perspective: analysis of Amazonian floristic compo-
sition and diversity. In: H. ter Steege (ed.), Plant
Diversity in Guyana, with recommendations for a
National Protected Area Strategy. Tropenbos Series 18:
19 – 34.
____, Pitman, N., Sabatier, D., Castellanos, H., Van Der
Hout, P., Daly, D. C., Silveira, M., Phillips, O. L.,
Vasquez, R., Van Andel, T., Duivenvoorden, J.,
Oliveira, A. A., Ek, R., Lilwah, R., Thomas, R., Van
Essen, J., Baider, C., Maas, P., Mori, S., Terborgh, J.,
Núñez Vargas, P., Mogollón, H. & Horchler, P. J.
(2003). A spatial model of tree alpha-diversity and
density for the Amazon Region. Biodivers. & Conserv.
12: 2255 – 2277.
____ & RAINFOR (2010). Contribution of current and
historical processes to patterns of tree diversity and
composition in the Amazon. In: C. Hoorn & F. P.
Wesselingh, Amazonia, landscape and species evolution:
a look into the past, pp. 349 – 359. Wiley-Blackwell,
Oxford.
Veloso, H. P., Rangel Filho, A. L. R. & Lima, J. C. A.
(1991). Classificação da Vegetação Brasileira, adaptada
a um sistema universal. Fundação Instituto Brasileiro
de Geografia e Estatística (IBGE), Rio de Janeiro.
Walker, R., Moore, N. J., Arima, E., Perz, S.,
Simmons, C., Caldas, M., Vergara, D. & Bohrer,
C. (2009). Protecting the Amazon with protected
areas. Proc. Natl. Acad. Sci. U.S.A. 106: 10582 –
10586.
White, A., Cannell, M. G. R. & Friend, A. D. (1999).
Climate change impacts on ecosystems and the
terrestrial carbon sink: a new assessment. Global
Environmental Change 9: S21 – S30.
Wittmann, F., Schöngart, J., Montero, J. C., Motzer, T.,
Junk, W. J., Piedade, M. T. F., Queiroz, H. L. &
Worbes, M. (2006). Tree species composition and
diversity gradients in white-water forests across the
Amazon basin. J. Biogeogr. 33: 1334 – 1347.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2011