Amazon Vegetation

Amazon vegetation: how much don't we know and how much does it matter?
Authors(s): William Milliken, Daniela Zappi, Denise Sasaki, Mike Hopkins and R. Toby
Pennington
Source: Kew Bulletin, Vol. 65, No. 4, Plant Conservation for the Next Decade: A Celebration
of Kew's 250 th Anniversary (2010), pp. 691-709
Published by: Springer on behalf of Royal Botanic Gardens, Kew
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KEW BULLETIN VOL. 65: 691-709 (2010)
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 existencia de um grande volume de informafao a respeito da diversidade vegetal da
Amazonia, uma serie de obstaculos ainda dificulta a tomada de decisSes devidamente informadas sobre con
servaâo e manejo sustentavel da regiao. Falta conhecimento com relaâo a distribuifao das especies, e as
interafoes entre diversidade e composifao da vegetafao, aos services do ecossistema e sua adaptabilidade e
resistencia as flutuafoes climaticas. A distribuiâo da diversidade filogenetica das especies em relafao as areas
atualmente protegidas ainda e desconhecida, e muito pouco e sabido sobre o significado pratico da variabil
idade intraespecifica. A interpretafao dos diferentes tipos de vegetaâo permanece extremamente simplificada,
com grandes areas da bacia para as quais nao ha dados botanicos disponiveis ou os mesmos sao insuficientes, e
muitas especies raras ou esparsamente distribuidas ainda aguardam descobrimento. Uma melhor compreensao
dos papeis ecologicos, da dinamica e das associates das especies mais importantes para a manutenfao de
estilos de vida sustentaveis e dos services do ecossistema, recuperafao de areas degradadas e adaptabilidade as
mudanfas climaticas, sao as grandes prioridades. Em muitos casos essas especies sao comuns e amplamente
distribuidas. Alguns desses temas sao explorados utilizando a regiao do Cristalino, no norte do Mato Grosso,
como um estudo de caso. A integraâo efetiva, o controle da qualidade, a sintese e a aplicaâo dos dados
existentes da diversidade vegetal da Amazonia serao fundamentais para a solufao dessas questoes. No entanto,
ainda e necessario gerar a informaâo relevante a partir do estudo da area. A priorizaâo das pesquisas futuras
requer um equilibrio cuidadoso e pratico entre o foco 'tradicional', interessado em especies raras e endemicas
e nas comunidades ricas em termos de especies e a necessidade crescente de compreender os elementos da
'estrutura' que ira determinar o futuro do bioma amazonico. Situafoes semelhantes estao sendo enfrentadas
nos tropicos como um todo: os institutos botanicos do seculo 21 precisam reavaliar urgentemente as suas
principals atividades e engajar-se de modo coordenado para incluir os topicos e desaiios causados pelo cres
cimento populacional desordenado, mudanfas climaticas e destruifao ambiental.
Accepted for publication November 2010.

HLAA, Royal Botanic Gardens, Kew, London, UK. e-mail: w.miiliken@kew.org, d.zappi@kew.org
Fundagao Ecol6gica Cristalino, Alta Floresta, Mato Grosso, Brazil.
Instituto de Pesquisas da AmazoniaINPA, Manaus, Brazil.

Royal Botanic Garden, Edinburgh, UK.
The Board of Trustees of the Royal Botanic Gardens, Kew, 2011
KEW BULLETIN VOL. 65(4)
Key Words. Alpha, Amazon, Amazonia, beta and gamma diversity, botanical survey, climate change, conservation
challenges, Mato Grosso, Parque Estadual Cristalino, vegetation.
Introduction
the State Secretariat for the Environment of Mato
Biodiversity and conservation in the Amazon

With increasing recognition ot 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
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.
1 he CSP lies in the basin of the Teles Pires River on the
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).
northern boundary of Mato Grosso (Map 1), spanning

the municipalities of Novo Mundo and Alta Floresta. It is
Biodiversity and ecosystem services are inextricably

linked and a thorough understanding of the former is
vital to successful management of the latter. Conserva
geomorphological complexity, characterised by gentle,

rounded hills and outcrops (principally sandstone and
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.
Ln this paper we provide a broad overview ol 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.
situated within the Southern Amazonian Marginal

Depression, an area of considerable geological 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 24C, annual rainfall around 2,400 mm, and a
well-defined dry season lasting 3-5 months (Nimer

1989; Koppen 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 Patrimonio
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
Para, 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
1 he question or how many plant species exist in
Amazonia arises repeatedly in the botanical literature,
^rograma Flora Cristalino

frograma flora (.nstalmo (PH,) was a collaborative
project developed between 2006 and 2009 by the Royal
Botanic Gardens Kew, the Cristalino Ecological Founda
tion (FEC), the University of Sao Paulo (USP), the

University of the State of Mato Grosso (UNEMAT) and
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).
s) The Board ot Trustees ot the Royal Botanic Gardens, Kew, 2011
amazon vegetation: how much don't we know and how much does it matter?
693
Para
Mato
Grosso
Alta Floresta
Guaranta do
Carlinda
Norte
Novo Mundo
*
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.
CEW BULLETIN VOL. 65(4)
594
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
A rapid query of the Institute) Nacional de Fesquisas

da Amazonia (INPA) herbarium database in Manaus,
which has been largely cleansed of inconsistencies in
spelling, names and synonymy that otherwise result in
mates arise partly from the low density of collections

and their highly skewed distribution towards a very few
high levels of data redundancy, indicates a total of

about 11,500 species recorded from the Brazilian
geographical centres (Nelson et al. 1990; Hopkins
Amazon states of Acre, Amapa, Amazonas, Mato Grosso,

Para 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
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 Belem, elsewhere vast areas remain totally
unexplored botanically.
The accuracy of our knowledge of the plant species
f 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
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
ing on type specimens, the National Science Founda

tion (NSF) project digitising the New York Botanical
Garden herbarium collections, and forthcoming proj
sparsely distributed species (Fig. 4). Given that many
ects from the Brazilian National Research Council
botanical collecting activity suggest that there remain
(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.
Amazonian species are locally rare and have limited

distributions, the low density and patchiness of
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
35
30
25
20
15
10
L.
UJ cn ^ kO ^
~iiii i ir
Njrorotorouiuiuiujuj ^ ia ^ ^ iî
u> cn *>J VO -i to vi nJ ^ tu w vJ iO U) vi xj io *
Fig. 2. Percentage of Sapotaceae specimens with x specimens in the IAN Herbarium; over halt are represented by three collections
or fewer.
amazon vegetation: how much don t we know and how much does it matter?
695
40
20
W Vl sj -
-TTTtThl mMln
~n~H m
_rsjrsjNjKjrsjujuiuiuju>XkXkk-u.ucnr^x^
UJ d <kU)v/i*-Jk0 Ui in \j -> Ui in M VO
i??
O rs> ^
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).
reserve on the outskirts of Manaus, which was
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
botanical research at INPA but does not reflect a
previously thought to have been comprehensively

catalogued. This helped promote a revitalisation of
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
100
90
80
70
60
50
40
30
20
10
0
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.
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
both in terms of extinction rates and causation: ter
ities across a range of tropical countries.
Amazon.
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
Data from two one-hectare terra firme forest plots at

Cristalino, inventoried in collaboration with the RAIN
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
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
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),
data, ranging from plot shape and size to site-selection
Melastomataceae (52), Moraceae (43), Malvaceae3 (38),

Euphorbiaceae (36), Apocynaceae (35) !and 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
criteria (habitat diversity versus homogeneity) and
diversity between different parts of the Amazon as there
species identification reliability. The first of these can
are few inventories that can be considered complete.

The inventory conducted at Cristalino is based on about
cipally terra firme) forest plots. There are intrinsic

consistency problems with some of the underlying
to some degree be overcome by the use of Fisher's a

(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
(varzea).
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
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 smsu 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
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,
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.
Malvaceae and Apocynaceae are here used in their broad sense

following Souza & Lorenzi (2005).
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 Amazonia).
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

698
phylogenies containing more exhaustive sampling of

species are needed for Amazonian plants, both to
would suggest. This emphasises the importance of
identification of areas that contain most lineage

diversity (e.g. Forest et al. 2007), which might be
found between the flora of Cristalino and Acre which
understand biogeographic history and to enable
ground truthing and vegetation studies in the field.

During the present work certain similarities were
will be discussed in more detail elsewhere (Milliken et
considered priorities for conservation. Nevertheless, in
al. 2011). Meaningful analysis of species-level biogeo
the Amazon recent syntheses have improved our
graphic patterns, however, is limited by problems with
understanding of patterns at higher taxonomic levels.

The major families in the terra firme tree flora, for
species distribution data, as discussed above. Quanti

tative sampling of the evergreen terra firme forest
example, show distinct geographical centres of abun

dance across the basin: Chrysobalanaceae and Lecy
thidaceae in the east and the Guianas; Myristicaceae
Burseraceae (principally due to abundance of Tetra
and Urticaceae in the west; Burseraceae and Laur

aceae in the south; Moraceae in the south-west; and
(Fig. 6) at family level showed a strong dominance of

gastris altissima (Aubl.) Swart), followed by Moraceae
and Leguminosae. In contrast, semi-deciduous forest

on nutrient-poor, sandy soils was heavily dominated by
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
Leguminosae (with Dialium guianense (Aubl.) Sand

with particularly abundant), whilst Rubiaceae and
Bignoniaceae dominated the deciduous forest (nota
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
bly Dialypetalanthus fuscescens Kuhlm., Tabebuia spp.).
geographical differences in composition across large

distances (ter Steege et al. 2010).
The capacity of vegetation to adapt to climate
species inventory, coincides with the patterns shown by

ter Steege et al. (2000) for woody plant families, and
change, and the nature of the changes that it is likely
nosae in the term firme forests.
influenced by its heterogeneity. Detailed mapping
on sandy soils and granite substrates, a few tree

species characteristic of the semi-arid region of
to undergo as a consequence, will doubtless be
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
The relatively high diversity of Moraceae in the

Cristalino region, both in tree plots and the overall
likewise the dominance of Burseraceae and Legumi

Within the semi-deciduous and deciduous forest
Eastern Brazil Tabebuia aurea (Silva Manso) Benth.

& Hook. f. ex S. Moore, Cordia alliodora (Ruiz & Pav.)
Oken, Coutarea hexandra (Jacq.) K. Schum. and
Simaroubaceae [3]
Nyctaginaceae [6]
Rubiaceae [9]
Celastraceae [12]
Arecaceae [11]
Apocynaceae [4]
Euphorbiaceae [14]
Urticaceae [11]
Lauraceae [13]
Rutaceae [12]
Mynsticaceae [15]
Lecythidaceae [7]
Malvaceae [17]
elastomataceae [13]
Vochysiaceae [9]
I Relative density
Meliaceae [19]
I Relative dominance
Sapotaceae [23]
I Relative frequency
Leguminosae [41]
Moraceae [61]
Burseraceae [92]
000
20.00 3000 4000
60 00
Importance Value Index
Fig. 6. Phytosociological data from evergreen terra firme forest at Cristalino (point-centred quarter transects of treesslO cm dbh),
showing strong dominance of Burseraceae.
699
Sterculia striata A. St.-Hil & Naud. were found that
a disjunction in Goias. It is associated with quartzitic/
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,
sandstone outcrops generally around or above
records from Cristalino have extended the known
distribution of species previously recorded only from
Peru and Bolivia (for example, Psychotria ownbeyi

Standi, ex C. M. Taylor, Passiflora miniata Vanderpl.,
Hydrocotyle bonplandiiA. 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. Briickn., Alchomeopsis Jloribunda
(Benth.) Mull. Arg., Manihot tristis Mull. 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.
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 Amazonia 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 Cristalxno 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 Para). On
flatter low-lying sandstone outcrops another vegeta

tion type occurs which, whilst closest to campinarana
gramineo-lenhosa in the IBGE vegetation classification
(Veloso et al. 1991), shows some physiognomic sim
According to the Instituto Brasileiro de Geograha e

Estatistica IBGE (2004), the vegetation types found
ilarity with campo rupestre, and indeed shares some of

the genera commonly found in Eastern Brazil (Vellozia,
Xyris, Cephalostemon, Microlicia, Hyptis and Perama).
in the Cristalino region are transitional between

evergreen forest, dry forest and savanna. Studies in
Amazonian campinaranas or occur also in the Guianan
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
However, the species are either widespread in the

and Venezuelan highlands and savannas. This vegeta
tion type, elsewhere described from Amapa 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 el al. 2003),
and, although the species composition overlaps to a
hills), campo rupestre da Amazonia (rocky scrub forma

tions on sandstone and granite outcrops), open forest
certain extent with the sandstone campo rupestre areas,
formations with bamboo and palms, seasonally
substrate (for example, Hymenocallis tubijlora "Salisb."
a number of species were found exclusively on this
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 Amazonia. According to Giulietti & Pirani
(1988), campo rupestre wegetation is largely restricted to
the Serra de Espinhaco, a range of mountains running
from the Brazilian state of Bahia to Minas Gerais with
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.
700
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 Standi.
(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 mfa "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
amazon vegetation: how much don't we know and how much does it matter? 701
Fig. 8. Campo rupestre da Amazonia vegetation on the Serra de Rochedo, Cristalino State Park.
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 el al. 2001), there have been very few if any
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
basin-wide studies of the genetic variability of these
vegetation that might replace it under potentially
species (Pennington & Dick 2010). This hampers the
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
ability to answer key questions in relation to historical
biogeography and biome history (ter Steege el 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.
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
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 8c Pav.) J. F. Macbr. (Moraceae)
Rinoreocarpns ulei (Melch.) Ducke (Violaceae)
Celtis schippii Standi. (Ulmaceae)
Theobroma speciosum Willd. ex Spreng. (Malvaceae)
Iriartea deltoidea Ruiz 8c Pav. (Arecaceae)
Qualea homosepala Ducke (Vochysiaceae)
Protium cf. nitidifolium (Cuatrec.) Daly (Burseraceae)
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 western Amazonia, Guianas and Central America
Widespread around southern, eastern and western Amazonia
Western South America and Central America
Few records from Rondonia and Acre (W Amazon)
Not known
(Marimon et al. 2006; Ratter 1992) demonstrate that
703
protected areas and strategic allocation of new ones
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
will be major factors in limiting these effects. Walker et
underway for the last three millennia and has yet to be
ing change from forest to semi-arid vegetation.

Proposed approaches include conservation corridors
reversed by anthropogenic climate change (Mayle et

al. 2007).
al. (2009) predict that effective conservation of

current protected areas in Brazil will provide a
sufficient buffer to avoid the 'tipping-point' precipitat
for species migration, minimising fragmentation, fire
control, watershed and river management, and the
Conservation planning and implementation:
protection of large-scale biological refuges. For corri

dors to be effective as mitigators of the effects of climate
identifying the priorities
change they will need to incorporate environmental
Traditional conservation planning in the Amazon has

placed a heavy emphasis on sites with high levels of
gradients, taking into account ecotypes, altitude, soils

and precipitation (Killeen & Solorzano 2008).
As discussed, there has been much work on
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
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 &
Solorzano 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 Avanâ 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
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
likely to prove useful.

It is in the nature of many taxonomists, as innate
inventory studies.
Analyses of conservation strategies for the Amazon
better understanding of the ecological roles and

adaptations, population genetics and distributions of
are becoming increasingly focused on managing the

predicted impacts of climate change (e.g. Malhi et al.
2008; Killeen & Solorzano 2008). Opinions vary as to
the likely changes that the Amazon will experience but
it is clear that successful management of existing
these framework species, will greatly benefit applied
'collectors' of rarity and novelty, to undervalue or

overlook common and abundant species. Yet gaining a
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
704
KEW BULLETIN VOL 65(4)
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
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
management.
cial mechanisms such as REDD Reduced
It is widely recognised that successful conservation

requires effective stakeholder engagement and this is
particularly relevant in the Amazon. In Brazil, indige
Emissions from Deforestation and Degradation),

adaptation to climate change and management of
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
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
although analysis shows no significant difference from
tionable financial transactions. Unauthorised 'ecotour
the level of protection afforded by unpopulated

protected areas, this may be due to their differing
contexts. The establishment of indigenous areas has
ism' infrastructure was established within the Park and
often been precipitated by contact with an advancing
subjected to selective logging, and in 2006 there was a

concerted, albeit unsuccessful, attempt by Mato Gros
development frontier, placing them directly in the

path of deforestation (Fig. 9). As such, several authors
have considered indigenous reserves to be the most
privately funded hydroelectric schemes initiated.

Some of the more accessible areas have also been
so's Vice-Governor to reduce its area by 27,500 ha,

including much of its most fertile land.
Fig. 9. The northern section of the Parakana Indigenous Reserve (Para, 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.
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 (Balee

& 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.
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
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
each one of the component habitats. Prioritisation
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
and zoning were not based on species criteria

(diversity, rarity and so on), but rather on the
large-scale change in the vegetation, biodiversity, ecology

and livelihoods of the southern Amazon, remains to be
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
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
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 Sao Paulo, Renato A. de Farias, Gracieli S.
Henicka, Jose H. Piva and all staff at Fundafao

Ecologica Cristalino, Cristalino Jungle Lodge and Hotel
Floresta Amazonica, Jose F. Ramos, Carlos Franciscon and
Jose Eduardo L. S. Ribeiro from INPA, Celia 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

Enio Beltrante), Jovita Yesilyurt, Nicky Biggs, Sue Frisby
and many colleagues and students who participated in the
fieldwork.
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
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Amazon vegetation: how much don't we know and how much does it matter?

KEW BULLETIN VOL. 65: 691-709 (2010)

Resumo. Apesar da existencia de um grande volume de informafao a respeito da diversidade vegetal da

Accepted for publication November 2010.

Instituto de Pesquisas da AmazoniaINPA, Manaus, Brazil.

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

KEW BULLETIN VOL. 65(4)

challenges, Mato Grosso, Parque Estadual Cristalino, vegetation.

the State Secretariat for the Environment of Mato

Biodiversity and conservation in the Amazon

services provided by the world's tropical forests,

of global terrestrial primary productivity, provides

Grosso (SEMA). Its objectives were to support conserva

management planning for the Cristalino State Park

through a programme of collaborative botanical

al. 1993; Muller-Karger et al. 1988; Salati & Santos

northern boundary of Mato Grosso (Map 1), spanning

Biodiversity and ecosystem services are inextricably

geomorphological complexity, characterised by gentle,

tion and management of the Amazon basin has

understand future trends both in climate and vegeta

are limited. It is vital that efforts be directed towards

the most pressing data requirements. This invites a re

effective decision-making, and of the impact of

state of knowledge of Amazonian vegetation and

knowledge and likewise of current data lacunae for

situated within the Southern Amazonian Marginal

granite) between 100 m and 400 m altitude (Brasil &

climate is hot and humid with average temperatures

above 24C, annual rainfall around 2,400 mm, and a

well-defined dry season lasting 3-5 months (Nimer

covers an area of 184,900 ha (Fig. 1). The adjacent

The state of knowledge of Amazonian

^rograma Flora Cristalino

tion (FEC), the University of Sao Paulo (USP), the

with the highest deforestation rate in the Brazilian Amazon

cfm?id=296443; Imazon http://www.imazon.org.br/

s) The Board ot Trustees ot the Royal Botanic Gardens, Kew, 2011

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

CEW BULLETIN VOL. 65(4)

with estimates ranging from 50,000 (e.g. Hubbell et al.

2008) to substantially lower (Silman 2007). These

A rapid query of the Institute) Nacional de Fesquisas

mates arise partly from the low density of collections

high levels of data redundancy, indicates a total of

geographical centres (Nelson et al. 1990; Hopkins

Amazon states of Acre, Amapa, Amazonas, Mato Grosso,

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

f Amazonia should improve through databasing of

list of all 33,885 currently known terrestrial plant

species has recently been published (Forzza et al.

crucially important herbaria, such as the Mellon

funded Latin American Plants Initiative (LAPI) focus

Though not specifically the subject of this paper,

analyses of collection frequency in Amazonian herba

plots most species are represented by very few

individuals (Fig. 3; Hubbell 2001), and that monogra

ing on type specimens, the National Science Founda

sparsely distributed species (Fig. 4). Given that many

ects from the Brazilian National Research Council

botanical collecting activity suggest that there remain

(CNPq) to digitise Brazilian collections from impor