holocene vegetation change and the mammal …...south america in the late pleistocene and holocene...

ORIGINALARTICLE

Holocene vegetation change and themammal faunas of South Americaand Africa

Mario de Vivo* and Ana Paula Carmignotto

Museu de Zoologia, Universidade de Sao

Paulo, Sao Paulo, Brazil

*Correspondence: Mario de Vivo, Museu de

Zoologia, Universidade de Sao Paulo, Av.

Nazare, 481 Ipiranga, Sao Paulo, SP 04263-000,

Brazil. E-mail: [email protected]

ABSTRACT

Aim Although sharing many similarities in their vegetation types, South America

and Africa harbour very dissimilar recent mammal faunas, not only

taxonomically but also in terms of several faunistic patterns. However late

Pleistocene and mid-Holocene faunas, albeit taxonomically distinct, presented

many convergent attributes. Here we propose that the effects of the Holocene

climatic change on vegetation physiognomy has played a crucial role in shaping

the extant mammalian faunistic patterns.

Location South America and Africa from the late Pleistocene to the present.

Methods Data presented here have been compiled from many distinct sources,

including palaeontological and neontological mammalian studies, palaeo-

climatology, palynology, and publications on vegetation ecology. Data on

Pleistocene, Holocene and extant mammal faunas of South America and Africa

allowed us to establish a number of similar and dissimilar faunistic patterns

between the two continents across time. We then considered what changes in

vegetation physiognomy would have occurred under the late Pleistocene last

glacial maximum (LGM) and the Holocene climatic optimum (HCO) climatic

regimes. We have ordained these proposed vegetation changes along rough

physiognomic seral stages according to assumptions based on current botanical

research. Finally, we have associated our hypothesized vegetation changes in

South America and Africa with mammalian faunistic patterns, establishing a

putative causal relationship between them.

Results The extant mammal faunas of South America and Africa differ widely in

taxonomical composition; the number of medium and large species they possess;

behavioural and ecological characteristics related to herbivore herding, migration

and predation; and biogeographical patterns. All such distinctions are mostly

related to the open formation faunas, and have been completely established

around the mid-Holocene. Considering that the mid-Holocene was a time of

greater humidity than the late Pleistocene, vegetation cover in South America and

Africa would have been dominated by forest or closed vegetation landscapes, at

least for most of their lower altitude tropical regions. We attribute the loss of

larger-sized mammal lineages in South America to the decrease of open

vegetation area, and their survival in Africa to the existence of vast savannas in

formerly steppic or desertic areas in subtropical Africa, north and south of the

equator. Alternative explanations, mostly dealing with the disappearance of South

American megamammals, are then reviewed and criticized.

Main conclusions The reduction of open formation areas during the HCO in

South America and Africa explains most of the present distinct faunistic patterns

between the two continents. While South America would have lost most of its

open formations within the 30� latitudinal belt, Africa would have kept large areas

Journal of Biogeography (J. Biogeogr.) (2004) 31, 943–957

ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi 943

INTRODUCTION

South America and Africa share many landscape similarities,

including the presence of extensive rain forests, savannas,

steppes and deserts, but their contemporary mammalian

faunas are quite distinct (Keast, 1972; Vrba, 1993). Convergent

patterns such as would be expected as a result of faunistic

evolution over similarly structured landscapes (Bourliere,

1973) are actually not striking (McNaughton et al., 1993;

Cristoffer & Perez, 2003). The faunas differ widely in

taxonomical composition, they differ in the number of

medium and large species each continent possess; behavioural

and ecological characteristics related to herbivore herding,

migration and predation; and biogeographical patterns. How-

ever, this has not always been the case. Both faunas have

presented, at least from the Miocene onwards, convergent

characteristics, with diversified forest and open-formation

faunas, the latter inhabited by medium to large grazers and

browsers, all preyed upon by specialized carnivores (Cooke,

1972; Patterson & Pascual, 1972). In the Quaternary, this

scenario remained almost unaltered in Africa but changed

dramatically in South America, where open-formation mam-

mal fauna lost practically all similarities to that of Africa.

Most authors believe that South America and Africa have

evolved their quite differently structured extant mammalian

faunas due entirely to distinct causes acting on each continent.

Considerable attention has been paid to the role of humans

(e.g. Martin, 1967, 1984), climatic change (e.g. Ochsenius,

1985; Cartelle, 1999) or the impact of North American

immigrant mammals on the ecosystems (e.g. May, 1978;

Marshall et al., 1982; Webb, 1985; Marshall & Cifelli, 1990). A

few faunistic comparisons have also appeared, but they have

been exclusively descriptive (Keast, 1972; Bourliere, 1973).

Vrba (1993) argued that global climatic events should be

employed in explaining faunistic patterns in South America

and Africa, however, she primarily described how climate

change affected the faunistic composition in South America as

a result of the Great American Interchange of the late Pliocene

and its after-effects during the Pleistocene. Additionally, all

previous attempts to study the faunas have assumed that the

late Pleistocene was the most significant period of change

(Martin & Klein, 1984; Ochsenius, 1985; Marshall & Cifelli,

1990; Webb & Rancy, 1996; Cartelle, 1999), but recent effort at

dating fossil mammals in South America revealed several

extinct species as living well into the middle Holocene (Faure

et al., 1999; Baffa et al., 2000). Faure et al. (1999) dated a fossil

assemblage from north-eastern Brazil including the camelid

Palaeolama, the horse Equus and the giant armadillos,

Glyptodon and Hoplophorus, as c. 8490 and 6890 yr bp. Baffa

et al. (2000) dated a Toxodon from a karstic cave in south-

eastern Brazil to between 8000 and 5400 yr bp. This places

these large mammals well into the middle Holocene, and

considerably changes all previous views that the South

American megafauna would have been extinct by that time.

Besides, it indicates that analogously similar faunistic patterns

between South America and Africa have persisted beyond the

late Pleistocene.

Here we will demonstrate how the same late Pleistocene and

Holocene global events could have simultaneously affected the

faunas and disrupted the pattern of shared analogous similar-

ities between South America and Africa. The impact of climate

change on vegetation at these times certainly affected floristic

composition. However, physiognomy change was probably

more significant for the mammalian faunas, with landscapes

alternatively showing denser and sparser facies through time.

Higher yearly average precipitation existing during the Holo-

cene climatic optimum (HCO) would favour denser vegetation

physiognomies, thus reducing the availability of savanna-like

habitats for open formation faunas. However, lower averages

would open existing physiognomies and open formation

faunas would be benefited. Our continental level reconstruc-

tion of the vegetation changes in South America and Africa

provide a model that indicates why the same climatic events

led to a diversification of savanna mammal patterns in Africa

while practically extirpating them in South America.

IDENTIFYING FAUNISTIC PATTERNS

Past and present faunistic patterns

South America and Africa have always possessed taxonomically

distinct mammal communities throughout the Tertiary and

Quaternary (Patterson & Pascual, 1972; Maglio & Cooke, 1978;

Wilson & Reeder, 1993). This is significant in the sense that

any past or present similarities in ecological or behavioural

patterns are analogous, not homologous.

Throughout the Tertiary and Quaternary, South America

harboured 20 orders of mammals and Africa 13 (only terrestrial

forms; aquatic and volant mammals excluded). The recent fauna

is much less diverse for South America, with only 12 extant

suitable to the open formation mammalian fauna in areas presently occupied by

desert and semi-arid vegetation. Thus, the same general climatic events that affected

South America in the late Pleistocene and Holocene also affected Africa, leading to

our present day faunistic dissimilarities by maintaining the African mammalian

communities almost unchanged while dramatically altering those of South America.

Keywords

Mammals, South America, Africa, vegetation change, HCO, LGM.

M. de Vivo and A. P. Carmignotto

944 Journal of Biogeography 31, 943–957, ª 2004 Blackwell Publishing Ltd

orders and 11 in Africa (eight ordinal extinctions for South

America and only two for Africa). Considering the extant faunas

of the two continents, seven orders are shared, but have only two

genera in common, the carnivores Panthera and Mustela, which

are practically cosmopolitan taxa (Nowak, 1999).

Below is a summary of ecological, behavioural and bioge-

ographical patterns that involve past and recent mammal

faunas in South America and Africa.

Diversification of continental faunas across weight categories

Figure 1 shows that Africa is richer in number of species for

any category above 5 kg; only in the weight category below

5 kg is South America richer than Africa (622 and 587 species,

respectively). There are no reliable weight estimates for extinct

mammals in both continents but Anderson (1984) furnishes

some adequate size estimates. If Pleistocene faunas could be

included in our graph, diversity within the weight categories

would have been quite similar between South America and

Africa. Most of the extinct South American medium and large-

sized mammals were associated with open formations

(MacFadden & Shockey, 1997; Cartelle, 1999; Rancy, 1999;

Cristoffer & Peres, 2003) therefore attracting attention to this

particular assemblage and its African counterpart.

Presence of grazer and mixed grazer-browser terrestrial

herbivores

Among the largest African herbivores are the elephant, zebra,

rhinoceros, giraffe and hippopotamus, but the bulk of its

savanna diversity lies in the multitude of medium and large-

sized bovids (Bigalke, 1972; Peters, 1983). The extant South

American counterparts are the comparatively much less

impressive tapir, capybara, deer and the Andean camelids.

All South American herbivores are either mixed grazers and

browsers, or exclusive browsers, while Africa has a large

number of grazers (McNaughton & Georgiadis, 1986). As

South America has extensive areas of open vegetation and did

possess a diversified grazer herbivore fauna (Patterson &

Pascual, 1972; Rancy, 1999; MacFadden, 2000), the loss of this

guild strongly affects any contemporary comparison. In fact,

bovids, which are responsible for much of the present day

distinctiveness of Africa, failed to enter South America at the

end of the Pliocene. This could lead us to think that their mere

absence would promote the faunistic distinction just described,

but South America did possess autochtonous herbivore orders,

such as Litopterna and Notoungulata which never extended

their distribution outside the Americas. Finally, even if all

bovids are excluded from the analysis, Africa would still

present an assemblage of large mammals for which no

equivalents can be found in modern South America.

Herding and migratorial behaviour

Group size among South American herbivores is usually small,

and solitary species are not infrequent, while Africa possesses

many herding herbivores, with groups ranging from dozens to

hundreds of individuals. These herds frequently migrate

seasonally in search of better pastures, while no herbivore

migration is known in the extant South American mammal

fauna except for small altitudinal shifts in the Andean camelids

(Kingdom, 1979; Nowak, 1999). The bulk of African mammal

species exhibiting herding and migratorial behaviours require

vast expanses of open vegetation to perform them

(Owen-Smith, 1988; Kappelman et al., 1997). The fossil record

shows that Plio-Pleistocene and present African mammal

faunas were similarly structured (Vrba, 1993; Kappelman

et al., 1997), but this is not clear for the Pleistocene mammals

of South America. Cartelle & Bohorquez (1982) believe that

the terrestrial sloth Eremotherium may have been gregarious,

while Webb (1999) envisages ‘vast herds of herbivores’ for the

continent. We suspect that South American gomphoteriid

mastodons may have lived in groups, as well as the horses

(Equus), as do phylogenetically close forms of today. The

Xenarthra is a monophyletic group and all extant xenarthrans

are solitary; the most parsimonious hypothesis is to suppose

that giant armadillos and terrestrial sloths would also have

been solitary, otherwise independent acquisitions of social

behaviour would have to be accepted. This is, of course,

possible, but less probable. Finally, we simply have no parallel

for how to characterize the herding behaviour of animals

belonging to extinct endemic orders of South American

mammals, such as the Litopterna and Notoungulata. These

animals could have formed ‘vast herds’, lived in small groups,

lived solitarily, or any combination of the former.

Specialized carnivore hunting and social behaviour: presence

of carrion eaters

All of the living South American predators hunt solitarily (with

the exception of canid Speothos venaticus, a small forest

dweller), while Africa has both solitary and group-organized

forms. Group organized hunters are mostly open vegetation

Nu

mb

er o

f sp

ecie

s

– – – –

Figure 1 Number of species per weight category; South America

in black and Africa in grey. Weight data from several sources,

mainly Nowak (1999). Africa presents more mammal species in all

categories above 5 kg, while South America is richer in the

category below 5 kg (see text).

Vegetation change and mammal faunas in South America and Africa

Journal of Biogeography 31, 943–957, ª 2004 Blackwell Publishing Ltd 945

forms, like lions (Panthera leo), wild hunting dogs (Lycaon)

and hyenas (Hyaena, Crocuta). As prey size and availability are

factors that strongly influence the predation patterns of

carnivores (Vrba, 1980), group hunting has probably evolved

as a response to herd herbivory and the complex behaviours

involved can only be fully performed in open areas. South

America presented, for instance, sabre tooth felids adapted to

preying on large mammals and even specialized ‘giant’ vampire

bats, but these predators disappeared with their prey

(Simpson, 1980; Reig, 1981; Trajano & de Vivo, 1991). Finally,

Africa has a group of socially organized carnivores (hyenas)

that behave as opportunistic carrion eaters, which has no

extant mammalian equivalent in South America. For carrion to

be a major food source, it is probable that a large herbivore

biomass must be available.

Contemporary biogeographical patterns

Today, South America has two distinct mammalian faunistic

provinces, the Brazilian and the Patagonian (Hershkovitz,

1972). The Brazilian subregion actually extends from Central

America to Colombia, west of the Andes, and to the northern

half of Ecuador. To the east of the Andean cordillera, it

encompasses almost the entire tropical and subtropical South

America southwards to Bolivia and from there across Paraguay

to southern Brazil and adjacent Uruguay. The Patagonian

subregion includes the remaining parts of the continent: all of

South America west of the Andes from central Ecuador

southwards, and east of the Andes from south Bolivia to

Argentina and Uruguay. The Andes have no parallel in Africa

in continental extension and average altitude. Their presence

distorts the expected zonal distribution of alternating latitu-

dinal humid and drier life zones due to their powerful

influence as a barrier to moisture carrying winds. To the west

of the Andes, dry and very dry climates prevail in tropical

latitudes even at sea level southwards from central Ecuador.

On the contrary, the expected zonal humid temperate climates

at latitudes around 60� appear only in Chile, and to the other

side of the Andes, the Argentinean Patagonia is steppic in

climate. The Patagonian province of South America includes

many genera and even families that do not occur in the

Brazilian subregion and vice versa. At least 27 genera of

rodents occur exclusively in the southern South America or at

the climatically equivalent high altitudes in the tropical Andes

(e.g. Lagostomus, Dolichotis, Abrocoma, Andalgalomys, Phyllo-

tis, Graomys and Eligmodontia), the camelids Llama and

Vicugna, the cervids Pudu and Hippocamelus, the armadillos

Chaetophractus, Chlamyphorus and Zaedyus, the carnivore

Lyncodon, while entire families are tropical (the tree sloths

Bradypodidae and Megalonychidae; the Myrmecophagidae

anteaters; Cebidae and Atelidae primates; Echimyidae, Ago-

utidae and Dinomyidae histricognath rodents; Redford &

Eisenberg, 1992; Eisenberg & Redford, 1999). In Africa, the

most common pattern is the replacement of vicariant species

and subspecies along the north–south axis of open formations

(Haltenorth & Diller, 1977; Kingdom, 1977, 1979). Genera

presenting vicariant patterns for species and subspecies include

giraffes (Giraffa), dik-diks (Madoqua), kudus (Tragelaphus),

oryx (Oryx), antelopes of the genus Hippotragus, the lechwe

(Kobus), reedbucks (Redunca), gazelles (Gazella), African

buffalo (Syncerus), zebras (Equus), white rhinoceros (Cerato-

therium), jackals (Canis), hyenas (Hyaena, Crocuta) and lions

(P. leo). The patterns just described probably emerged due to

the disappearance, in South America, of open vegetation large

mammals and their survival in Africa. The fossil record reveals

the presence of some vicariant species in South America

represented by Patagonian and Brazilian provinces species

(Cartelle, 1999). Nevertheless the fossil record is not accurate

enough to support or reject the presence of vicariant taxa along

the South American continent as a whole.

RESULTS AND DISCUSSION

Commentaries on faunistic patterns

The patterns 1 to 4 described above are real: they underline the

distinctiveness of the extant South American and African

mammalian faunas. One question is if these patterns can in

fact be associated with the open formations. Another question

relates to the forest faunas: are they just as distinctive as the

open formation ones?

Our assumption is that large size, herding and migratory

behaviour, and social hunting techniques are strongly related to

open formations. This is not only intuitive, but also supported by

several lines of research (Clutton-Brock & Harvey, 1983; Peters,

1983; Owen-Smith, 1988; Kappelman et al., 1997). The terrest-

rial medium to large herbivores of the African rain forest are

either small-sized lineages (e.g. Cephalophinae bovids) or

dwarfed vicariants to savanna taxa. The duikers (Cephalophi-

nae) represent a bovid lineage that is clearly forest adapted. Their

size is small (around 20 kg body mass), and they are either

solitary or live in small groups, up to three individuals

(Haltenorth & Diller, 1977; Peters, 1983). The forest buffalo

(Syncerus caffer nanus) is a distinct taxon than the savanna

forms, its body mass reaches a maximum of 300 kg, a weight

comparable with that of the South American tapirs. The two

savanna subspecies are much larger, reaching up to 800 kg body

mass. The differences in group size are remarkable: the forest

buffalo lives in groups of three to 12 individuals, while the

savanna’s taxa range from 20 to 2000 (Haltenorth & Diller,

1977). The forest elephant (Loxodonta cyclotis, a distinct species

according to Roca et al., 2001), is also quite smaller, with up to

half the weight and size of the savanna form. Average group sizes

are also quite distinct. The forest elephant averages 3.2 individ-

uals per group, while savanna elephants average 10, but the latter

can form loose aggregations of up to 1000 individuals in the

rainy season (Haltenorth & Diller, 1977). It is revealing that

forest elephants actually use mostly forest clearings for much of

their activities; moving through the forest along paths to reach

alternative clearings (Vanleeuwe & Gautier-Hion, 1998).

The case of the forest buffalo and elephant is highly

significant because it indicates that elephants and buffalos



cannot be considered habitat generalists. The fact that forest

and savanna present vicariant taxa means that speciation, or at

least significant differentiation, has occurred. It is beyond the

scope of this article to discuss the history of speciation in the

African rain forests, but it is plausible that the large herbivores

presently living in African rain forests represent populations

once living in the open, that have been encircled by expanding

forest sometime in the past. These populations would have

become isolated from their savanna sisters and differentiated.

Analogously, it can be assumed that, while a fraction of the

large-sized savanna mammalian fauna of South America could

have vicariously inhabited South American rain forests, it

would still represent a minority of the taxa involved. However,

there is no evidence for the existence of such a large-sized

fauna in South American forests (Kay & Madden, 1997;

MacFadden & Shockey, 1997; Rancy, 1999; Cristoffer & Perez,

2003).

Cristoffer & Perez (2003) discuss the rain forest mammals of

South America and Africa, and indicate that forest lineages in

Africa have consistently evolved more terrestrial offshoots (in

the case of the African primate Cercopithecus). To this example

we could add the evolution of baboons (Papio) and our own

(Homo) (Reed, 1996). If African rain forests repeatedly

produced terrestrially adapted lineages, they are quite poor

in specialized arboreal locomotory adaptations. Prehensile

tails evolved independently at least six times in South

America: twice in Primates (Cebus and the ateline genera,

e.g. Brachyteles), rodents (Sphiggurus and Coendou), carnivores

(Potos), xenarthrans (Tamandua and Cyclopes) and marsupials

(all South American genera). In Africa, only pangolins (Manis)

possess prehensile tails. Emmons & Gentry (1983) showed that

African rain forests not only practically lack prehensile tailed

mammals, but also are very poor on gliding forms (only three

genera, Anomaluridae rodents), which abound in Southeastern

Asia. Emmons & Gentry (1983) attributed the evolution of

specialized locomotory adaptations in the world’s rain forests

to selective pressures derived from their differently structured

forest canopies and strata. It is not our aim to discuss the forest

patterns, but it is intriguing that South America has so many

independent acquisitions of prehensile tails, and Southeastern

Asia possesses several gliding vertebrates, while Africa shows a

remarkable poverty of both, and its forest lineages have

frequently evolved terrestrial offshoots.

Climate change and its influence on vegetation

Climate change has been evoked as playing an important role

in the megamammal Pleistocene extinctions (Graham &

Lundelius, 1984; Guilday, 1984; Ochsenius, 1985; Cartelle,

1999), but besides pointing out general consequences such as

drought, authors have refrained from building a model in

which tropical vegetation types would function under distinct

climatic regimes.

For the purpose of evaluating the impact of late Pleistocene

and Holocene climatic changes on the vegetation, we have

made the following assumptions.

The geographical distribution of vegetation types

We have considered climatic changes affecting the present

geographical configuration of major vegetation types in South

America and Africa. There is a considerable body of research

on botanical palaeocommunities which reveals that floristic

composition has changed in the past 20 Ka (Coetzee, 1993;

Houerou, 1997; Colinvaux et al., 2000). Despite the growing

palynological knowledge, these reconstructions, however,

remain local and would render impossible any analysis that

considers South America and Africa simultaneously with all

their distinct biomes. Besides, we know very little about past

relationships between extinct mammal lineages and particular

floristic elements beyond the general inferences about grazing

and/or browsing habits. Thus, our focus is on physiognomic

changes of vegetation rather than floristic (see below).

How average precipitation levels affect vegetation

physiognomy

Our second assumption is generally applicable to tropical

landscapes, in that plant formations tend to present denser

concentrations of woody elements with greater water availab-

ility and vice versa (Fig. 2; Furley & Newey, 1983; Walter,

1984; Rizzini, 1997; Furley, 1999). The density of woody

species directly reflects on the amount of sunlight reaching the

soil and thus affects grass biomass.

One way to demonstrate that density of woody elements in a

vegetation varies with water availability through changing

precipitation levels is to examine the literature on areas

monitored for the effects of fire on vegetation. This is, of

course, an indirect approach as the climate (precipitation level)

has not actually changed in these areas. We assume that a

severe dry period lasting for a large number of consecutive

years, as would be the case at the beginning of a glacial period,

enhances the openness of a landscape through fires, either

natural or man made, while lesser deficits diminish the impact

of natural fire, as attested by several sources (Walter, 1984;

Miranda et al., 2002; Oliveira-Filho & Ratter, 2002).

In Africa, a number of studies have focused on the vegetation

changes due to the effects of seasonal fire and herbivory on the

vegetation. Salvatori et al. (2001) and Mapaure & Campbell

(2002) have shown that fire and herbivory play a significant

role in the decrease of woodland area and its transformation in

grasslands, but these authors have not been able to properly

distinguish between the independent contributions of fire and

herbivores. However, Dublin et al. (1990) showed that decrease

in woodland cover was due primarily to fire, large herbivores

playing a part mainly in the lack of regeneration of woodland

after its eradication by fire. Swaine et al. (1992) were able to

monitor controlled areas and demonstrated that trees become

more abundant if savannas are protected from fire. These

authors showed that in protected areas of savanna close to

forest vegetation, the regeneration included an expansion of

forest species, while in areas away from forests, the savanna

trees increased their density.



In South America, there are no large wild herbivores as in

Africa, but the net results are essentially the same. Studies on

the density of tree cover in the Venezuelan (San Jose & Farinas,

1983, 1991) and Central Brazilian savannas (Moreira, 2000;

Hoffmann & Moreira, 2002) have shown that protection from

fire leads to denser physiognomies. McNaughton et al. (1993)

provide comparisons between vegetation structures of arid and

semi-arid regions of South America and Africa, concluding

that great general similarities can be observed, including the

response of open and dense savannas and woodland to fire.

Our model assumes that vegetation is much less vulnerable to

fire if precipitation levels are consistently higher, as would occur

during the prevalence of wetter climates and vice versa. The role

of soil in areas covered by savannas is, of course, essential, but

not as important in determining physiognomies than it is for

floristics. Following Oliveira-Filho & Ratter (2002), high fertility

unflooded soils will always sustain some kind of forest, from

semi-evergreen to dry. Low fertility unflooded soils will support

savanna vegetation, but if water availability is high and

seasonality not strongly marked, tall Cerrado (a kind of savanna

forest) and dense savanna occur; even semi-evergreen forests

can be found. Low fertility soils with strong water deficits will

sustain several kinds of open savanna.

The estimated effect of average water availability on

vegetation physiognomy is summarized in Fig. 2. This figure

presents two distinct paths of vegetation physiognomic change,

both commencing at levels of good water availability and

degrading to steppic or desertic conditions. The first series

ranges from rain forests and the second from dense arboreal

savanna. By keeping these two series separate we underline the

fact that savannas are soil dependent (Furley & Newey, 1983;

Sarmiento, 1984; Walter, 1984; Rizzini, 1997; Oliveira-Filho &

Ratter, 2002).

How general were physiognomic changes in vegetation

Any vegetation map for South America and Africa is a

simplified view of how the floras and physiognomies are

spatially distributed. A vegetation such as the ‘Amazonian rain

forest’ actually encloses what we understand as ‘typical’ rain

forest and many other subtypes and enclaves. This makes our

assumptions (1) and (2) not immediately applicable at regional

or local scales, but adequate at continental level. Thus, when

we generalize the alteration of a vegetation physiognomy from

‘evergreen forest’ to ‘semi-evergreen forest’ or ‘dry forest’, we

are not stating that the entire biome has changed uniformly in

these directions, but only that the indicated resulting physi-

ognomy would be the predominant one.

Our focus on the LGM and at the HCO makes distinctions

between the two periods straightforward: at the LGM the

prevalent climate over South America and Africa was drier,

while at the HCO it was wetter. We assume that the distinct

climatic regimes of the LGM and HCO prevailed over both

continents, acting over their entire range of different kinds of

vegetation cover.

The intensity and kind of the climatic change

We have considered a single variable in our model: the

precipitation levels. The rationale is the same as that we have

employed for the vegetation, i.e. we are aware that climate

change involves alteration in a vast number of variables, such

as temperature, aerial and marine circulation patterns, sea-

sonality and others. However, not only are there no available

climatic reconstructions at such detailed level for South

America and Africa simultaneously, but also if such recon-

structions were available, our data on the mammals would

Evergreen forest

Semi-evergreen forest

Dry forest

Dry forest and shrub

Steppe

Tall Cerrado Woodland

Savanna

Mixed grass and shrub

Grassland

Steppe

Figure 2 Vegetation type physiognomies

under increasingly drier and/or more mark-

edly seasonal climates (from top to bottom).

Elephants indicate those habitats suitable to

medium to large savanna mammals. We

believe that tall Cerrado, a kind of forested

savanna of Central Brazil and known as

‘Cerradao’ by South American botanists,

could not have supported large mammals

because it is denser and have less grasses than

typical African woodland.



certainly lag behind. We find that the best climatic variable

that can be reasonably studied with regard to its effect on

vegetation physiognomy is water availability, translated as

yearly average precipitation. As we cannot evaluate how much

the average precipitation would drop or increase, nor if

seasonal patterns would be maintained, we therefore have

assumed that under wetter and drier climates, all regions

would receive more or less rain by approximately the same

amount, thus affecting vegetation physiognomy in one direc-

tion. Finally, as we cannot know how much more or less

precipitation a continent received under a certain climatic

regime, we cannot ascertain whether a vegetation change

would be slight or else range across two or more of the

categories depicted in Fig. 2.

Below we briefly describe the present vegetation types of

both continents and the effect of climate change in the savanna

habitats and their mammalian faunas.

Present vegetation physiognomy in South America

and Africa

In South America the main areas covered today by evergreen

forest vegetation are the Amazonian rain forest (including, to

the west of the Andes, Colombia and northern Ecuador) and

the Atlantic rain forest of eastern Brazil (Fig. 3a). These forests

are mainly tropical, but a significant temperate rain forest

occurs in Chile, south of Valdivia. Between the Amazonian and

Atlantic tropical rain forests lies a vast strip of more or less

open vegetation types, extending across the continent. In

north-eastern Brazil is the Caatinga, a type classified as dry

forest and shrub (Eva et al., 2002). The central Brazilian

Cerrado is generally described as a savanna, but actually

includes savanna physiognomies and mesophytic forests as

well. In Bolivia, Paraguay and northern Argentina there is also

a dry forest and shrub, known as Chaco. An important area of

savannas occur in Colombia and Venezuela (the Llanos), and

two major enclaves of savannas can be found in northern

Brazil and adjacent Guyana (Gran Sabana). To the south,

Argentina has the extensive grasslands of the Pampas,

which gradually changes into steppe as we move south into

Patagonia. Just north of the Chilean temperate rain forest,

climates become ever more drier and the vegetation changes

accordingly: northwards from Valdivia, vegetation is gradually

altered into deciduous forest, shrub forest, steppe and desert

(Atacama). A large section of the Chaco is tropical to

temperate, and the Pampas and the Patagonian steppe are

entirely temperate. Thus, most of South America can be

described as consisting of three major areas of evergreen

forest – the Amazonian, the Atlantic, and the Chilean – with a

succession of more or less open vegetation types extending

from equatorial to temperate latitudes in between the first two

and northwards from the third (Hueck, 1966; Hueck & Seibert,

1981; Rizzini, 1997; Eva et al., 2002).

Africa presents a similar picture, albeit distinctively geo-

graphically organized (Fig. 3a). The main body of evergreen

forest is equatorial, occurring from the Atlantic coast to central

parts of Africa, at the Albertine Rift, with an eastern arc

sweeping through Mozambique, Tanzania and southern

Kenya. Savanna, and savanna and forest mosaics surround

this forest. To the north, increasingly seasonal and drier

climates predominate, constituting a transitional region

between the savannas and the Sahara, known as the Sahel;

the vegetation becomes ever more open till the desertic

Saharan landscape. To the east and south of the equatorial

rain forests the same general pattern of open formations can

be verified, but more important areas of woodland and

mixed savanna and woodland can be found, from central

Angola to Tanzania and Mozambique. To the south of this

vast ‘woodland’ area, climate becomes drier and vegetation

changes accordingly into the xeric regions of the Namibian

desert and the Kalahari. Forests or forest mosaics appear in

the south-eastern coast of Africa, in Mozambique, mirroring

the South American Atlantic forest. In summary, most of the

African open formations lie surrounding evergreen forests,

within regions dominated by tropical and subtropical

climates. Africa does not possess sizeable land within

temperate domains, but the moderately high areas of the

Ethiopian and East African plateaus include altitudinal

vegetation gradients analogous to those found in parts of

the South American Andes (Cooke, 1972; Dickson, 1992;

McMaster, 1992; Carroll, 2001).

Vegetation physiognomies in South America

and Africa in the late Pleistocene and Holocene

Accepting the assumptions described above, we may discuss

what changes would occur over the continents in terms of their

vegetation cover. Considering that, world-wide, the LGM of

the late Pleistocene was drier than the present (Flenley, 1979;

Whitmore & Prance, 1987; Clapperton, 1993; Colinvaux et al.,

2000; Whitlock et al., 2001), we associate our maximum

vegetational openness scenario with this phase.

Figure 3b shows our reconstruction of vegetation physiog-

nomies for both continents during the LGM. As can be seen,

subjected to lower precipitation averages, evergreen forests

would degrade to semi-evergreen and could as well present

more open patches of savanna-like physiognomies in a mosaic

pattern. This kind of change would occur along vast areas of

rain forest if precipitation dropped significantly. That this kind

of change took place in South America can be verified by the

fact that the Amazonia is mainly covered by rain forest, but

include hundreds of isolated patches of savanna (Hueck &

Seibert, 1981). Areas covered by semi-evergreen forests, dry

forests, woodlands and savannas would generally open up as

well. Forest would be mostly restricted to mountain ranges

subjected to orographic rain and to other areas where rainfall

would still be high enough. The retracting forests would leave

space for drier seral stages in the forest succession (as seen in

Fig. 2), and in areas with poor soils, savannas could be

established. Vegetation types that are presently subjected to

semi-arid and arid climates, such as deserts, South American

grasslands, steppes and the south African areas of mixed grass



and shrub would become even drier. Mountain ranges would

be affected somewhat differently, in that altitude and not only

latitude would play a part. Altitudinal zonation would

probably be found at correspondingly lower altitudes than

those found today.

As we enter the Holocene, climatic conditions changed.

Figure 3c shows our reconstruction of the prevailing vegetation

physiognomies in South America and Africa at the HCO. For

Africa, there is a well-established pattern of ever more humid

climate reaching the hypsithermal interval, or the HCO at the

middle Holocene, followed by a return to drier conditions in

the present (Coetzee, 1993; Houerou, 1997). During the HCO,

African rain forests would be under ideal humidity conditions,

probably expanding to tropical areas where soil could support

it. Open woodlands would likewise present denser vegetation.

Thus, open formations would be eliminated in certain areas or

reduced in others, but would actually expand into regions

previously too dry to support it, such as desertic areas. For

South America, the HCO is less consensual. Some authors

(Bradbury et al., 1981; Bigarella & de Andrade-Lima, 1982;

Hare, 1992; Joly et al., 1999; de Oliveira et al., 1999) propose

the same pattern as that just described for Africa, that is, the

HCO occurring from early to middle Holocene. However,

palynological studies throughout the continent resulted in

somewhat conflicting results in the sense that some regions

would have received more rainfall with others experiencing

desiccation at the same moment, even within the tropics

(Ledru et al., 1998; Salgado-Labouriau et al., 1998; Behling &

Hooghiemstra, 2001). The same can be said of researches

relying on different or broader data bases (Clapperton, 1993;

Colinvaux et al., 2000). Generally, they sustain that most of the

Holocene was drier than the present, gradually becoming ever

more humid in the last 5000 years, approximately – although

not uniformly – over the entire tropical South America.

Irrespective of a middle or late Holocene HCO for South

America, both Amazonian and Atlantic rain forests would be

under their best climatic conditions, while dry forests such as

those of Caatinga and Chaco would turn into semi-evergreen

forests. Tropical savannas could maintain their floristic

identity but its woody component would have become densely

packed, with marked reduction of grassland areas. Grasslands

and wetlands within tropical South America could probably

still be found, but predominant physiognomies would be dense

savanna and tall, forest savannas.

Figure 3d presents a summary of our model, showing that

areas within and without the 30� latitudinal belt would present

complementary and opposite characteristics during the LGM

and the HCO. A major distinction between South America and

Africa is established in that, during the HCO, the latter

continent would maintain large expanses of suitable habitats

for medium- and large-sized open vegetation mammals, both

to the north and to the south (the Sahara and the southern

xeric regions of Namibia and the Kalahari), whereas South

America would present its physiognomically suitable areas

only at the colder, temperate lower latitudes of the southern

part of the continent.

Mammal faunas and vegetation change

Changing climate effects on vegetation and mammal faunas is

a well-documented feature of the evolutionary history of

mammals. The Miocene, with its world-wide expansion of

grassland habitats is associated with the radiation of several

herbivore lineages both in South America and Africa; at the

end of the Miocene, when grasslands retreated, extinction

of part of these lineages followed suit, at least in South

America (Patterson & Pascual, 1972; Janis, 1993; Vrba, 1993;

MacFadden, 2000). We would expect the same pattern

occurring during the Pleistocene–Holocene climatic and

vegetation change of the magnitude proposed here. In

addition, we should expect the entire mammal fauna to be

affected, not only its larger-sized lineages. Roughly, faunistic

assemblages adapted to open vegetation environments should

either become extinct or isolated within pockets if this

vegetation is replaced by forest, and when open landscapes

replace forests, the same would happen with the forest-adapted

taxa. This is indeed the case, as we demonstrate below.

Pardinas et al. (2002) summarized several dramatic changes

in the composition of small mammal communities in southern

South America from the late Miocene to the Holocene. The

ecological implications of these changes in the taxonomic

structure of the communities is not entirely understood, but

these authors attributed most of the small mammal distribu-

tional changes to climate and its effect on vegetation. Evidence

that the Amazonian rain forest was widely penetrated by open

formations is provided by some studies on the distribution of

small mammals with pronounced habitat fidelity. The marsu-

pial Lutreolina occurs from the Buenos Aires province in

Argentina northwards until the Brazilian States of Sao Paulo,

Goias and Mato Grosso do Sul, always being associated with

open vegetation habitats. Its distribution is then interrupted by

the Amazonian rain forest, and the same species reappear in

the open areas of Venezuela and Colombia (Nowak, 1999).

Nunes (2001) studied small mammal communities within

several isolated savanna enclaves within the Amazon basin and

found that some of its rodents represented disjunct popula-

tions from species inhabiting the northern and central South

American savannas, which implies that these rodents had a

wider distribution in open habitats and survived in savanna

pockets when the forests returned. We have already mentioned

above that the presence of vicariant species such as the forest

elephant (L. cyclotis) and the forest buffalo (S. caffer nanus)

within the African rain forest may represent the same

phenomenon of forest retraction followed by expansion and

‘capture’ of open enclaves for that continent, although the

dates involved may be different. In the case of the South

American small mammals, most of the species did not speciate,

indicating that the disjunction events may be quite recent.

The study of extinct Amazonian mammals reinforces the

same scenario. Webb & Rancy (1996) and Rancy (1999)

describe in detail the Amazonian Pleistocene open formation

mammal fauna and concluded that there was a need for

savanna habitats to sustain them. However, Colinvaux et al.



(2000), working on western Amazonia Pleistocene palynolog-

ical record, sustained that most of the Amazon lowlands

remained as a forest throughout glacial cycles; proposing that

these extinct mammals actually lived in forests, and that all

extinct grazers would be restricted to river margins, where

grasses could be found. We agree with most authors (Webb &

Rancy, 1996; Rancy, 1999; Cristoffer & Perez, 2003) in that

Amazonian forest as it is today could not support the

megafauna. In our view the question of physiognomy vs.

floristics is crucial here. Arboreal or grass pollen do not furnish

an immediate picture of how densely or sparsely forested a

region would be; instead they reveal floristic assemblages. Thus,

for instance, arboreal pollen might be recovered from a site

sustaining a woodland or dry forest (see Pennington et al.,

2000), which is rather more open and more suitable to the

megafauna than a rain forest. Spatial organization of plants is

not immediately revealed by pollen lacustrine deposits, it is

only suggsted by it, as structural reconstruction depends on the

knowledge of our present-day floristic communities.

If the HCO resulted in the spread of closed, forest

environments over previously open landscapes, we should

expect that forest-adapted mammals would leave signals of

their presence in areas presently covered by open formations.

Voss & Myers (1991), Emmons & Vucetich (1998) and Silva

et al. (2003) documented that several species of small sigm-

odontine and histricognath rodents that are otherwise still

living in forest areas elsewhere have been found as ‘Pleistocene’

fossils in the central Brazilian site of Lagoa Santa, in the State

of Minas Gerais. This locality is today covered by savanna

vegetation, and most of the rodents listed by those authors are

30 N

30 S

EquatorEvergreen forest

Savanna

Savanna

Savanna

Dry forest Semi evergreenand evergreenforest

Evergreen forest

Grassland

Evergreenforest

Grassland / Steppe

Desert

DesertWoodland

Woodland

Mixed grass and shrub

Mosaic of open forestsand savannas

Mosaic of open forestsand savannas

Open savannas

Open savannas

Open savannas

Desert

Desert

Grassland / Steppe

Desert / Cold Steppe

Evergreen forestEvergreen forest

Evergreen andsemi evergreen forests

Evergreen semi evergforests

andreen

Savanna

Savanna

Dense savanna

Dense savanna

Dense savanna

Temperate formationsEvergreenforest

Suitable during HCOUnsuitable during LGM


Suitable during LGMUnsuitable during HCO

Suitable during LGMUnsuitable during HCO


(a)(b)

(c) (d)

Dry forest

Figure 3 Present day major plant formations of South America and Africa (a) and geographical vegetation change for the two continents at

the LGM (b) and the HCO (c). A summary of this change is depicted in (d), where darker grey are for areas of predominantly suitable

megafauna habitat at the LGM and unsuitable habitats at the HCO; and lighter grey are for areas predominantly suitable habitats at the HCO

and unsuitable at the LGM. White areas in the maps of South America mostly indicate complex Andean vegetation not considered here.



forest dwellers. Their disappearance from a savanna locality

and survival in forests elsewhere indicates that ecological

changes in vegetation distribution were probably responsible

for their local extinction. Cartelle & Hartwig (1996) described

two Pleistocene extinct folivorous monkeys from a fossil site

within the present-day Caatinga (dry forest and shrub). This

kind of vegetation cannot support highly herbivorous

monkeys, and thus they must have inhabited considerably

moister forests. de Vivo (1997) showed several cases of

disjunction between species of arboreal mammals occurring

in eastern Amazonian rain forest and north-eastern Brazilian

Atlantic rain forest. Between these two rain forests, there is the

Caatinga, which cannot presently support these species. These

disjunctions appear to be quite recent because no differentia-

tion can be detected for the populations involved.

In Africa, there is evidence for the expansion of savanna

mammal communities to areas covered today by desert.

Savanna invaded what is today the Saharan desert at the

Holocene HCO, as is beautifully attested by the rock art left by

humans from 8000 to 1000 yr bp. Wild savanna mammals,

such as giraffe, hippopotamus, elephant and several bovids

were depicted in hunting scenes, and gradually were replaced

in the paintings by cattle. Finally, at the end of the HCO even

cattle are no longer depicted and the Sahara took its present

arid configuration (Camps, 1974; Scarre, 1988; Houerou,

1997).

Thus, there is considerable evidence that open and forest-

adapted mammal faunas have changed their distribution

patterns in the past few thousand years, probably as a result

of change in the vegetation physiognomy. If these vegetation

changes have occurred as depicted in Fig. 3, the large-sized

South American mammal lineages, mostly adapted to open

formations, would have found reduced adequate landscapes in

tropical and subtropical regions during the HCO, with its

massive forest expansion. Most populations would become

extinct and the few isolated ones would be much more

vulnerable to accidental extinction or to human predation. In

Africa, the savanna-adapted mammals would still find vast

expanses of suitable habitats both in the subtropical north and

south parts of the continent, and the populations would be

under much less stress.

As glacial and interglacial cycles occurred throughout the

Quaternary, some of the faunistic patterns may have been

established earlier. However, there is evidence that the last

glacial and interglacial cycle was particularly intense and

abrupt relative to previous cycles, at least relative to the last

100 Ka (Hambrey & Harland, 1981; Haffer, 1982; Bartlein &

Prentice, 1989; Roy et al., 1996). All Quaternary glaciation

cycles supposedly had the same general effect on vegetation,

albeit with varying intensity. Thus, we think that most of the

previous Pleistocene extinctions could be attributed to the

alternating glacial and interglacial cycles, but the final coup

would have been administered by the last one.

It remains to be explained why the southern cone of South

America, which was subjected to colder and drier conditions at

the LGM, would not have supported most of the savanna

mammals at the Holocene HCO. The mammalian fossil record

tells some of the story. Cartelle (1999) compiled data from

several independent researchers of fossil Argentine mammals

of the late Pleistocene (LGM) which show that this temperate

fauna migrated to northern, more tropical localities at this

period, separated by severe cold and steppic environments.

(see also Ficcarelli et al., 1997; Coltorti et al., 1998; Nunez

et al., 2001). During the HCO, expanding savannas could have

become available again only in the temperate southern cone of

South America. Animals adapted to tropical conditions would

find it a harsh environment and thus only a fraction of the late

Pleistocene tropical savanna fauna would have returned. Our

hypothesis thus predicts that any Holocene assemblage of

mammals from the temperate southern cone will be poorer

than contemporary tropical ones. Most of the present day

dissimilarities between Africa and South America would then

be established at the Holocene.

A critique of previous hypotheses on the extinction

of the South American megafauna

Alternative hypotheses have been proposed, namely (1) the

action of human hunters entering the American continent and

overexploiting the fauna (Martin, 1967, 1984); (2) the

elimination of phyletic lineages due to a faunistic imbalance

caused by the invasion of North American mammals in South

America after the Great American Interchange of the late

Pliocene (May, 1978; Marshall et al., 1982; Webb, 1985); and

(3) the intense dry phase of the late Pleistocene (Ochsenius,

1985; Cartelle, 1999).

Martin (1967, 1984) proposed that humans entering the

Americas some 12 000 years ago overhunted a ‘naive’ mammal

fauna that, having never faced such resourceful predators,

provided easy and abundant meat. We reject it for several

reasons discussed below.

Martin (1984) proposed two possible features that would be

discordant to his model: (1) the extinction of small mammals

(not vulnerable to human impact) and (2) the extinction of

large mammals before prehistoric human arrival. Both phe-

nomena have been recorded for South America.

The extinction of small forest mammals in areas presently

covered by savannas has already been explored in the previous

section. Ficcarelli et al. (1997) and Coltorti et al. (1998)

described and dated sites in the Andes of Ecuador and in the

northern Peruvian coast where mastodons and giant sloths

became locally extinct before the arrival of man at the end of

the Pleistocene (between 16 670 and 12 350 yr bp), while for

areas in the Equatorial Andes more subjected to cold climates,

the extirpation of the megafauna dated from c. 20 980 yr bp.

Ficcarelli et al. (1997) cite other similar cases for the Argentine

plains. Nunez et al. (2001) also describe and date a site in

northern Chile where the arrival of humans found the

megafauna already gone, and archaeological sites reveal

remains of extant mammals (late Pleistocene, c. 10 800 yr

bp). Ficcarelli et al. (1997), Coltorti et al. (1998) and Nunez

et al. (2001) attributed the disappearance of large mammals



from these sites to climatic change (excessive cold and dryness

at the end of the Pleistocene).

Additionally, Martin (1984, p. 360) argues that if the

disappearance of large mammals through human agency was

‘truly swift and devastating’, few kill sites would have been

found. Indeed, no kill sites have been found in South America,

and the extended presence of large mammals in the mid-

Holocene (described above) suggests that the absence of such

sites is not favourable to Martin’s scenario. There are a few

archaeological sites where evidence of some large mammals

hunted by humans have been found (Taima-Taima, Venezuela,

Gruhn & Bryan, 1984; southern Patagonia, Markgraf, 1985;

north-eastern Brazil, Guerin et al., 1996; Tagua-Tagua and

Monteverde, Chile, Nunez et al., 2001). The analysis of these

publications clearly indicates that humans did hunt large

mammals in South America, but the sites can hardly be

described as ‘kill and processing sites’. The dating of mid-

Holocene large mammals in South America (summarized

earlier) far extends the presence of these animals in the

continent simultaneously with the presence of humans,

contradicting an extremely rapid rate of megamammal extir-

pation through excessive hunting. We believe that human

predation of large South American mammals did occur, but

that it may have played a marginal role, probably in the

elimination of small isolated populations.

Beck (1996) disagrees with Martin’s blitzkrieg hypothesis for

North America, based on the geographical and temporal

distribution of the megafaunal remains which do not fit the

direction and timing of human migration from north-west to

south-east North America, as should have happened under

Martin’s assumptions. Beck (1996) emphasizes that his data

does not preclude the possibility that changes in climate were a

more important factor.

Martin (1984) argues that the long coevolutionary history of

humans and the megafauna in Africa and southern Asia would

account for the survival of the animals in those regions. The

animals would have learned to recognize humans as threaten-

ing and developed avoidance behaviours. As large mammals

were present in the South American Holocene, the ‘overkill’

period would have to be extended from at least 12 000 to

c. 6000 yr bp (some authors claim earlier dates for human

entry in the American continent; see Rudgley, 1999). With

such long period of co-habitation, South America would not

have any ‘naive’ mammals left; they would all have learned to

recognize and avoid humans, which is exactly what large

mammals in Africa and Asia did.

A second hypothesis dealing with (but not only) the South

American mammal extinctions at the end of the Pleistocene is

related to predictions contained in the Equilibrium Theory

(MacArthur & Wilson, 1967), in which land masses are

believed to reach relatively stable numbers of lineages given

enough time. May (1978), Marshall et al. (1982), Webb (1985)

and Marshall & Cifelli (1990), applying the Equilibrium

Theory to the palaeontological history of New World mammal

faunas, generally considered it as a viable alternative explana-

tion for the South American extinctions. To them, the number

of taxa that resulted from the sum of North American invaders

plus the southern native mammals far exceeded the carrying

capacity of the South American continent after the Great

American Interchange at the end of the Pliocene. We reject this

hypothesis. If the Equilibrium Theory explains the South

American extinctions, we should expect a decrease in the

number of lineages after the Great American Interchange, and

that indeed occurred. However, it is not at all clear why these

extinctions were concentrated on larger mammals, as demon-

strated by Lessa et al. (1997). If an excessive number of

lineages is the factor behind extinctions, we should expect that

small, medium and large mammals would have been equally

affected, but that is not what happened as we associate larger

body sizes to the habitat preferences of the mammals (as

discussed above, larger mammals tending to live in open

formations), we believe that our hypothesis furnishes a better

explanation for the known facts.

Ochsenius (1985) and Cartelle (1999) proposed that the

South American megafauna extinctions occurred in the late

Pleistocene due to climatic change, relating it to the cold and

dry conditions of the LGM. We agree with both authors that

this indeed occurred, at least locally (Ficcarelli et al., 1997;

Coltorti et al., 1998; Nunez et al., 2001). Our hypothesis,

however, considers the presence in the mid-Holocene of some

lineages of large mammals, which was ignored by both authors.

The LGM climatic conditions cannot be assumed as the sole

explanation for the extinctions, and a new perspective must be

considered. The Holocene and its climatic events should be

taken into account.

CONCLUDING REMARKS

All characteristics related to the special savanna attributes of

herbivores and predators are largely dependent on the existence

of vast extensions of open spaces with abundant grasses. In

South America these spaces were severely reduced in tropical

and subtropical areas during the HCO. The present savannas of

Africa were probably affected in the same manner, but then

Africa had ‘escape’ areas in its deserts, north and south. These

deserts, under a more humid climate, would present savanna-

like physiognomies, thus sustaining entire savanna communi-

ties. The same climatic events could additionally explain the

contemporary existence of vicariant patterns in the African

savanna vs. distinct faunistic assemblages in the South

American open formations (tropical and temperate). In Africa,

the savannas invading former desertic regions, plus savanna

isolates inside the tropical areas of the continent, could sustain

isolated savanna mammal communities, which suffered differ-

entiation resulting in the current vicariant pattern. Evidently,

this could have occurred before, during the several glacial and

interglacial cycles of the Quaternary, not only in the last one,

thus reinforcing differentiation. However, South America

would have lost most of its tropical and subtropical savannas.

Grasslands in temperate South America, already somewhat

depopulated in the late Pleistocene (see above), did not recover.

What remained in the tropical and subtropical vs. the



temperate areas of the continent were the distinct faunistic

complements found today.

In conclusion, we believe that South America and Africa did

share similar faunistic patterns until quite recently (middle

Holocene) and that vegetation changes associated with fluctu-

ating rainfall levels physiognomically altered habitats available

for the open formation fauna. Present-day faunistic differences

between these continents would arise not because the events

affecting them were different, but because open formations

survived in far greater extensions in Africa than in South

America during the more humid phases of the Holocene.

ACKNOWLEDGMENTS

Erika Hingst-Zaher, Mario de Pinna, Eliana Marques Cancello,

Alexandre Reis Percequillo, Daniel Jelin and two anonymous

referees read earlier versions of this manuscript and offered

valuable criticism and suggestions. This research was suppor-

ted by FAPESP (grants 98/05075-7 and 00/06642-4).

REFERENCES

Anderson, E. (1984) Who’s who in the Pleistocene: a mam-

malian bestiary. Quaternary extinctions. A prehistoric revo-

lution (ed. by P.S. Martin and R.G. Klein), pp. 40–89. The

University of Arizona Press, Tucson, AZ.

Baffa, O., Brunetti, A., Karmann, I. & Neto, C.M.D. (2000)

ESR dating of a toxodon tooth from a Brazilian karstic cave.

Applied Radiation and Isotopes, 52, 1345–1349.

Bartlein, P.J. & Prentice, I.C. (1989) Orbital variations, climate,

and palaeoecology. Trends in Ecology and Evolution, 4, 195–

199.

Beck, M.W. (1996) On discerning the cause of late Pleistocene

megafaunal extinctions. Paleobiology, 22, 91–103.

Behling, H. & Hooghiemstra, H. (2001) Neotropical savannah

environments in space and time: late Quaternary inter-

hemispheric comparisons. Interhemispheric climate linkages

(ed. by V. Markgraf), pp. 307–323. Academic Press, San

Diego.

Bigalke, R.C. (1972) The contemporary mammal fauna of

Africa. Evolution, mammals, and southern continents (ed. by

A. Keast, F.C. Erk and B. Glass), pp. 141–194. State Uni-

versity of New York Press, Albany.

Bigarella, J.J. & de Andrade-Lima, D. (1982) Palaeoenviron-

mental changes in Brazil. Biological diversification in the

tropics (ed. by G.T. Prance), pp. 27–40. Columbia University

Press, New York.

Bourliere, F. (1973) The comparative ecology of rain forest

mammals in Africa and Tropical America: some introduc-

tory remarks. Tropical forest ecosystems in Africa and South

America: a comparative review (ed. by B.J. Meggers, E.S.

Ayensu and W.D. Duckworth), pp. 279–292. Smithsonian

Institution Press, Washington DC.

Bradbury, J.P., Leyden, B., Salgado-Labouriau, M., Lewis W.

M. Jr, Schubert, C., Binford, M.W., Frey, D.G., Whitehead,

D.R. & Weibezahn, F.H. (1981) Late Quaternary environ-

mental history of Lake Valencia, Venezuela. Science, 24,

1299–1305.

Camps, G. (1974) Les Civilisations Prehistoriques de l’Afrique du

Nord et du Sahara. Doin, Paris.

Carroll, A. (2001) Africa’s natural realms. National Geographic

Magazine, Map Supplement, 200 (3), September.

Cartelle, C. (1999) Pleistocene mammals of the Cerrado and

Caatinga of Brazil. Mammals of the Neotropics, the Central

Neotropics, vol.3, Ecuador, Peru, Bolivia, Brazil (ed. by J.F.

Eisenberg and K.H. Redford), pp. 27–46. The University of

Chicago Press, Chicago and London.

Cartelle, C. & Bohorquez, G.A. (1982) Eremotherium laurillardi

(Lund, 1842). 1. Determinacao especıfica e dimorfismo

sexual. Iheringia, Serie Geologica, 7, 45–63.

Cartelle, C. & Hartwig, W.C. (1996) A new extinct primate

from the Pleistocene megafauna of Bahia, Brazil. Proceedings

of the National Academy of Sciences, 93, 6405–6409.

Clapperton, C.M. (1993) Nature of environmental changes in

South America at the Last Glacial Maximum. Palaeogeog-

raphy, Palaeoclimatology, Palaeoecology, 101, 189–208.

Clutton-Brock, T.H. & Harvey, P.H. (1983) The functional

significance of variation in body size among mammals.

Advances in the study of mammalian behaviour (ed. by J.F.

Eisenberg and D.G. Kleiman), pp. 632–663. Special Publi-

cations no.7, The American Society of Mammalogists,

Pittsburgh, Pennsylvania.

Coetzee, J.A. (1993) African flora since terminal Jurassic. Biolo-

gical relationships between Africa and South America (ed. by

P. Goldblatt), pp. 37–61. Yale University Press, New Haven.

Colinvaux, P.A., de Oliveira, P.A. & Bush, M.B. (2000) Ama-

zonian and neotropical plant communities on glacial time-

scales: the failure of the aridity and refuge hypotheses.

Quaternary Science Reviews, 19, 141–169.

Coltorti, M., Ficcarelli, G., Jahren, H., Moreno Espinosa, M.,

Rook, L. & Torre, D. (1998) The last occurrence of Pleis-

tocene megafauna in the Ecuadorian Andes. Journal of South

American Earth Sciences, 11, 581–586.

Cooke, H.B.S. (1972) The fossil mammal fauna of Africa.

Evolution, mammals, and southern continents (ed. by A.

Keast, F.C. Erk and B. Glass), pp. 89–139. State University of

New York Press, Albany.

Cristoffer, C. & Perez, C.A. (2003) Elephants versus butterflies:

the ecological role of large herbivores in the evolutionary

history of two tropical worlds. Journal of Biogeography, 30,

1357–1380.

Dickson, K.B. (1992) The land. Africa, The New Encyclopaedia

Britannica, 13, 45–51.

Dublin, H.T., Sinclair, A.R.E. & McGlade, J. (1990) Elephants

and fire as causes of multiple stable states in the Serengeti-

Mara woodlands. Journal of Animal Ecology, 59, 1147–1164.

Eisenberg, J.F. & Redford, K.H. (1999) Mammals of the Neo-

tropics, the Central Neotropics, vol.3, Ecuador, Peru, Bolivia,

Brazil. The University of Chicago Press, Chicago and London.

Emmons, L.H. & Gentry, A.H. (1983) Tropical forest structure

and the distribution of gliding and prehensile-tailed verte-

brates. The American Naturalist, 121, 513–524.



Emmons, L.H. & Vucetich, M.G. (1998) The identity of

Winge’s Lasiuromys villosus and the description of a new

genus of echimyid rodent (Rodentia: Echimyidae). American

Museum Novitates, 3223, 1–12.

Eva, H.D., de Miranda, E.E., Di Bella, C.M., Gond, V., Huber,

O., Sgrenzaroli, M., Jones, S., Coutinho, A., Dorado, A.,

Guimaraes, M., Elvidge, C., Achard, F., Belward, A.S.,

Bartholome, E., Baraldi, A., de Grandi, G., Vogt, P., Fritz, S.

& Hartley, A. (2002) A vegetation map of South America.

Official Publications of the European Communities,

Luxembourg.

Faure, M., Guerin, C. & Parenti, F. (1999) The Holocene

megafauna from the Toca do Serrote do Artur (Sao Rai-

mundo Nonato archaeological area, Piauı, Brazil). Comptes

Rendus de l’Academie des Sciences, serie II fascicule A- Sci-

ences de la Terre et des Planetes, 329, 443–448.

Ficcarelli, G., Azzaroli, A., Bertini, A., Coltorti, M., Mazza, P.,

Mezzabotta, C., Moreno Espinosa, M., Rook, L. & Torre, D.

(1997) Hypothesis on the cause of extinctions of the South

American mastodons. Journal of South American Earth Sci-

ences, 10, 29–38.

Flenley, J.R. (1979) The equatorial rain forest: a geological his-

tory. Butterworths, London.

Furley, P.A. (1999) The nature and diversity of neotropical

savanna vegetation with particular reference to the Brazilian

cerrados. Global Ecology and Biogeography, 8, 223–241.

Furley, P.A. & Newey, W.W. (1983) Geography of the biosphere.

An introduction to the nature, distribution and evolution of

the world’s life zones. Butterworths, London.

Graham, R.W. & Lundelius, E.L., Jr (1984) Coevolutionary

disequilibrium and Pleistocene extinctions. Quaternary

extinctions. A prehistoric revolution (ed. by P.S. Martin and

R.G. Klein), pp. 223–249. The University of Arizona Press,

Tucson, AZ.

Gruhn, R. & Bryan, A.L. (1984) The record of Pleistocene

megafaunal extinctions at Taima-taima, northern Venezue-

la. Quaternary extinctions. A prehistoric revolution (ed. by

P.S. Martin and R.G. Klein), pp. 128–137. The University of

Arizona Press, Tucson, AZ.

Guerin, C., Curvello, M.A., Faure, M., Hugueney, M. &

Mourer-Chauvire, M. (1996) The Pleistocene fauna of Piauı

(northeastern Brazil): palaeoecological and biochronological

implications. Revista da Fundacao Museu do Homem

Americano, 1, 55–103.

Guilday, J.E. (1984) Pleistocene extinction and environmental

change: case study of the Appalachians. Quaternary extinc-

tions. A prehistoric revolution (ed. by P.S. Martin and R.G.

Klein), pp. 250–258. The University of Arizona Press, Tuc-

son, AZ.

Haffer, J. (1982) General aspects of the refuge theory. Biological

diversification in the tropics (ed. by G.T. Prance), pp. 6–24.

Columbia University Press, New York.

Haltenorth, T. & Diller, H. (1977) Saugetiere Afrikas und

Madagaskars. BLV Verlagsgesellschaft mbH, Munchen.

Hambrey, M.J. & Harland, W.B. (1981) The evolution

of climates. The evolving earth (ed. by L.R.M. Cocks),

pp. 137–152. British Museum of Natural History and

Cambridge University Press, London and Cambridge.

Hare, F.K. (1992) Climate and weather. Climate change. The

new encyclopaedia Britannica, Vol. 16, 15th edn (ed. by R.

McHenry), pp. 484–497. Encyclopaedia Britannica, Inc.,

Chicago.

Hershkovitz, P. (1972) The recent mammals of the Neotropical

region: a zoogeographic and ecological review. Evolution,

mammals, and southern continents (ed. by A. Keast, F.C. Erk

and B. Glass), pp. 311–431. State University of New York

Press, Albany.

Hoffmann, W.A. & Moreira, A.G. (2002) The role of fire in

population dynamics of woody plants. The Cerrados of

Brazil (ed. by P.S. Oliveira and R.J. Marquis), pp. 159–177.


Houerou, H.N. (1997) Climate, flora and fauna changes in the

Sahara over the past 500 million years. Journal of Arid

Environments, 37, 619–647.

Hueck, K. (1966) Die Waelder Sudamerikas. Gustav Fischer

Verlag, Stuttgart.

Hueck, K. & Seibert, P. (1981) Vegetationskarte von Suda-

merika. Gustav Fischer Verlag, Stuttgart.

Janis, C.M. (1993) Tertiary mammal evolution in the context

of changing climates, vegetation, and tectonic events.

Annual Review of Ecology and Systematics, 24, 467–500.

Joly, C.A., Aidar, M.P.M., Klink, C.A., McGrath, D.G.,

Moreira, A.G., Moutinho, P., Nepstad, D.C., Oliveira, A.A.,

Pott, A., Rodal, M.J.N. & Sampaio, E.V.S.B. (1999) Evolu-

tion of the Brazilian phytogeography classification systems:

implications for biodiversity conservation. Ciencia e Cultura

Journal of the Brazilian Association for the Advancement of

Science, 51, 331–348.

Kappelman, J., Plummer, T., Bishop, L., Duncan, A. &

Appleton, S. (1997) Bovids as indicators of Plio-Pleistocene

paleoenvironments in East Africa. Journal of Human Evo-

lution, 32, 229–256.

Kay, R.F. & Madden, R.H. (1997) Mammals and rainfall:

paleoecology of the middle Miocene at La Venta (Colombia,

South America). Journal of Human Evolution, 32, 161–199.

Keast, A. (1972) Comparisons of contemporary mammal

faunas of southern continents. Evolution, mammals, and

southern continents (ed. by A. Keast, F.C. Erk and B. Glass),

pp. 433–501. State University of New York Press, Albany.

Kingdom, J. (1977) East African mammals. An atlas of evolution

in Africa, vol. III part A (carnivores). The University of

Chicago Press, Chicago.

Kingdom, J. (1979) East African mammals. An atlas of evolution

in Africa, vol. III part B (large mammals). The University of

Chicago Press, Chicago.

Ledru, M.-P., Salgado-Labouriau, M.L. & Lorscheitter, M.L.

(1998) Vegetation dynamics in southern and central Brazil

during the last 10,000 yr BP. Review of Palaeobotany and

Palynology, 99, 131–142.

Lessa, E.P., Van Valkenburgh, B. & Farina, R.A. (1997)

Testing hypotheses of differential mammalian extinctions

subsequent to the Great American Biotic Interchange.



Palaeogeography, Palaeoclimatology, Palaeoecology, 135,

157–162.

MacArthur, P.H. & Wilson, E.O. (1967) The theory of island

biogeography. Princeton University Press, Princeton, New

Jersey.

MacFadden, B.J. (2000) Cenozoic mammalian herbivores from

the Americas: reconstructing ancient diets and terrestrial

communities. Annual Review of Ecology and Systematics, 31,

33–59.

MacFadden, B.J. & Shockey, B.J. (1997) Ancient feeding

ecology and niche differentiation of Pleistocene mammalian

herbivores from Tarija, Bolivia: morphological and isotopic

evidence. Paleobiology, 23, 77–100.

McMaster, D.N. (1992) Animal life. Africa. The New Encyclo-

paedia Britannica, 13, 56–59.

McNaughton, S.J. & Georgiadis, N.J. (1986) Ecology of African

grazing and browsing mammals. Annual Review of Ecology

and Systematics, 17, 39–65.

McNaughton, S.J., Sala, O.E. & Oesterheld, M. (1993) Com-

parative ecology of African and South American arid to

subhumid ecosystems. Biological relationships between Africa

and South America (ed. by P. Goldblatt), pp. 548–567. Yale

University Press, New Haven.

Maglio, V.J. & Cooke, H.B.S. (1978) Evolution of African

mammals. Harvard University Press, Cambridge.

Mapaure, I.N. & Campbell, B.M. (2002) Changes in miombo

woodland cover in and around Sengwa Wildlife Research

Area, Zimbabwe, in relation to elephants and fire. African

Journal of Ecology, 40, 212–219.

Markgraf, V. (1985) Late Pleistocene faunal extinctions in

southern Patagonia. Science, 228, 1110–1112.

Marshall, L.G. & Cifelli, R.L. (1990) Analysis of changing

diversity patterns in Cenozoic land mammal age faunas,

South America. Palaeovertebrata, 19, 169–210.

Marshall, L.G., Webb, S.D., Sepkoshi, J.J. & Raup, D.M. (1982)

Mammalian evolution and the great American interchange.

Science, 215, 1351–1357.

Martin, P.S. (1967) Prehistoric overkill. Pleistocene extinctions:

the search for a cause (ed. by P.S. Martin and H.E. Wright

Jr), pp. 75–120. Yale University Press, New Haven.

Martin, P.S. (1984) Prehistoric overkill: the global model.

Quaternary extinctions. A prehistoric revolution (ed. by P.S.

Martin and R.G. Klein), pp. 354–403. The University of

Arizona Press, Tucson, AZ.

Martin, P.S. & Klein, R.G. (1984) Quaternary extinctions. A

prehistoric revolution. The University of Arizona Press,

Tucson, AZ.

May, R.M. (1978) The evolution of ecological systems. Scien-

tific American, 239, 118–133.

Miranda, H.S., Bustamante, M.M.C. & Miranda, A.C. (2002)

The fire factor. The Cerrados of Brazil (ed. by P.S. Oliveira

and R.J. Marquis), pp. 51–68. Columbia University Press,

New York.

Moreira, A.G. (2000) Effects of fire protection on savanna

structure in Central Brazil. Journal of Biogeography, 27,

1021–1029.

Nowak, R.M. (1999) Walker’s mammals of the world, Vol.

I/II. Johns Hopkins University Press, Baltimore and

London.

Nunes, A. (2001) Gradientes estruturais dos habitats em savanas

amazonicas: implicacoes sobre a distribuicao e ocorrencia das

especies de pequenos mamıferos terrestres (Rodentia, Didel-

phimorphia). Doctorate Thesis, Universidade Federal do Rio

de Janeiro, Rio de Janeiro.

Nunez, L., Grosjean, M. & Cartajena, I. (2001) Human

dimensions of late Pleistocene/Holocene arid events in

southern South America. Interhemispheric climate linkages

(ed. by V. Markgraf), pp. 105–117. Academic Press, San

Diego.

Ochsenius, C. (1985) Late Pleistocene aridity in the Neotropic

as extinction cause of the South American Landmegafauna.

Zentralblatt fuer Geologie und Palaeontologie Teil I Allgeme-

ine Angewandte Regionale und Historische Geologie, H.11/12,

1691–1699.

de Oliveira, P.E., Barreto, A.M.F. & Suguio, K. (1999) Late

Pleistocene/Holocene climatic and vegetational history of

the Brazilian Caatinga: the fossil dunes of the middle Sao

Francisco River. Palaeogeography, Palaeoclimatology,

Palaeoecology, 152, 319–337.

Oliveira-Filho, A.T. & Ratter, J.A. (2002) Vegetation physiog-

nomies and woody flora of the Cerrado biome. The Cerrados

of Brazil (ed. by P.S. Oliveira and R.J. Marquis), pp. 91–120.


Owen-Smith, R.N. (1988) Megaherbivores – the influence of

very large body size on ecology. Cambridge studies in ecology.

Cambridge University Press, Cambridge.

Pardinas, U.F.J., D’Elıa, G. & Ortiz, P.E. (2002) Sigmodontinos

fosiles (Rodentia, Muroidea, Sigmodontinae) de America

del Sur: estado actual de su conocimiento y prospectiva.

Mastozoologia Neotropical, 9, 209–252.

Patterson, B. & Pascual, R. (1972) The fossil mammal fauna of

South America. Evolution, mammals, and southern continents

(ed. by A. Keast, F.C. Erk and B. Glass), pp. 247–309. State

University of New York Press, Albany.

Pennington, R.T., Prado, D.E. & Pendry, C.A. (2000)

Neotropical seasonally dry forests and Quaternary vegeta-

tion changes. Journal of Biogeography, 27, 261–273.

Peters, R.H. (1983) The ecological implications of body size.

Cambridge studies in ecology. Cambridge University Press,

Cambridge.

Rancy, A. (1999) Fossil mammals of the Amazon as a portrait

of a Pleistocene environment. Mammals of the Neotropics,

the Central Neotropics, vol. 3, Ecuador, Peru, Bolivia, Brazil

(ed. by J.F. Eisenberg and K.H. Redford), pp. 20–26. The

University of Chicago Press, Chicago and London.

Redford, K.H. & Eisenberg, J.F. (1992) Mammals of the Neo-

tropics, the Southern Cone, vol. 2, Chile, Argentina, Uruguay,

Paraguay. The University of Chicago Press, Chicago and

London.

Reed, K.E. (1996) Early hominid evolution and ecological

change through the African Plio-Pleistocene. Journal of

Human Evolution, 32, 289–322.



Reig, O.A. (1981) Teoria del origen y desarolo de la fauna de

mamıferos de America del Sur. Monographiae Naturae, 1,

1–162.

Rizzini, C.T. (1997) Tratado de Fitogeografia do Brasil. Aspectos

Ecologicos, Sociologicos e Florısticos. Ambito Cultural Edicoes,

Rio de Janeiro.

Roca, A.L., Georgiadis, N., Pecon Slattery, J. & O’Brien, J.

(2001) Genetic evidence for two species of elephant in

Africa. Science, 293, 1473–1477.

Roy, K., Valentine, J.W., Jablonski, D. & Kidwell, S.M. (1996)

Scales of climatic variability and time averaging in Pleisto-

cene biotas: implications for ecology and evolution. Trends

in Ecology and Evolution, 11, 458–463.

Rudgley, R. (1999) The lost civilisations of the Stone Age. The

Free Press, New York, NY.

Salgado-Labouriau, M.L., Barberi, M., Ferraz-Vicentini, K.R. &

Parizzi, M.G. (1998) A dry climatic event during the late

Quaternary of tropical Brazil. Review of Palaeobotany and

Palynology, 99, 115–129.

Salvatori, V., Egunyu, F., Skidmore, A.K., de Leeuw, J. & van

Gils, H.A.M. (2001) The effects of fire and grazing pressure

on vegetation cover and small mammal populations in the

Maasai Mara National Reserve. African Journal of Ecology,

39, 200–204.

San Jose, J.J. & Farinas, M.R. (1983) Changes in tree density

and species composition in a protected Trachypogonsavanna,

Venezuela. Ecology, 64, 447–453.

San Jose, J.J. & Farinas, M.R. (1991) Changes in tree density

and species composition in a protected Trachypogon savanna

protected for 25 years. Acta Oecologica, 12, 237–247.

Sarmiento, G. (1984) The ecology of Neotropical savannas.

Harvard University Press, Cambridge.

Scarre, C. (1988) Past worlds. The Times atlas of archaeology.

Times Books, London.

Silva, C.R., Percequillo, A.R., Iack-Ximenes, G.E. & de Vivo,

M. (2003) New distributional records of Blarinomys brevi-

ceps (Winge, 1888) (Sigmodontinae, Rodentia). Mammalia,

67, 147–152.

Simpson, G.G. (1980) Splendid isolation. The curious history of

South American mammals. Yale University Press, New

Haven.

Swaine, M.D., Hawthorne, W.D. & Orgle, T.K. (1992) The

effects of fire exclusion on savanna vegetation at Kpong,

Ghana. Biotropica, 24, 166–172.

Trajano, E. & de Vivo, M. (1991) Desmodus draculae Morgan,

Linares and Ray, 1988, reported for southeastern Brazil, with

palaeoecological comments (Phyllostomidae, Desmodonti-

nae). Mammalia, 55, 456–459.

Vanleeuwe, H. & Gautier-Hion, A. (1998) Forest elephant

paths and movements at the Odzala National Park, Congo:

the role of clearings and Marantaceae forests. African Journal

of Ecology, 36, 174–182.

de Vivo, M. (1997) Mammalian evidence of historical ecological

change in the Caatinga semiarid vegetation of northeastern

Brazil. Journal of Comparative Biology, 2, 65–73.

Voss, R.S. & Myers, P. (1991) Pseudoryzomys simplex

(Rodentia: Muridae) and the significance of Lund’s collec-

tions from the caves of Lagoa Santa, Brazil. Bulletin of the

American Museum of Natural History, 206, 414–432.

Vrba, E.S. (1980) The significance of bovid remains as indi-

cators of environment and predation patterns. Fossils in the

making (ed. by A.K. Behrensmeyer and A.P. Hill), pp. 247–

271. University of Chicago Press, Chicago.

Vrba, E.S. (1993) Mammal evolution in the African Neogene

and a new look at the great American interchange. Biological

relationships between Africa and South America (ed. by

P. Goldblatt), pp. 393–432. Yale University Press, New

Haven.

Walter, H. (1984) Vegetation und Klimazonen – Grundriss der

globalen Okologie. Eugen Ulmer GmbH & Co., Stuttgart.

Webb, S.D. (1985) Late Cenozoic dispersals between the

Americas. The Great American Interchange (ed. by F.G.

Stehli and S.D. Webb), pp. 357–386. Plenum, New York.

Webb, S.D. (1999) Isolation and interchange: a deep history

of South American mammals. Mammals of the Neotropics,

the Central Neotropics, vol. 3, Ecuador, Peru, Bolivia, Brazil

(ed. by J.F. Eisenberg and K.H. Redford), pp. 13–19. The

University of Chicago Press, Chicago and London.

Webb, S.D. & Rancy, A. (1996) Late Cenozoic evolution of the

Neotropical mammal fauna. Evolution and environments in

tropical America (ed. by J. Jackson, N. Budd and A. Coates),

pp. 335–358. University of Chicago Press, Chicago.

Whitlock, C., Bartlein, P.J., Markgraf, V. & Ashworth, A.C.

(2001) The midlatitudes of North and South America dur-

ing the Last Glacial Maximum and Early Holocene: similar

paleoclimatic sequences despite differing large-scale con-

trols. Interhemispheric climate linkages (ed. by V. Markgraf),

pp. 391–416. Academic Press, San Diego.

Whitmore, T.C. & Prance, G.T. (1987) Biogeography and Qua-

ternary history in tropical America. Clarendon Press, Oxford.

Wilson, D.E. & Reeder, D.M. (1993) Mammal species of the

world. A taxonomic and geographic reference. Smithsonian

Institution Press and American Association of Mammalo-

gists, Washington and London.

BIOSKETCHES

Mario de Vivo is Chair of Vertebrates and Curator of

Mammals and Birds at the Museu de Zoologia, Universidade

de Sao Paulo. His main research interests are the systematics of

South American mammals and their biogeography.

Ana Paula Carmignotto is in the PhD Program of Zoology

at the Instituto de Biociencias, Universidade de Sao Paulo,

currently developing her thesis research at the Museu de

Zoologia, USP. Her interests are ecology and biogeography of

open formation small mammals.



holocene vegetation change and the mammal …...south america in the late pleistocene and holocene...

Documents