Cladistics

Cladistics 30 (2014) 202214

10.1111/cla.12039

Cladistic biogeography of the Neotropical region: identifying the

main events in the diversification of the terrestrial biota
Juan J. Morrone*
Departamento de Biolog
a Evolutiva, Museo de Zoolog
a Alfonso L. Herrera, Facultad de Ciencias, Universidad Nacional Aut

onoma de M

exico
(UNAM), Mexico, DF, Mexico
Accepted 29 April 2013

Abstract
A cladistic biogeographical analysis was undertaken to identify the main events in the biotic diversification of the terrestrial
Neotropical biota. For the 36 animal and plant taxa analysed, a component 9 area matrix was constructed, associating geographical data only with informative nodes, and it was analysed under implied weights using the software TNT. The general
area cladogram obtained shows that the Neotropical region constitutes a monophyletic unit, with a first split separating the
Antilles and a second one dividing the continental areas into a north-western and a south-eastern component. Within the northwestern component the areas split following the sequence northern Amazonia, south-western Amazonia, north-western South
America, and Mesoamerica. Within the south-eastern component the areas split following the sequence south-eastern Amazonia,
Chaco, and Parana. The three main components are treated as subregions: Antillean, Amazonian (northern Amazonian, southwestern Amazonian, Mesoamerican, and north-western South American dominions), and Chacoan (south-eastern Amazonian,
Chacoan, and Parana dominions). Dispersal and vicariant events postulated to explain these pattens might have occurred during
the Cretaceous, when the Caribbean plate collided with the Americas, a combination of eustatic sea-level changes and tectonic
deformations of the continental platform exposed large parts of South America to episodes of marine transgressions, and the
Andean uplift reconfigured the Amazonian area. Tertiary and Quaternary events are assumed to have later induced the diversification within these large biogeographical units.
The Willi Hennig Society 2013.

The tropics of the Americas are well known for their

remarkable biodiversity, which is due to habitat heterogeneity and a complex geological history, both being
responsible for the patterns of geographical distribution of species and clades. Forests are among the most
common Neotropical biomes, particularly the Amazon
forest, but there are also extensive open biomes, e.g.
the diagonal of South America comprising the Pampa,
Chaco, Cerrado, and Caatinga. One of the first
hypotheses to account for the diversification of the
Amazonian terrestrial biota was provided by Wallace
(1852), who considered that the rivers from the Amazon basin have acted in the past as barriers to dispersal. An alternative explanation, the Pleistocene
*Corresponding author:
E-mail address: juanmorrone2001@yahoo.com.mx
The Willi Hennig Society 2013

refugia theory, postulated that the Amazonian forest

was fragmented through Pleistocenic climatic changes,
resulting in an archipelago of patches or refuges (Haffer, 1969, 1974, 1997; Vanzolini and Williams, 1970;
Prance, 1982; Lourenco, 1986). Some authors have
since postulated that Pleistocene climatic changes have
not been arid enough to fragment the forest and that
vicariance was caused by the formation of islands in
elevated areas (Colinvaux et al., 1996, 2000; Colinvaux, 1997, 1998; Colinvaux and Oliveira, 1999) and
others even invalidated the existence of refugia by considering them to be sampling artefacts (Nelson et al.,
2009). Other authors postulated that more ancient,
pre-Quaternary vicariant events explained general patterns of distribution of Amazonian taxa (Croizat,
1958, 1976; Cracraft and Prum, 1988; Bush, 1994; Patton et al., 2000; Amorim, 2001; Nihei and Carvalho,

J. J. Morrone / Cladistics 30 (2014) 202214

2004, 2007; Jaramillo et al., 2006; Rull, 2008; Hoorn

et al., 2010). The open biomes have been also analysed
recently (Ramos and Melo, 2010; Werneck, 2011), but
their integration with analyses of the areas with forest
biomes is rare (e.g. Sigrist and Carvalho, 2009; Pires
and Marinoni, 2010; Costa, 2003).
The formal definition of the Neotropical region
began in the 19th century. De Candolle (1820) did not
recognize the Neotropical region, but he listed five
regions that correspond to it: Mexico, tropical America, Chile, southern Brazil and Argentina, and Tierra
del Fuego. Sclater (1858), based on bird taxa, divided
the world into six zoogeographical regions; and Wallace (1876) later accepted this scheme, applying it to
other vertebrate taxa. According to the SclaterWallace system, the Neotropical region comprises South
and Central America, reaching as far north as central
Mexico. This scheme has been largely accepted ever
since, especially by zoogeographers working with vertebrates (Cox, 2001). Several biogeographers working
with plants or with invertebrates, however, have
adopted a more restrictive definition of the Neotropical region, excluding the southern portion of South
America, because of its closest links to Australia, New
Guinea, and New Zealand (Engler, 1879, 1882; Drude,
1884; Gill, 1885; Allen, 1892; Lydekker, 1896; Diels,
1908; Monr

os, 1958; Kuschel, 1964; Cabrera and Willink, 1973; Good, 1974; Takhtajan, 1988; Amorim and
Tozoni, 1994;
Morrone, 2002a; Moreira-Mu~
noz,
2007). In this restricted sense, the Neotropical region
corresponds to the tropics of the New World, i.e.,
most of South America, Central America, southern
Mexico, the West Indies, and southern Florida (Morrone, 2006; p. 477), explicitly excluding the Andean
area of South America, which is assigned to the
Andean region, and northern Mexico, which is
assigned to the Nearctic region. The Mexican Transition zone represents the overlap between the Nearctic
and Neotropical regions, whereas the South American
Transition zone represents the overlap between the
Neotropical and Andean regions (Morrone, 2006,
2010a). If both transition zones are included within it,
we have the Neotropical region sensu lato, and if they
are not included, the Neotropical region sensu stricto
(Fig. 1).
Regarding the biogeographical regionalization of
the Neotropical region, there have been several proposals recognizing subregions or dominions within it
(S

anchez Os
es and P
erez-Hern

andez, 1998; Morrone,
2010b). Wallace (1876) identified four subregions:
Mexican (southern Mexico and Central America),
Antillean (West Indies), Brazilian (tropical South
America), and Chilean (southern or temperate South
America). Cabrera and Willink (1973) recognized five
dominions: Caribbean (Mexico and the Antilles),
Amazonian, Guyanan, Chacoan, and Andean-Patago-

203

Fig. 1. The Neotropical region, with the Mexican and South American transition zones represented by dashed lines. The Neotropical
region sensu stricto does not include these transition zones, whereas
the Neotropical region sensu lato encompasses them.

nian. For South America, several authors have recognized two subregions: one named Guyano-Brazilian
or Brazilian, and the other named Andean-Patagonian, Patagonian, Argentinean, Chilean, or Austral
(Cabrera and Yepes, 1940; Ringuelet, 1961, 1975; Fittkau, 1969; Hershkovitz, 1969; Kuschel, 1969; Sick,
1969; Smith, 1983; Rivas-Mart
nez and Navarro,
1994; Almir

on et al., 1997; Kreft and Jetz, 2010;
Proches and Ramdhani, 2012). This main division
within South America has been also evidenced by
ecogeographical (Bailey, 1998) and macroecological
analyses (Ruggiero et al., 1998; Ruggiero and
Ezcurra, 2003). Morrone (2001) proposed a classification of the Neotropical region, based on panbiogeographical analyses, dividing it into four subregions:
Caribbean, Amazonian, Chacoan, and Parana. Morrone and Coscar

on (1996) undertook a parsimony
analysis of endemicity based on South American Peiratinae (Heteroptera: Reduviidae), hypothesizing that
the gradual development of an open vegetation diagonalcomprising the Chaco, Cerrado, and Caatinga
separated a former forest into a north-western part
(north-western South America and Amazonian forest)
and a south-eastern part (Parana and Atlantic

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J. J. Morrone / Cladistics 30 (2014) 202214

forests). This hypothesis was corroborated by a cladistic biogeographical analysis (Morrone and Coscar

on, 1998). Other cladistic biogeographical analyses
(Amorim, 2001; Nihei and Carvalho, 2007; Sigrist
and Carvalho, 2009; Pires and Marinoni, 2010;
Echeverry and Morrone, in press) have questioned
the naturalness of the Amazonian and Caribbean
subregions of Morrone (2006).
My objective is to analyse the relationships of different areas of the Neotropical region to test its naturalness, revise its regionalization, and identify the main
events in the biotic diversification of the terrestrial
Neotropical biota.

Material and methods

Areas
I analysed eight areas (Fig. 2), which have been
identified as biogeographical units in previous studies
(Rapoport, 1968; Savage, 1982; Amorim and Pires,
1996; Amorim, 2001; Morrone, 2001, 2006 Nihei and
Carvalho, 2007; Ramos and Melo, 2010; Echeverry
and Morrone, in press). The Caribbean and Amazonian subregions, as previously recognized (Morrone,
2006), were divided into three smaller units each to
test their naturalness. The eight areas analysed are as
follows:
1 Mesoamerica: lowlands of southern and central
Mexico and most of Central America (Guatemala,
Belize, Honduras, El Salvador, and northern Nicaragua).
2 Antilles: West Indies (Bahamas, Greater Antilles,
and Lesser Antilles).
3 North-western South America: western Ecuador
and Colombia, north-western Venezuela, Panama,
Costa Rica, and southern Nicaragua.
4 Northern Amazonia: Amazonian forest, basically
north of the Amazon river.
5 South-eastern Amazonia: Amazonian forest,
south-east of the Amazon river.
6 South-western Amazonia: Amazonian forest,
south-west of the Amazon river.
7 Chaco: open vegetation areas in northern and
central Argentina, southern Bolivia, western and central Paraguay, Uruguay, and central and north-eastern
Brazil.
8 Parana: forests from north-eastern Argentina,
eastern Paraguay, southern Brazil (west of the Serra
do Mar and toward central Rio Grande do Sul), and
eastern Brazil.
Additionally, four external areas were included to
test the naturalness of the Neotropical region: the
Nearctic and Andean regions, and the Mexican and
South American transition zones (Morrone, 2006).

Fig. 2. Units of the analysis. (a) Nearctic region; (b) Mexican Transition Zone; (c) Mesoamerica; (d) Antilles; (e) north-western South
America; (f) northern Amazonia; (g) south-eastern Amazonia; (h)
south-western Amazonia; (i) Chaco; (j) Parana; (k) South American
Transition Zone; (l) Andean region.

Taxa
Thirty-six taxa (Table 1) were analysed. They
include genera and species groups of insects, spiders,
vertebrates, and plants, which were chosen because
phylogenetic hypotheses were available for them and
their species are distributed in the areas analysed
herein. Although there are other potential taxa with
published phylogenetic analyses that could be considered, some of them lack information on the geographical distribution of the species analysed and others
include only a limited sample of the terminal taxa, as
occurs in most molecular analyses. Some of the taxa
analysed herein have been considered in previous cladistic biogeographical studies (Morrone and Coscar

on,
1998; Nihei and Carvalho, 2007; Sigrist and Carvalho,
2009; Pires and Marinoni, 2010; Echeverry and Morrone, in press).
Analysis
A cladistic biogeographical analysis is based on a
correspondence between phylogenetic relationships

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J. J. Morrone / Cladistics 30 (2014) 202214

Table 1
Taxa analysed, with the paralogy-free subtree derived from them
Taxa
Insecta: Coleoptera
Agaporomorphus
Chroaptomus
Entinmini
Hypselotropis
Ptychoderes
Rhinostomus
Insecta: Diptera
Coenopsia
Polietina
Pseudoptilolepis
Sepedonea
Insecta: Hemiptera
Eidmannia
Nicomia
Rasahus
Rhodnius
Serdia
Thoreyella and related genera
Insecta: Lepidoptera
Amorbia
Charis gynaea species group
Cliniodes
Insecta: Orthoptera
Abracrini
Arachnida: Araneae
Anelosinus ethicus species group
Vertebrata: Aves
Amazona ochrocephala complex
Chlorospingus ophthalmicus species complex
Pionopsitta
Vertebrata: Mammalia
Alouatta
Ateles
Caluromys
Marmosa murina
Metachirus nudicaudatus
Oryzomys megacephalus species group
Rhipidomys
Plants
Exostema
Hillia
Prosopis
Sabal
Stigmatopteris

Paralogy-free subtrees

Reference(s)

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

4a
4b
4c,d
4eg
4h,i
4j

Miller (2001)
Chani-Posse (2006)
Morrone (2002b), Vanin and Gaiger (2005)
Mermudes (2005)
Mermudes and Napp (2006)
Morrone and Cuevas (2002)

Fig.
Fig.
Fig.
Fig.

4k
4l
4m
4n

Nihei and Carvalho (2004)

Nihei and Carvalho (2007)
Schuehli and Carvalho (2005)
Pires and Marinoni (2010)

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

4o
4p
4q,r
4s,t
4u,v
4w

Coscar

on (1989), Morrone and Coscar

on (1998)
Albertson and Dietrich (2005)
Morrone and Coscar

on (1998)
Paula et al. (2007)
Fortes and Grazia (2005)
Bernardes et al. (2009)

Fig. 4xz, aa
Fig. 4bb
Fig. 4ccff

Phillips-Rodr
guez and Powell (2007)

Hall and Harvey (2001)
Hayden (2011)

Fig. 4ggii

Da Costa et al. (2010)

Fig. 4jj

Agnarsson (2005)

Fig. 4kk
Fig. 4ll
Fig. 4mm

Eberhard and Bermingham (2004)

Bonaccorso et al. (2008)
Ribas et al. (2005)

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

4nn
4oo
4pp
4qq
4rr
4ss
4tt

Cort
es-Ortiz et al. (2003)

Collins and Dubach (2000)
Costa (2003)
Costa (2003)
Costa (2003)
Costa (2003)
Costa (2003)

Fig.
Fig.
Fig.
Fig.
Fig.

4uu
4vvxx
4yy, zz
4aaa
4bbb, ccc

McDowell et al. (2003)

Taylor (1994)
Catalano et al. (2008)
Zona (1990)
Moran (1991)

and area relationships (Morrone, 2005a, 2009; Parenti

and Ebach, 2009). It comprises three basic steps: (1)
construction of taxonarea cladograms from taxon
cladograms, by replacing the terminal taxa by the
area(s) inhabited by them; (2) resolution of the problems due to widespread taxa, redundant distributions
and missing areas; and (3) derivation of general area
cladogram(s) representing the most logical solution
for all the taxa analysed (Morrone and Carpenter,
1994). General area cladograms represent hypotheses
on the biogeographical history of the taxa analysed
and the areas where they are distributed (Morrone,
2009).

To obtain the general area cladogram(s), a parsimony analysis of paralogy-free subtrees (Nelson and
Ladiges, 1996; Contreras-Medina et al., 2007; Le

on
Paniagua and Morrone, 2009; Morrone, 2009) was
undertaken. This method consists of four steps
(Fig. 3):
1 For each taxonarea cladogram, areas duplicated
or redundant in the descendants of a node are eliminated (Fig. 3a), so that geographical paralogy is eliminated or reduced significantly, and data are associated
only with informative nodes.
2 Components are identified on each of the paralogy-free subtrees obtained (Fig. 3b).

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J. J. Morrone / Cladistics 30 (2014) 202214

(a)

(b)

(c)

(d)

Fig. 3. Steps of the cladistic biogeographical analysis. (a) Original taxonarea cladograms; (b) paralogy-free subtrees derived from the taxon
area cladograms, with informative components marked on them; (c) data matrix of areas 9 components; (d) general area cladogram obtained.

3 A data matrix is compiled, scoring with 1 the

presence of a component in an area and 0 its
absence (Fig. 3c).
4 A parsimony analysis of the data matrix is undertaken to identify the general area cladogram (Fig. 3d).
Analysis of the 36 taxonarea cladograms identified
55 paralogy-free subtrees (Fig. 4), 85 informative components were extracted from them, and a data matrix
was constructed (Table 2). The matrix was analysed
using TNT (Goloboff et al., 2008), performing
searches of the most parsimonious cladograms with
the heuristic traditional search algorithm of TNT,
with 1000 replications, and tree-bisection-reconnection
branch-swapping (TBR), holding ten trees during each
replication. The effect of homoplasy on the results was
explored by conducting different implied weights analyses (Goloboff, 1993), with constants of concavity (k)
set to a different integer value of 130, where 1 is
weighted most severely against homoplastic characters.

Results
Two most parsimonious cladograms were obtained
analysing the matrix under equal weights. Implied
weights calculated with different constants of concavity (k = 130) consistently produced one of them
(Fig. 5). This general area cladogram shows that the

Neotropical areas in the strict sense constitute a monophyletic unit, whereas the Mexican and South American transition zones are the sister areas to the Nearctic
and Andean regions, respectively. Within the Neotropical region in the strict sense, a first dichotomy separates the Antilles from the continental areas, which in
a second dichotomy are arranged into a north-western
and a south-eastern component. Within the north-western component the areas split according to the
sequence northern Amazonia, south-western Amazonia, north-western South America, and Mesoamerica.
Within the south-eastern component the areas split following the sequence south-eastern Amazonia, Chaco,
and Parana. According to these results, the Caribbean
and Amazonian subregions as formerly recognized by
Morrone (2006) under a panbiogeographical framework do not represent natural areas: within the former, the Antilles do not group with Mesoamerica and
north-western South America; and within the latter,
south-eastern Amazonia is closer to Chaco and Parana
than to the remaining Amazonian areas.
Based on the general area cladogram, the following
regionalization is proposed:
Neotropical region
Antillean subregion
Amazonian subregion
Northern Amazonian dominion
South-western Amazonian dominion

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J. J. Morrone / Cladistics 30 (2014) 202214

(a)

(d)

(g)

(f)

(e)

(c)
(b)

(l)

(i)
(h)
(j)

(k)

(m)
(o)

(s)

(q)

(p)

(r)

(n)

(v)

(u)

(x)

(y)

(w)

(z)

(aa)

(dd)
(bb)
(cc)

(ee)

(ff)
(gg)
(ll)

(hh)
(ii)

(mm)

(kk)

(jj)

(rr)

(pp)
(qq)
(oo)
(nn)

(ww)
(uu)

(tt)

(vv)

(ss)
(yy)

(zz)

(xx)

(aaa)
(ccc)

(bbb)

Fig. 4. Paralogy-free subtrees analysed. (a) Agaporomorphus; (b) Chroaptomus; (c,d) Entimini; (eg) Hypselotropis; (h, i) Ptychoderes; (j) Rhinostomus; (k) Coenopsia; (l) Polietina; (m) Pseudoptilolepis; (n) Sepedonea; (o) Eidmannia; (p) Nicomia; (q,r) Rasahus; (s,t) Rhodnius; (u,v) Serdia;
(w) Thoreyella and related genera; (xz, aa) Amorbia; (bb) Charis gynaea species group; (ccff) Cliniodes; (ggii) Abracrini; (jj) Anelosinus ethicus
species group; (kk) Amazona ochrocephala complex; (ll) Chlorospingus ophthalmicus complex; (mm) Pionopsitta; (nn) Alouatta; (oo) Ateles; (pp)
Caluromys; (qq) Marmosa murina; (rr) Metachirus nudicaudatus; (ss) Oryzomys megacephalus species group; (tt) Rhipidomys; (uu) Exostema; (vv
xx) Hillia; (yy, zz) Prosopis; (aaa) Sabal; (bbb, ccc) Stigmatopteris. Areas as in Fig. 2.

Mesoamerican dominion
North-western South American dominion
Chacoan subregion
South-eastern Amazonian dominion
Chacoan dominion
Parana dominion

Some of the events implied by the general area cladogram (Fig. 6) can be associated with known tectonic
and geological information available:
1 The oldest event corresponds to the former
connection between the North American and South
American landmasses during the Early Jurassic to

208

Table 2
Data matrix, where files correspond to the areas analysed and columns to the components identified in the paralogy-free subtrees of Fig. 4

J. J. Morrone / Cladistics 30 (2014) 202214

root
00000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000
Nearctic region
00000000000000000000001000011111011000000000000000000000000000001001000000000000000011101000000000000000000000000000000000000000000000000000011111000000000
Mexican Transition Zone
00111000000000000000001000011111010000000000000000000000000000101000000000000000000000000000000000000000000000000000000000000000001111111000000000000000000
Mesoamerica
00111010000000010100101111111110011111100000001000000110010000110001111110001111011011111100000000100101000100000110100000000000001111111001100111111111111
Antilles
00000000000000000000001111100000000000000000000000000000000000001100001000001100010100000000000000000000000000000000000000000000000000000101100111101000010
North-western South America
00110000000110000110001100011100011100110100101111000111010000001111111111111000010111001111010100110111110111110110100000000000000001110111000111111111111
Northern Amazonia
00000011010111000000111110011000000000110110110011111001100000001111100011101110000000000001100010111001101111100100000100101111001001110100000000001110000
South-eastern Amazonia
10000000000001111111111111000000011100000000000010000000000000001110000000000000000000000001100000000001101110001111011111001111110000000100000000000000000
South-western Amazonia
11100111011101100111111111011000000000000011000000011000011000001111101011111110000011010001110100111111110111110111000110111001101001000111000110001111000
Chaco
11000011100000011111001111010000111110111000000011110000000111001110000011111111100000001111111011000000000111000000010000001110001110000000011110000000000
Parana
11100111111000011111110111110000111111111011111111111001100111111110001111001111111010010001111011000000000110001000011111111101110001100100000000001100000
South American Transition Zone
00000000000000000000000000000000000000000000000000011100011100000000000000000000000000000000000000000000000000000000000000000000001100000000010100000000000
Andean region
00000000000000000000000000000000000000000000000000000000000110000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000

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J. J. Morrone / Cladistics 30 (2014) 202214

Fig. 5. General area cladogram obtained.

(a)

(b)

(d)

(g)

(e)

(c)

(f)

(h)

Fig. 6. Sequence of events implied by the general area cladogram. (a) Geodispersal of the Neotropical biota from South to North America; (b)
vicariance of the Antilles and the rest of the Neotropical region; (c) vicariance of the north-western and south-eastern continental components;
(d) vicariance of northern Amazonia and the rest of the north-western component; (e) vicariance of south-western Amazonia and north-western
South AmericaMesoamerica; (f) vicariance of north-western South America and Mesoamerica; (g) vicariance of southern Amazonia and ChacoParana; (h) vicariance of Chaco and Parana.

Early Cretaceous (190148 Ma). It allowed the geodispersal of the Neotropical biota to North America and
would explain the presence of ancient Neotropical elements in northern Mexico (Halffter, 1987; Morrone,
2005b).

2 The vicariance between the Antilles and the rest

of the Neotropical region can be linked to the severance of the temporal connection represented by the
leading edge the Caribbean plate during its north-eastward drift (Echeverry and Morrone, in press). The

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J. J. Morrone / Cladistics 30 (2014) 202214

leading edge of the Caribbean plate reconnected temporarily with North and South America at 125
100 Ma. This connection began to be severed in the
Late Cretaceous (80 Ma) and finished in the Miocene
to Middle Pliocene (Pitman et al., 1993). The continental mainland constituting the rest of the Neotropical region shares extensive hydrological connections
and there is evidence that it constituted a superbasin
that has persisted in relative isolation since at least the
Late Cretaceous (Albert et al., 2011).
3 The vicariance between the north-western and
south-eastern continental components of the Neotropical region might have begun with the formation of a
lake along the Amazon, Madeira, and Mamor
e rivers,
in the Late Cretaceous; and finished with the formation of a wide epicontinental sea by water invasion
through the north, east, and south portal seaways, in
the Miocene (Amorim, 2001; Frailey, 2002; Nihei and
Carvalho, 2004, 2007).
4 The vicariance between northern Amazonia and
the remaining areas of the north-western continental
component can be linked to the Romeral Fault Zone
and/or the final uplift of the northern Andes. The
Romeral Fault, of Cretaceous age, is an active and
continuous fault system almost 700 km long that comprises three or four parallel regional faults, which form
the boundary between autochthonous continental
rocks to the east and accreted oceanic arc rocks
related to Caribbean terranes in the west (Kennan and
Pindell, 2009; Heads, 2012; Echeverry and Morrone,
in press). The uplift of the Andes began in the Cretaceous, but has been more conspicuous since the Miocene, 237 Ma, and finished in the Pliocene (Lundberg
et al., 1998; Kennan, 2000; Cort
es-Ortiz et al., 2003;
Garzione et al., 2008; Hoorn et al., 2010).
5 The vicariance between south-western Amazonia
and north-western South AmericaMesoamerica can
be linked to the formation of an epicontinental sea by
water invasion through the Maracaibo and Amazon
basins in the Pliocene (Rodr
guez-Olarte et al., 2011).
6 The vicariance between Chaco and Parana can be
linked to the connection of the Parna
ba and Paran
a
basins in the Palaeocene (Amorim, 2001; Nihei and
Carvalho, 2004).

Discussion
Previous analyses have suggested that the Amazonian and Caribbean subregions might not be natural
areas. Amorim and Pires (1996) considered that
Amazonia consisted of two non-related areas: northwestern Amazonia and south-eastern Amazonia. Amorim (2001) also considered that the Amazon forest did
not correspond to a natural biogeographical area,
being geographically delimited, based on the Amazon

river basin. Costa (2003) undertook a comparative

phylogeographical analysis of forest-dwelling small
mammals distributed between and within the Amazon
and Atlantic forests, finding that sequence similarity
was often greater between samples from the Atlantic
Forest and either Amazon or central Brazilian forests
than it was within each of both forest areas. The
Atlantic Forest clades were either not reciprocally
monophyletic or were the sister group to all the other
clades, and the central Brazilian area did not behave
as a separate region but as complementary to either
the Amazon or the Atlantic forests. Nihei and Carvalho (2007) tested two previous proposals (Amorim
and Pires, 1996; Morrone, 2006), by undertaking different cladistic biogeographical analyses based on the
areas implied by these authors, and concluded that
Amazonia should be regarded as a composite area,
because north-western Amazonia was closely related to
the Caribbean subregion, whereas south-eastern
Amazonia was related to the Chacoan and Parana
subregions. Sigrist and Carvalho (2009) analysed the
historical relationships among Neotropical areas of
endemism using Brooks parsimony analysis (BPA) to
examine whether the inclusion of open area formations
influences area relationships of the surrounding forests.
They found a basal split between the Amazonian and
Atlantic forests, suggesting that they have been isolated for a long period of time, and corroborated the
hypothesis that Amazonia is a composite area; however, when they added two areas with open formations
(Cerrado and Caatinga), internal relationships within
Amazonia changed, so they concluded that a biogeographical classification comprising both forests and
open formations should be preferred given their complementary history. Pires and Marinoni (2010) applied
BPA to species of Sepedonea (Diptera: Sciomyzidae),
finding that when the Cerrado and Caatinga were
included the Atlantic forest was monophyletic, whereas
the Amazonian forest was not, and concluded that a
single history of the current distribution of taxa in the
area analysed was unlikely. A recent panbiogeographicalcladistic biogeographical analysis of the Caribbean
subregion based on plant and animal taxa (Echeverry
and Morrone, in press) showed that it is not a natural
area. Differences among these analyses may be due to
different delimitations of the areas of endemism,
absence of non-forested areas in most of the analyses,
and the fact that most analyses did not use testable
procedures (Sigrist and Carvalho, 2009).
The major events postulated herein might have
occurred during the Cretaceous, when the Caribbean
plate collided with the Americas (Echeverry et al.,
2012), a combination of eustatic sea-level changes and
tectonic deformations of the continental platform
exposed large parts of South America to episodes of
marine transgressions (Albert and Reis, 2011), and

J. J. Morrone / Cladistics 30 (2014) 202214

Andean uplift reconfigured the Amazonian area (Garzione et al., 2008; Hoorn et al., 2010). Tertiary and
Quaternary events are assumed to have induced the
diversification within these large biogeographical units
(Bush, 1994; Werneck, 2011). The vicariance between
the Antilles and the rest of the Neotropical region,
and the vicariance between the north-western and
south-eastern continental components are coincident
with previous cladistic biogeographical analyses (Amorim and Pires, 1996; Amorim, 2001; Nihei and Carvalho, 2004). The dispersal of the Neotropical biota
from South to North Americaa prerequisite for the
first vicariant eventhas been supported by several
authors, working on different taxa, who have previously recognized two cenocrons: Old Southern or
Ancient Neotropical that dispersed between the Cretaceous and the Palaeocene; and a younger one that
dispersed between the Pliocene and Pleistocene, after
the establishment of the Panama Isthmus (Rosen,
1976; Gentry, 1982; Savage, 1982; Bussing, 1985;
Halffter, 1987; Morrone, 2005b). It is interesting to
note that the vicariance between north-western South
America and Mesoamerica occurs more recently than
detected in some previous analyses (Amorim and Pires,
1996; Amorim, 2001; Camargo and Pedro, 2003),
where it was considered as an older vicariant event. In
the present analysis this event has a younger age, as
suggested by Camargo (1996) and Camargo and Moure (1996).
Several authors (Bush, 1994; Bates et al., 1998;
Marks et al., 2002; Costa, 2003; Nihei and Carvalho,
2007) have concluded that it is not possible to postulate a single hypothesis explaining the current distribution of the Neotropical terrestrial biota. Pires and
Marinoni (2010) have even suggested that in the
absence of temporal information (inferred from molecular phylogenies) one cannot be sure that a general
pattern is due to a common history of biotic diversification. Additionally, it might be argued that taxa
showing so different dispersal means may not show a
clear, congruent pattern. I think, however, that biogeographical regionalizations are useful as general reference models (Ribichich, 2002). Their heuristic value
should be explored by examining the geographical distribution of other plant and animal taxa. Molecular
phylogenetic analyses that have been calibrated may
allow us to disprove the proposed sequence of vicariant events and help to clarify the time frame of the
events that led to the biotic diversification of the Neotropics.

Acknowledgments
I thank Amparo Echeverry and Dalton de Souza
Amorim for interesting discussions on the biogeogra-

211

phy of the Neotropics. Amparo Echeverry, Michael

Heads, and two anonymous reviewers made useful
comments that helped improve the manuscript.
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