Tropical Diversity (2022) 2(1): 1-21.

ISSN: 2596-2388

DOI: 10.5281/zenodo.10978657

RESEARCH ARTICLE

1

Distributional patterns of Characiformes in the Chacoan

Sub-region (Neotropical Region)

Padrões de distribuição de Characiformes na Sub-Região Chaquenha (Região Neotropical)

Vinícius Amaral Corrêa1*, Francisco José de Figueiredo2 https://orcid.org/0000-0002-1238-7447, Valéria Gallo1

1Laboratório de Sistemática e Biogeografia, Departamento de Zoologia, Instituto de Biologia, Universidade do Estado

do Rio de Janeiro (UERJ).

2Laboratório de Ictiologia, Departamento de Zoologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro

(UERJ)

*Email: viniciusamaral1986@gmail.com

Received: November 9, 2020 / Accepted: January 21, 2022 / Published: February 9, 2022

Resumo A maior diversidade de peixes de água

doce ocorre na Região Neotropical. Porém, há

poucos estudos em Biogeografia Histórica

incluindo um número representativo de espécies

ou áreas de grande extensão geográfica. Seus

padrões biogeográficos têm sido comparados com

aqueles identificados para táxons de organismos

terrestres, com o intuito de recuperar a história da

biota Neotropical. A história geológica de rios e

lagos costeiros do componente sudeste da Região

Neotropical é predominantemente recente,

apresentando complexidades bióticas singulares,

que resultaram em grande diversidade ictiológica.

Neste estudo, foi aplicado o método pan-

biogeográfico de análise de traços, utilizando o

software Martitracks, para identificar padrões de

distribuição de Characiformes da Sub-região

Chaquenha. Foram usadas 13.410 ocorrências

referentes a 132 espécies nominais, que

resultaram em 16 traços generalizados e um nó

biogeográfico. Os Characiformes apresentaram

padrões de distribuição concordantes com aqueles

de outros táxons. Os padrões obtidos estão de

acordo com o estado atual de conhecimento da

história geológica da Sub-região Chaquenha e se

ajustam aos Padrões Biogeográficos A, B e C

formalmente reconhecidos, determinados por

eventos geológicos antigos, intermediários e

recentes, respectivamente.

Palavras-Chave: Characiformes, Biogeografia

Histórica, análise de traços, Sub-região Chaquenha,

Região Neotropical.

Abstract The greatest diversity of freshwater

fishes occurs in the Neotropical Region. However,

there are a few Historical Biogeography studies

including a large amount of species or areas with

a great geographical extension. Their

distributional patterns have been compared with

those identified for taxa of terrestrial organisms,

with the purpose to recover the history of the

Neotropical biota. The geological history of

coastal rivers and lakes of the southeastern

component of the Neotropical Region is mainly

recent, showing singular biotic complexities,

resulting in great ichthyological diversity. In this

study, we applied the panbiogeographic method of

track analysis using the Martitracks software to

identify distributional patterns of Characiformes

from the Chacoan Sub-region. We used 13,410

occurrences related to 132 nominal species,

resulting in 16 generalized tracks and a single

biogeographic node. The Characiformes showed

distributional patterns matching with those of

other taxa. The patterns obtained is according to

the current state of knowledge about the

geological history of the Chacoan Sub-region and

fit in with the Biogeographic Patterns A, B and C,

formally recognized, determined by ancient,

intermediate, and recent geological events,

respectively.

Keywords:Characiformes, Historical Biogeography,

track analysis, Chacoan Sub-region, Neotropical

Region.

Corrêa et al., 2022

Historical Biogeography of Characiformes

2

Corrêa et al., 2022

Historical Biogeography of Characiformes

3

Introduction

The greatest diversity of freshwater fishes

occurs in the tropics, especially in the Neotropical

Region (Helfman et al., 2009). In absolute

numbers, about 7,000 nominal species of fishes

live in South and Central America (Albert & Reis,

2011), and globally one out of five known species

occur in the Neotropical Region (Vari &

Malabarba, 1998).

Characiformes is a diverse lineage of

actinopterygian fish from the continental waters of

Africa and the Americas (Betancur-R et al.,

2017). The clade comprises 24 families (Nelson et

al., 2016), from which four having more than 200

species distributed throughout Africa while the

rest is found in the Americas, with about 2,300

species. The group comprises a great diversity of

forms, behaviors and ecologies and includes the

piranhas, tetras, lambaris and dourados so

familiar to both fishermen and aquarists (Géry,

1977).

The richness of the Neotropical fish fauna

has been the focus of some discussions and some

efforts have been made in order to understand its

origin and history. Several studies have presented

evidence that speciation and genetic divergence of

the Neotropical fish fauna is related to geological

events between lowland water bodies and those of

adjacent crystalline shields and involves process

such as the catchment of rivers and headwaters

(Ribeiro, 2006; Albert & Reis, 2011; Ribeiro &

Menezes, 2015).

River systems in eastern part of Brazil,

including river mouths such as São Francisco in

Alagoas, have been considered important areas of

endemism for fish (Menezes, 1996). A list drawn

up by Bizerril (1994) included reference to

possible vicariant events that would have

promoted the differentiation of the ichthyofauna.

Several authors (e.g., Menezes, 1988; Ribeiro,

2006; Buckup, 2011; Echeverry & Gallo, 2015;

Ribeiro & Menezes, 2015) have discussed the

historical relationship between geotectonic

processes and differentiation of the ichthyofauna.

Thus, although the history of southeastern

Neotropical rivers and lakes is relatively recent

from a geochronological point of view, they have

unique and complex biotic history, which include

outstanding ichthyological diversity (Albert &

Reis, 2011).

Different tools of Historical Biogeography

could be used to reconstruct and understand the

history of isolation, divergence, and

diversification of fish species. Panbiogeography is

one approach to Historical Biogeography

consisting of an amount of methods with the

widest application (Heads, 2012; Morrone, 2015).

It was founded and developed from studies

realized by the Italian American Léon Camille

Marius Croizat (1894-1982), with components of

global fauna and flora. The results of applying his

method were published in some works, such as

Panbiogeography (Croizat, 1958) and Space,

Time, Form: The Biological Synthesis (Croizat,

1964). Later, the panbiogeographic methodology

was improved by several authors (Craw et al.,

1999; Grehan, 2011; Miranda & Dias, 2012) using

statistics and exact algorithms.

The study presented here aims to identify

distributional patterns of freshwater fishes,

specifically characiforms, from the Chacoan Sub-

region (Neotropical Region) applying

panbiogeographic method of track analysis, and

Corrêa et al., 2022

Historical Biogeography of Characiformes

4

comparing the results with other taxa, including

those that served as basis for the regionalization

proposed by Morrone (2014).

Materials and Methods

The panbiogeographic method of track

analysis used here delineates the individual tracks

of each taxon that are drawn as line graphs on

maps. These are superimposed to determine

generalized tracks and the intersecting localities

(nodes) between generalized tracks. This method

involves the following steps: 1) map each taxon

by connecting localities as a minimum length line

graph; 2) recognizing similar individual tracks

that have overlapping localities as a generalized

track; 3) recognizing nodes in areas where two or

more generalized tracks meet; 4) map the

individual and generalized tracks, and the nodes

(Heads, 2004; Morrone, 2004).

The individual track is the basic unit of

the study. It is a line that connects the locations

(geographical coordinates of each geographic

locality) for the distribution of a species or a

supraspecific taxon. This line is formed from the

union of points by the minimum distance

connecting all points (Grehan, 2011). The

Generalized Track (GT) is the overlap of two or

more individual tracks, which indicates that they

have common ancestral range subject to the same

vicariant events (geological, tectonic, or climatic)

(Craw, 1988; Craw et al., 1999; Morrone, 2004;

Grehan, 2011), or they represent a common path

of dispersion (i.e., geodispersion), or isolated

dispersion events (Morrone & Crisci, 1995; Craw

et al., 1999; Nihei & Carvalho, 2005).

Additionally, they can also indicate putative areas

of endemism (Morrone, 2004; Nihei & Carvalho,

2005).

We named the “Generalized Tracks” in

this study as GT - n, where “n” is an integer. For

those subunits that are inserted in major GTs, we

used GT n1-n2.

Nodes are represented here as the

intersection of two or more generalized tracks

(Craw et al., 1999; Crisci et al., 2003). According

to Grehan (2011), the nodes represent the

intersection of different ecologies, phylogenies, as

well as distributions. Nodes can also be

characterized as areas of biological endemism,

phylogenetic diversity, limits of geographic or

phylogenetic distribution and geographical

disjunction. The nodes correlate the biological

characteristics with the origin and/or geological

process that formed the biotas (Heads, 1989;

Crisci et al., 2003; Nihei & Carvalho, 2005).

Species included in this work were

selected based on geographic data (“type locality”

and “distribution”) from Buckup & Menezes

(2007) and records from freshwater bodies

(Arroio, Bacia, Baía, Córrego, Lago, Lagoa,

Laguna, Riacho, Rio and Tributário) of the

hydrographic regions according to ANA (2015).

The 132 species selected for analysis are listed in

the Appendix.

We used maps of hydrographic regions

available in

http://www3.ana.gov.br/portal/ANA/aguas-no-

brasil/panorama-das-aguas/copy_of_divisoes-

hidrograficasand

http://portal1.snirh.gov.br/ana/apps/webappviewer

/index.html?id=9cc5900ceb0d4c279305d4319798

0dd8 (accessed in July 25th, 2015).

Corrêa et al., 2022

Historical Biogeography of Characiformes

5

The geographical data used to estimate the

distribution of taxa and to perform track analysis

were obtained from the database of the following

collections: The Academy of Natural Sciences

Fish Collection (ANSP-Ichthyology, Philadelphia,

Pennsylvania, United States); Fish Collection of

the Departamento de Zoologia e Botânica,

Universidade Estadual Paulista (DZSJRP-Pisces,

São José do Rio Preto, São Paulo, Brazil);

Coleção Ictiológica da Universidade Federal do

Espírito Santo (CIUFES, Vitória, Espírito Santo,

Brazil); Coleção Ictiológica do Acervo Biológico

da Amazônia Meridional, Campus Sinop,

Universidade Federal de Mato Grosso (ABAM,

Sinop, Mato Grosso, Brazil); Coleção Zoológica

Delta do Parnaíba – Pisces, Universidade Federal

do Piauí (CZDP-Pisces, Parnaíba, Piauí, Brazil);

Laboratório de Ictiologia do Grupo de Ecologia

Aquática, Universidade Federal do Pará (GEA,

Belém, Pará, Brazil); Coleção de Peixes INPA,

Instituto Nacional de Pesquisas da Amazônia

(INPA-Peixes, Manaus, Amazonas, Brazil);

Coleção de Peixes do Laboratório de Ictiologia de

Ribeirão Preto, Universidade de São Paulo,

Campus Ribeirão Preto (LIRP, Ribeirão Preto,

São Paulo, Brazil); Coleção de Peixes do Museu

de História Natural Capão da Imbuia (MHNCI–

Peixes, Curitiba, Paraná, Brazil); Coleção

Científica da Divisão de Peixes do Museu de

Zoologia, Universidade Estadual de Feira de

Santana (MZFS, Feira de Santana, Bahia, Brazil);

Subcoleção Ictiológica do Campus Parnaíba

daUniversidade Estadual do Piauí (UESPI PHB,

Parnaíba, Piauí, Brazil); Coleção de Peixes do

Laboratório de Ictiologia Sistemática da

Universidade Federal do Tocantins (UNT, Porto

Nacional, Tocantins, Brazil); Zoneamento

Ecológico Econômico do Acre – Ictiofauna (ZEE-

ICTIO, Rio Branco, Acre, Brazil); Museum of

Comparative Zoology, Harvard University (HU-

Zoo, Cambridge, Massachusetts, United States);

Coleção de Peixes, Instituto Nacional da Mata

Atlântica (INMA), Museu de Biologia Prof. Mello

Leitão (MBML-Peixes, Santa Teresa, Espírito

Santo, Brazil); Coleção de Peixes, Pontifícia

Universidade Católica do Rio Grande do Sul

(MCP-Peixes, Porto Alegre, Rio Grande do Sul,

Brazil); Museu de Zoologia da Universidade

Estadual de Londrina, Coleção de Peixes

(MZUEL-Peixes, Londrina, Paraná, Brazil);

Coleção de Peixes do Museu de Zoologia da

Universidade de São Paulo (MZUSP - São Paulo,

São Paulo, Brazil); US-Animalia, National

Museum of Natural History, Extant Specimen and

Observation Records, Smithsonian Institution

(NMNH-Animalia, Washington, DC, United

States); Coleção de Peixes, Universidade Federal

do Rio de Janeiro (NUPEM/UFRJ, Macaé, Rio de

Janeiro, Brazil); Coleção Ictiológica do Nupélia,

Universidade Estadual de Maringá (NUP,

Maringá, Paraná, Brazil); Coleção de Peixes,

Universidade Federal do Rio Grande do Sul

(UFRGS - Porto Alegre, Rio Grande do Sul,

Brazil); Coleção de Peixes do Museu de Zoologia

da Universidade Estadual de Campinas (ZUEC-

PIS, Campinas, São Paulo, Brazil); Coleção

Zoológica de Referência da Universidade Federal

de Mato Grosso do Sul (ZUFMS-PIS, Campo

Grande, Mato Grosso do Sul, Brazil); Coleção

Ictiológica do Museu Nacional, Universidade

Federal do Rio de Janeiro (MNRJ, Rio de Janeiro,

Rio de Janeiro, Brazil).

The geographical coordinates were obtained

from the Global Biodiversity (GBIF, 2016) and

Corrêa et al., 2022

Historical Biogeography of Characiformes

6

the Species Link (SPLINK, 2016). We used only

“original” (those defined by the collectors) and

“not suspect” coordinates (those that coincide

with the names of the municipalities registered in

the collection).

For the construction of generalized tracks

and nodes, tools from Martitracks (Echeverría-

Londoño & Miranda-Esquivel, 2011) were used.

The following parameters were established: c = 2;

r = 2.5 (lmin), 3 (lmax), 4 (lmax.line); and m = 0.8

(min-SI). Where “c” is the Cut Value that

eliminates the redundant points of the analysis;

“r” is the Congruence Rule that checks whether

the individual tracks are congruent with each

other; “m” - min-Sl is a Similarity Index

(minimum congruence).

Firstly, the default parameters [c = 0.25; r =

0.25 (lmin), 0.50 (lmax), 0.75 (lmax.line); and m

= 0.85 (min-SI)] were used, but they generated

uninformative results. For this reason, we

modified them based on the simulation made by

Ferrari et al. (2013), who used the data provided

by Moreira et al. (2011). This simulation

generated similar results to those found in the

manual analysis performed by Moreira et al.

(2011). Also, Echeverría-Londoño & Miranda-

Esquivel (2011) tested the application with data

from Alzate et al. (2008), with the same

parameters herein used, as well as by Ferrari et al.

(2013)

The map in shapefile format of the

Neotropical Region (Morrone, 2014) for the

presentation of the results was available in

Löwenberg-Neto (2014). We used the software

QGIS 2.18 (QGISBRASIL, 2017) for plot and

analyze the tracks and nodes.

Track analysis was applied to 13,410

geographic coordinates for 132 species. After

each figure (Figs. 1-9) there is a description of the

generalized tracks and their biogeographic

location.

Results

From the analysis of 13,410 geographic

data for 132 species, 16 generalized tracks and

one node were obtained. The maps with the GTs

and nodes and their description are shown below.

Corrêa et al., 2022

Historical Biogeography of Characiformes

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Corrêa et al., 2022

Historical Biogeography of Characiformes

8

Figures 1-9 - Generalized tracks and nodes of Characiformes.

Corrêa et al., 2022

Historical Biogeography of Characiformes

9

 GT1 composed of Apareiodon affinis, A.

ibitiensis, A. vittatus, A. vladii, A. itapicuruensis,

A. hasemani, A. davisi, A. piracicabae and

Parodonhilari, located in Cerrado (Chacoan);

Atlantic and Parana Forest (Parana);

 GT2 composed of Curimatella lepidura,

Cyphocharax gilbert, Psectrogaster rhomboides,

P. saguiru, Steindachnerina elegans and S.

notonota, located in Caatinga and Cerrado

(Chacoan); Atlantic and Parana Forest (Parana);

 GT3 composed of Cyphocharax modestus, C.

nagelii, C. vanderi, C. santacatarinae and

Steindachnerina insculpta, located in Atlantic,

Parana Forest and Araucaria Forest (Parana);

 GT2-3 formed by line segment in common

between GT2 and GT3. Located in Atlantic and

Parana Forest (Parana);

 GT4 composed of Cyphocharax saladensis, C.

spilotus, C. voga and Steindachnerina biornata,

located in Chacoan and Pampean (Chacoan),

Araucaria Forest (Parana);

 GT5 composed of Prochilodus argenteus, P.

costatus, P. hartii, P. vimboides, P. brevis and P.

lineatus, in Madeira (South Brazilian), Caatinga,

Cerrado (Chacoan), Atlantic and Parana Forest

(Parana);

 GT6 composed of Characidium bahiense and

C. bimaculatum, crossing in Caatinga (Chacoan)

and Atlantic (Parana);

 GT7 composed of Characidium alipioi, C.

fasciatum, C. grajahuense, C. interruptum, C.

japuhybense, C. lagosantense, C. lauroi and C.

schubarti, crossing Cerrado (Chacoan), Atlantic

and Parana Forest (Parana);

 GT8 composed of Characidium timbuiense

and C. vidali, located in Parana Forest (Parana);

 GT9 composed of Characidium lanei, C.

oiticicai, C. pterostictum, C. occidentale, C.

orientale, C. rachovii, C. serrano and C. tenue,

located in Atlantic and Araucaria Forest (Parana);

 GT7-8 formed by line segment in common

between GT7 and GT8 e located in Atlantic and

Parana Forest (Parana);

 GT7-9 formed by line segment in common

between GT7 and GT9, located in Atlantic, Parana

Forest and Araucaria Forest (Parana);

 GT10 composed of Bryconferox, B. insignis, B.

nattereri, B. opalinus, B. orthotaenia and

Henochilus wheatlandii, crossing Cerrado

(Chacoan), Atlantic and Parana Forest (Parana);

 GT11 composed of Galeocharax gulo,

Phenacogaster calverti, P. franciscoensis and

Roeboides xenodon, spanning through Pará

(Boreal Brazilian), Cerrado (Chacoan).

 GT12 composed of Charax stenopterus e

Galeocharax knerii spanning Rondônia (South

Brazilian), Cerrado, Chacoan (Chacoan), Parana

Forest and Araucaria Forest (Parana);

 N1 formed by GT11 and GT12 located in

Cerrado (Chacoan).

 GT13 composed of Acinocheirodon

melanogramma, Serrapinnus heterodon,

Serrapinnus piaba, Compsura heterura and

Kolpotocheirodon theloura, crossing Caatinga,

Cerrado (Chacoan) and Parana Forest (Parana);

 GT14 composed of Cheirodon ibicuhiensis, C.

interruptus, Heterocheirodon jacuiensis,

Macropsobrycon uruguayanae, Serrapinnus

calliurus, S. notomelas, Spintherobolus leptoura,

S. papilliferus and S. ankoseion, crossing Cerrado,

Pampa (Chacoan), Atlantic, Parana Forest and

Araucaria Forest (Parana);

Corrêa et al., 2022

Historical Biogeography of Characiformes

10

 GT15 composed of Diapoma speculiferum, D.

terofali, Mimagoniates inequalis, M. lateralis, M.

microlepis, M. rheocharis, M. sylvicola,

Planaltina glandipedis, Pseudocoryno

pomadoriae and P. heterandria, crossing Pampean

(Chacoan), Atlantic, Parana Forest and Araucaria

Forest (Paraná);

 GT16 composed of Triportheus guentheri and

T. signatus, crossing Cerrado (Chacoan) and

Parana Forest (Parana);

Discussion

Overall, the generalized tracks and nodes

found here were corroborated by distributional

patterns of dipterans Cyrtoneurina,

Cyrtoneuropsis and Bitoracochaeta (Carvalho et

al., 2003) and Polietina (Nihei & Carvalho,

2005); hymenopteran Bombus (Abrahamovichet

al., 2004); and coleopteran Entimini (Romo &

Morrone, 2011), as follows:

1. GTg and GTf of Cyrtoneurina (cf.

Carvalho et al., 2003) match GT2-3 of

Curimatidae and GT7-9 of

Crenuchidae.

2. GTt of Cyrtoneuropsis (cf. Carvalho

et al., 2003) is congruent to GT2 of

Curimatidae and GT11 of Characinae;

GTu and GTvto GT2-3 of

Curimatidae, GT7-9 of Crenuchidae,

and GT15 of Glandulocaudinae.

3. GT CDE of Bithoracochaeta (cf.

Carvalho et al., 2003) match GT2-3 of

Curimatidae, GT7-9 of Crenuchidae,

GT15 of Glandulocaudinae.

4. GT7 of Bombus (cf. Abrahamovich et al.,

2004) match GT2-3 of Curimatidae, GT7-

9 of Crenuchidae, GT 5 of

Prochilodontidae, GT 14 of

Cheirodontinae, GT 15 of

Glandulocaudinae.

5. GT2 of Polietina (cf. Nihei & Carvalho,

2005) is congruent to GT6 of

Crenuchidae and GT2 of Curimatidae;

GT3 of Polietina(cf. Nihei & Carvalho,

2005) match GT6 de Crenuchidae; the

line segment formed by GT 4 and 6 de

Polietina (cf. Nihei & Carvalho, 2005)

match GT 5 of Prochilodontidae; line

segment formed by GT 5 and 7 of

Polietina (cf. Nihei & Carvalho, 2005)

match GT 7-9 of Crenuchidae; GT 7 of

Polietina (cf. Nihei & Carvalho, 2005)

match GT 2-3 of Curimatidae, GT 14 of

Cheirodontinae (Characidae), GT 15 of

Glandulocaudinae.

6. GT b Entimini (cf. Romo & Morrone,

2011) match GT2-3 and GT3 of

Curimatidae, GT7-9 of Crenuchidae, GT9

of Crenuchidae, GT14 of Cheirodontinae

(Characidae), GT 15 of Cheirodontinae

(Characidae).

Carvalho et al. (2003) discussed possible

vicariant events that could have shaped the

distribution of Muscidae (Diptera) with reference

to Amorim & Pires (1996). The authors

emphasized the connection between the Parnaíba

and Paraná basins, which occurred in the Late

Cretaceous. GT 11 and GT 12, and N1 of

Characinae (Characidae) (Fig. 6) were formed

from the overlapping of localities of rivers and

streams of the Paraná and Parnaíba basins.

Throughout its history, the Parnaíba River has

also been connected with other rivers in

northeastern Brazil, such as the São Francisco

River (Ribeiro, 2006).Due to these connections,

Corrêa et al., 2022

Historical Biogeography of Characiformes

11

the hydrographic basins of northeastern Brazil

allowed a faunal interchange resulting in a diverse

ichthyofauna (Ribeiro, 2006). In the present work,

the GT 16 formed by Triportheinae species (Fig.

9) joins localities or streams that flow from the

Parnaíba River to the São Francisco River.

Congruence found in the southeast of the

Brazilian coast between GT 2, GT 2-3, GT 7-9,

GT 15 and literature data is probably related to the

rise of Serra da Mantiqueira (Amorim & Pires,

1996; Carvalho et al., 2003) or Serra do Mar

(Ribeiro, 2006). New studies are necessary to

confirm this spatial correlation shaped by the

uplift of both mountains.

Comparing the distributional

patterns of fish with the results obtained

by Nihei & Carvalho (2005) for Polietina,

it is clear that there is congruence in the

distribution between Crenuchidae,

Curimatidae, Prochilodontidae,

Cheirodontinae (Characidae),

Glandulocaudinae (Characidae) and the

cited Diptera.

Generalized tracks may indicate

ancestral range subject to the same

vicariance events or isolated dispersion

events. If phylogenetically supported, the

generalized track indicates an area of

endemism or the preexistence of an

ancestral biota (Nihei & Carvalho, 2005).

Phylogenetically supported tracks are

those formed by sister species or closely

related species. According to phylogenetic

studies, it was found that:

 Melo et al. (2016) recognized the

following relationships:

(Prochiloduscostatus, P. lineatus) + (P.

argenteus, P. hartii).These species is in GT5;

 In the composition of GT10 of

Bryconinae (Characidae) there are

Bryconferox and B. insignis that are sister

species (Hilsdorf et al., 2008);

 In the composition of GT14, there are

Cheirodon interruptus and Serrapinnus

calliurus, which are sister species, according

to Mirande (2010);

 GT15 has in its composition

Mimagoniates lateralis, M. microlepis and M.

sylvicola which, according to the cladogram

presented by Menezes & Weitzman (2009)

the relationship between these species is not

fully understood, but they belong to the same

clade.

According to Nihei & Carvalho (2005),

generalized tracks formed by sister species or

closely related species are relevant to

understanding the history of species

diversification. It should be noted here that the

generalized tracks mentioned above are formed by

species belonging to the same group of species or

clade or are sister species. It is likely that the areas

in which these tracks are identified have an

effective historical determination in the

speciation.

There is biogeographic congruence

between Entimini and Curimatidae, Crenuchidae,

Cheirodontinae (Characidae) in the Atlantic and

Paraná Forest provinces in southeastern Brazil

(see figures 2, 4 e 7). As for the distribution of

Entimini, Romo & Morrone (2011) mention that

the establishment of the Savana Corridor (Schmidt

Corrêa et al., 2022

Historical Biogeography of Characiformes

12

& Inger, 1951; Prado & Gibbs, 1993; Morrone,

2006; Romo & Morrone, 2011) or Diagonal of

Open Formations (Vanzolini, 1963; Romo &

Morrone, 2011) along the Chacoan Sub-region

(north-central Argentina, southern Bolivia,

central-western Paraguay, Uruguay and central-

northeast Brazil) (Morrone, 2006), would have

been an important vicariant event for the fauna of

the Brazilian Sub-region and Chacoan Sub-region.

Regarding this work and the fish

distribution studied here, it should be considered

that only distributional patterns of the Chacoan

Sub-region were analyzed, therefore, there is no

possibility to assess the impact of the emergence

of the Savanna Corridor for characiformes. A

broader study would be necessary for this analysis

and there is a possibility that this event, like so

many other events, will have a reduced impact on

the distribution of fish, given the recurrence of

“headwater capture” and the connections between

the large hydrographic basins (Lundberg et al.,

1998; Ribeiro, 2006; Albert & Reis, 2011).

For the GTs 4, 12, 13 and 16 and node

N1, no similar examples were found in the

literature, due to probably the lack of collections

and studies (Morrone, 2004). There is also the

possibility that the incongruity is the result of

particularities of the analyzed taxon. In this sense,

many of the species of bony fish that contributed

to the tracks and node may have a unique

evolutionary history or linked to the histories of

the rivers that shelter them. A possible lack of

exact biological information for these species in

these mentioned tracks and node difficult to make

any argument to improve the biogeographic

understanding. The incongruity may be related to

the program used for the analysis of tracks, such

as Ferrari et al. (2013) stated that the subjective

definition of parameters can lead to an imprecise

analysis.

In the Late Cretaceous, large clades of

Neotropical ichthyofauna, including

Characiformes, had already diversified, as shown

by Lundberg et al. (1998). Although the oldest

record of characiform in South America is from

the late Campanian/early Maastrichtian (~83 to 72

Mya) (Gayet, 1991), probably the diversification

of this clade occurred before the final separation

of Africa and South America, also considering the

gaps in the fossil record. This means that the

history of the largest clades of bony fish occurred

before the emergence of modern rivers or their

present geography and catchment relationships

(Lundberg et al., 1998).

The geological history of the Neotropical

Region, specifically in the crystalline shield and

coastal drainages (Chacoan Sub-region), resulted

in three distinct biogeographic patterns (Ribeiro

2006).

Pattern A: coastal rivers of Brazil are

inhabited by taxa that have an ancient

biogeographic history (Stiassny & Pinna, 1994;

Ribeiro, 2006) dating from the Cretaceous, with

the diversification of an endemic ichthyofauna.

They are old taxa with few species and restricted

geographic distribution. The Characiformes has its

origin by at least in the Cretaceous (Brito et al.,

2007; Albert & Reis, 2011) with a minimum fossil

age of 83 to 72 Mya (Gayet, 1991). This coastal

pattern applies to the Crenuchidae (GT 6, 7, 8 and

9, Fig. 4).

Pattern B: generic level relationships

between the endemic ichthyofauna of coastal

drainages with the crystalline shield (Ribeiro,

Corrêa et al., 2022

Historical Biogeography of Characiformes

13

2006). None of the taxa sampled here show this

relationship.

Pattern C: is the result of faunal

exchange between the rivers of the crystalline

shield and the coastal drainages (Ribeiro, 2006).

The distribution along the lines of the Pattern C

can be corroborated by the generalized tracks that

connect or that are present in hybrid areas, which

are areas that have undergone neotectonics

processes that have led to fauna exchange.

Through the examples of hybrid areas given by

Ribeiro (2006) and the generalized tracks, it is

noticed that there is Pattern C in:

 In the region that includes the Upper

Uruguay River, Jacuí and the Patos Lagoon

System (Rio Grande do Sul, Brazil) and Negro

and Salado (Argentina). In this area were

found GT4 Curimatidae (Fig. 2).

 In the region that includes the Alto Rio

Tietê, Ribeira de Iguape, the tributaries of the

Paraná Basin, Rio Iguaçu and Rio

Paranapanema. In this area are: GT3 from

Curimatidae (Fig. 2) and GT14 from

Cheirodontinae (Fig. 7).

 The region that includes the northern

portion of the Paraná basin, São Francisco,

Paraíba do Sul, Itapicuru, Itapemirim, and the

mouth of the Rio Doce. In this area were

found: the GT 1 of Parodontidae (Fig. 1); GT2

Curimatidae (Fig. 2); GT 10 Bryconinae (Fig.

5); GT 13 from Cheirodontinae (Fig. 7); GT 15

from Glandulocaudinae (Fig. 8).

 The region that includes the São

Francisco River and the Parnaíba River: GT 16

from Triportheinae (Fig. 8).

In addition to the areas mentioned above, we have:

 The region that extends from the Paraná

River Basin to the Parnaíba River: The

GT 11 and GT 12, and the N1 of

Characinae (Fig 6) were formed from the

overlapping of localities of rivers and

streams of the Paraná basin and of

Parnaíba basin. Thus indicating a previous

connection of these, as previously

mentioned (Amorim & Pires, 1996).

 The region from the Paraná Basin to the

Amazon (Lundberg et al., 1998): GT 5 of

Prochilodontidae (Fig. 3) links Amazonas

to the tributaries of the Paraná, Rio São

Francisco, Paraíba do Sul. There is a

possibility that group took advantage of

sea level changes throughout history. The

distribution may be related to the

connection of the Paraná River with the

Amazon River, through the Paraná Sea

(Lundberg et al., 1998).

According to Ribeiro (2006), the

distributional patterns A, B and C are

consequences of old, intermediate, and recent

geological events, respectively.

According to the hypothesis that

corroborates Pattern A, cladogenetic events are

related to the origin of the first drains that flowed

into the Atlantic Ocean. Coastal rivers were

structurally oriented by megadomes, major flaws

and grabens. Ribeiro (2006) stated that this is the

case about the rivers that were established in the

megadomes Mantiqueira-Angola and Brazil-

Niger, where a fault system was responsible for

structuring the drainage pattern.

Corrêa et al., 2022

Historical Biogeography of Characiformes

14

The Pattern B suggests an interchange of

fauna between the crystalline shield and coastal

rivers across the Cenozoic. The continuous

erosive retraction on the east limit of the platform

was responsible for the transfer of fauna from the

rivers of the central plateau to the rivers of the

coastal plains. With taxa undergoing subsequent

diversification in both the coastal plain and

plateau drainages. The tectonism, through

reactivations and movements of large blocks, led

to the capture of hydrographic systems. This has

an impact on the distributional patterns of aquatic

biotas (Ribeiro, 2006).

The geological mechanism associated

with the Pattern C refers to the concept of

Neotectonics (Saadi, 2013). According to recent

models, the widespread rift system and other

crustal discontinuities present along the Atlantic

coast of South America act as areas of weakness

more prone to tectonic activity and deformations

(Saadi et al., 2002). Several hydrological

anomalies are probably related to tectonic

activations, most importantly the stream capture

(Cobbold et al., 2001).

The areas where the Pattern C occurs have

been identified as active tectonic areas, some of

which have a recent activity of approximately 1.6

MA (Saadi et al., 2002). This is what occurs in the

crystalline shield of southeastern Brazil, which

shares a mixed fauna with the drainages of the

coastal plain such as, for example, the area that

includes the headwaters of the Ribeira do Iguape,

Iguaçu and Paranapanema rivers and the upper

Tietê. It is important to note that in this area we

find the GT3 from Curimatidae (Fig. 2) and the

GT14 from Cheirodontinae (Fig. 7).

The generalized tracks in the regions of

Paranapanema, Iguaçu and Ribeira do Iguape can

be explained by the presence of the Ponta Grossa

Arch. It has a general tendency to uplift (Almeida

&Carneiro, 1998) and suffered tectonic activity

during the Cenozoic (Almeida & Carneiro, 1998;

Souza & Souza, 2002). The arc's tectonic activity

may have resulted in a change in the fluvial

dynamics of the area, accelerating the faunal

exchange between water bodies, such as the

coastal part of Ribeira do Iguape and the plateau

portions of Iguaçu and Paranapanema. The

distributional pattern of Curimatidae (Fig. 2) and

Cheirodontinae (Fig. 7) may be a consequence of

this.

The idea of southeastern Brazil being a

tectonically active area (Cobbold et al., 2001) is

supported by complex distributional patterns that

include the Pattern C. This tectonic activity

explains why the Paraná Basin has contributed

more to the development of ichthyofauna coastal

drainage, exemplified by the patterns of

Parodontidae (Fig. 1), Curimatidae (Fig. 2),

Bryconinae (Fig. 5), Cheirodontinae (Fig. 7) and

Glandulocaudinae (Fig. 8).

The tectonic control over the

distributional patterns of fish fauna in coastal and

plateau drainages is a recurrent process,

suggesting a high degree of faunal exchange

between basins (Ribeiro, 2006). The mixed nature

of hydrographic basins has already been noted by

Costa (2001), who indicated headwaters as

effective areas of faunal exchange. The same

tectonic process that allowed the interchange

between Brazilian coastal rivers with neighboring

drainages probably also occurred between basins

located in the interior of the continent in the

Corrêa et al., 2022

Historical Biogeography of Characiformes

15

crystalline shield. This pattern is found in

Characinae (Fig. 6), which occurs in the

crystalline shield and joins the Paraná and São

Francisco rivers.

The distributional patterns of

ichthyofauna in the Neotropical Region are related

to multiple factors: changes in the sea level

(Weitzman et al., 1988); retraction of the east

limit (Ribeiro & Menezes, 2015); old and recent

historical events; and, mainly, the capture of

rivers, headwaters, or entire hydrographic

systems.

It was found that the same clade can

present different and conflicting distributional

patterns. This is possible if we consider the

premise that the fish fauna of the Neotropical

Region is modern, and that the main groups of

fish have a minimum Cretaceous age (Brito et al.,

2007).

Conclusion

Through tracks analysis it was possible to

identify that the distributional patterns of

characiformes from Chacoan Subregion fit into

the Patterns A, B and C already described in the

literature.

The patterns found are corroborated by

others presented by different taxa with different

dispersion capacities, but which may have

something in common when compared to their

histories.

In addition, the results indicate that the

history of Neotropical fish and the rivers that

shelter them is intricate and of great complexity,

resulting from ancient and recent geological

processes, from the setting of a watercourse to the

interaction between them through headwaters

capture, streams, rivers, and even entire

hydrographic systems.

ACKNOWLEDGMENTS

We are grateful to John Grehan, Mauro

Cavalcanti, Michael Heads and Paulo Roberto

Lopes for insightful review of the manuscript. VG

is partially supported by a CNPq grant and

receives anUERJ/FAPERJ Prociência grant.

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Historical Biogeography of Characiformes

20

APPENDIX- Fish species included in this study.

- CHARACIFORMES – Parodontidae:

Apareiodon affinis (Steindachner, 1879), A. davisi

(Fowler, 1941), A. hasemani Eigenmann, 1916, A.

ibitiensis Campos, 1944, A. itapicuruensis

Eigenmann & Henn, 1916, A. piracicabae

(Eigenmann, 1907), A. vittatus Garavello, 1977,

A. vladii Pavanelli, 2006 and Parodon hilarii

Reinhardt, 1866; Curimatidae: Curimatella

lepidura (Eigenmann & Eigenmann, 1889),

Cyphocharax gilbert (Quoy & Gaimard, 1824), C.

modestus (Fernández-Yépez, 1948), C. nagelii

(Steindachner, 1881), C. saladensis (Meinken,

1933), C. santacatarinae (Fernández-Yépez,

1948), C. spilotus (Vari, 1987), C. vanderi

(Britski, 1980), C. voga (Hensel, 1869),

Psectrogaster rhomboids Eigenmann &

Eigenmann, 1889, P. saguiru (Fowler, 1941),

Steindachnerina biornata (Braga &

Azpelicueta, 1987), S. elegans (Steindachner,

1874), S. insculpta (Fernández-Yépez, 1948)

andS. notonota (Miranda Ribeiro, 1937);

Prochilodontidae: Prochilodus argenteus

Agassiz, 1829, P. brevis Steindachner, 1874, P.

costatus Valenciennes, 1850, P. hartii

Steindachner, 1874, P. lineatus (Valenciennes,

1836) and P. vimboides Kner, 1859;

Anostomidae: Leporellus vittatus (Valenciennes,

1850), Leporinus aguapeiensis Campos, 1945, L.

amblyrhynchus Garavello & Britski, 1987, L.

bahiensis Steindachner, 1875, L. conirostris

Steindachner, 1875, L. copelandii Steindachner,

1875, L. crassilabris Borodin, 1929, L. elongates

Valenciennes, 1850, L. garmani Borodin, 1929, L.

lacustris Campos, 1945, L. marcgravii Lütken,

1875, L. melanopleura Günther, 1864, L.

mormyrops Steindachner, 1875, L. obtusidens

Valenciennes, 1836, L. octofasciatus

Steindachner, 1915, L. paranensis Garavello &

Britski, 1987, L. piau Fowler, 1941, L. reinhardti

Lütken, 1875, L. steindachner iEigenmann, 1907,

L. striatus Kner, 1858, L. taeniatus Lütken, 1875,

L.thayeri Borodin, 1929, Schizodon australis

Garavello, 1994, S. intermedius Garavello &

Britski, 1990, S. jacuiensis Bergman, 1988, S.

knerii (Steindachner, 1875) and S. nasutus Kner,

1858; Crenuchidae: Characidium alipioi

Travassos, 1955, C. bahiense Almeida, 1975, C.

bimaculatum Fowler, 1941, C. fasciatum

Reinhardt, 1866, C. interruptum Pellegrin, 1909,

C. japuhybense Travassos, 1949, C. lagosantense

Travassos, 1947, C. lanei Travassos, 1967, C.

lauroi Travassos, 1949, C. occidentale Buckup&

Reis, 1997, C. oiticicai Travassos, 1967, C.

orientale Buckup& Reis, 1997, C. pterostictum

Gomes, 1947, C. rachovii Regan, 1913, C.

schubarti Travassos, 1955, C. Serrano Buckup&

Reis, 1997, C. tenue (Cope, 1894), C. timbuiense

Travassos, 1946, C. vestigipinne Buckup& Hahn,

2000 and C. vidali Travassos, 1967; Characidae

– Bryconinae: Bryconferox Steindachner, 1877,

B. hilarii (Valenciennes, 1849), B. insignis

Steindachner, 1877, B. nattereri Günther, 1864, B.

opalinus (Cuvier, 1819), B. orthotaenia Günther,

1864 and Henochilus wheatlandii Garman, 1890;

Characidae – Characinae: Charax stenopterus

(Cope, 1894), Galeocharax gulo Cope, 1870, G.

knerii Steinachner, 1875, Phenacogaster calverti

(Fowler, 1941), P. franciscoensis Eigenmann,

1911 and Roeboides xenodon (Reinhardt, 1851);

Characidae – Cheirodontinae: Acinocheirodon

melanogramma Malabarba & Weitzman, 1999,

Cheirodon ibicuhiensis Eigenmann, 1915, C.

Corrêa et al., 2022

Historical Biogeography of Characiformes

21

interruptus (Jenyns, 1842), Compsuraheterura

Eigenmann, 1915, Heterocheirodon jacuiensis

Malabarba & Bertaco,1999, Kolpotocheirodon

figueiredoi Malabarba, Lima & Weitzman, 2004,

K. theloura Malabarba & Weitzman, 2000,

Macropsobrycon uruguayanae Eigenmann, 1915,

Serrapinnus calliurus (Boulenger, 1900), S.

heterodon (Eigenmann, 1915), S. notomelas

(Eigenmann, 1915), S. piaba (Lütken, 1875),

Spintherobolu sankoseion Weitzman &

Malabarba, 1999, S. broccae Myers, 1925, S.

leptoura Weitzman & Malabarba, 1999 and S.

papilliferus Eigenmann, 1911; Characidae –

Glandulocaudinae: Diapoma speculiferum Cope,

1894, D. terofali Géry, 1964, Glandulocauda

melanogenys Eigenmann, 1911, G. melanopleura

Eigenmann, 1911, Hysteronotus megalostomus

Eigenmann, 1911, Mimagoniates inequalis

Eigenmann, 1911, M. lateralis Nichols, 1913, M.

microlepis Steindachner, 1877, M. rheocharis

Menezes & Weitzman, 1990, M. sylvicola

Menezes & Weitzman, 1990, Planaltina britski

Menezes, Weitzman & Burns, 2003, P.

glandipedis Menezes, Weitzman & Burns, 2003,

Pseudocoryno pomadoriae Perugia, 1891 and P.

heterandria Eigenmann, 1914; Characidae –

Iguanodectinae: Piabucus melanostomus

Holmberg, 1891; Characidae – Serrasalminae:

Myleus altipinnis (Valenciennes, 1850), M.

micans (Lütken, 1875), Myloplustiete (Eigenmann

& Norris, 1900), Pygocentrus piraya (Cuvier,

1819), Serrasalmus brandti (Lütken, 1875) and S.

marginatus Valenciennes, 1836; Characidae –

Stethaprioninae: Orthospinus franciscensis

(Eigenmann, 1914); Characidae –

Tetragonopterinae: Tetragonopterus chalceus

Spix & Agassiz, 1829; Characidae –

Triportheinae: Lignobrycon myersi (Miranda

Ribeiro, 1956), Triportheus guentheri (Garman,

1890) and T. signatus (Garman, 1890).