PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL

FACULDADE DE BIOCIÊNCIAS
PROGRAMA DE PÓS-GRADUAÇÃO EM BIOCIÊNCIAS – ZOOLOGIA

TAXONOMIA E FILOGENIA DO GÊNERO Hollandichthys EIGENMANN, 1909

(TELEOSTEI: CHARACIDAE) DO SUL E SUDESTE DO BRASIL

Vinicius Araújo Bertaco

Orientador: Dr. Luiz Roberto Malabarba

DISSERTAÇÃO DE MESTRADO
PORTO ALEGRE – RS – BRASIL
2003
 SUMÁRIO

Dedicatória ...............................................................................................................................iv

Agradecimentos ........................................................................................................................vi

Apresentação ...........................................................................................................................viii

Taxonomy and phylogeny of Hollandichthys Eigenmann, 1909 (Teleostei: Characidae) from

South and Southeastern Brazil ...................................................................................................1

Abstract ......................................................................................................................................2

Resumo .......................................................................................................................................3

Introduction ................................................................................................................................4

Methods and Materials ...............................................................................................................6

Results ........................................................................................................................................8

Character descriptions ....................................................................................................8

Phylogenetic relationships of Hollandichthys ..............................................................31

Phylogenetic relationships of Hollandichthys species .................................................45

Taxonomic revision of Hollandichthys.........................................................................49

Key of the species of Hollandichthys............................................................................52

Hollandichthys multifasciatus.......................................................................................54

Hollandichthys affinis...................................................................................................61

Hollandichthys perstriatus............................................................................................67

Hollandichthys sp. n. A.................................................................................................74

Hollandichthys sp. n. B.................................................................................................80

Hollandichthys sp. n. C.................................................................................................85

Hollandichthys sp. n. D.................................................................................................91

Hollandichthys sp. n. E.................................................................................................96

Biogeography of Hollandichthys............................................................................................103

ii
Discussion ..............................................................................................................................107

List of the comparative material.............................................................................................107

Acknowledgments ..................................................................................................................109

Literature cited .......................................................................................................................110

Figures ....................................................................................................................................118

Tables .....................................................................................................................................140

Table and figure legends ........................................................................................................153

Appendix ................................................................................................................................160

iii
 A minha Família,
e a Verônica.

iv
Estou convicto de que, hoje, nossa sociedade como um todo encontra-se numa crise análoga.
Temos taxas elevadas de inflação, desemprego, crise energética e na assistência a saúde,
poluição e desastres ambientais. Tudo isso são facetas diferentes de uma só crise, que é,
essencialmente, uma crise de percepção. Precisamos, pois, de um novo “paradigma” – uma
nova visão da realidade...uma visão holística e ecológica...

Fritjof Capra

Acredito que a visão de mundo sugerida pela física moderna seja incompatível com a nossa
sociedade atual, a qual não reflete o harmonioso estado de inter-relacionamento que
observamos na natureza. Para se alcançar tal estado de equilíbrio dinâmico, será necessária
uma estrutura social e econômica radicalmente diferente: uma revolução cultural na
verdadeira acepção da palavra. A sobrevivência de toda a nossa civilização pode depender
de sermos ou não capazes de realizar tal mudança.

Fritjof Capra
Trechos retirados do livro:
Ponto de Mutação - 1982

v
AGRADECIMENTOS
Ao meu Orientador, Prof. Dr. Luiz Roberto Malabarba, pela dedicação e excelente
orientação, amizade e apoio durante todo o desenvolvimento deste trabalho. Também, estou
muito grato pela ajuda nas fotos e por redigir e revisar o texto em inglês. Gracias!
A minha Família, por me incentivar, apoiar, ajudar e mostrar o caminho da sabedoria e
do sucesso e, principalmente, por me dar condições de chegar até aqui.
Um agradecimento especial para a minha Mãe, Heloisa de A. Bertaco, e a minha Vó,
Wanda M. de Araújo, para mim, exemplos de vida.
A Verônica C. Baumbach, pelo companheirismo, amor, apoio e incentivo durante
todas as etapas deste estudo. "Hoje, ela já sabe até coletar Hollandichthys".
Ao diretor do Museu de Ciências e Tecnologia da PUCRS, Jeter J. Bertoletti, por
providenciar suporte técnico durante a coleta dos peixes.
Aos professores e pesquisadores do Laboratório de Ictiologia do Museu de Ciências e
Tecnologia da PUCRS, Carlos A. S. de Lucena, Zilda Margarete S. Lucena e Roberto E. dos
Reis, pelo grande apoio e dedicação nas várias discussões levantadas sobre taxonomia e
filogenia de Characiformes nas aulas do curso de mestrado, e durante o desenvolvimento
deste trabalho.
Aos colegas do Laboratório de Ictiologia do Museu de Ciências e Tecnologia da
PUCRS, Alexandre R. Cardoso, Cíntia Kaefer, Cristina Buhrnhein, Edson Henrique L.
Pereira, Ignacio B. Moreno, José F. Pezzi da Silva, Pablo Lehmann (O Colombiano), Paulo H.
F. Lucinda, Thiago Borges, Tiago Carvalho, pela colaboração em alguma etapa deste estudo.
Ao Mineiro, Paulo H. F. Lucinda, pelas discussões taxômicas e filogenéticas do grupo
em estudo, pela ajuda em alguma instância para redigir o texto em inglês e nos programas
envolvidos na análise filogenética.
Ao Dr. Fernando G. Becker, mais conhecido como Fritz, pelo material coletado na
bacia do rio Maquiné e, com isso, o grande incentivo para estudar os Hollandichthys.
A Miriam S. Ghazzi e ao Flávio Lima, MZUSP, por calcularem as coordenadas de
algumas localidades do estado de São Paulo.
Ao Dr. Fernando G. Becker, José F. Pezzi da Silva, Juan Anza, Prof. Dr. Luiz R.
Malabarba, Marco A. Azevedo, P. Colombo, Pablo Lehmann, Thomas Hasper, pela ajuda nas
várias expedições de coleta.
Ao Marco A. Azevedo (UFRGS), pelos cortes histológicos em gônadas de alguns
espécimes de Hollandichthys.
Ao Dr. Stanley Weitzman, pelas discussões e sugestões via e-mail.

vi
 Aos curadores ou responsáveis de coleções científicas pelo empréstimo do material
para o estudo: Mark Sabaj e John Lundberg (ANSP), David Catania (CAS), Francisco
Langeani (DZSJRP), Barry Chernoff (FMNH), Ivan Mojica (ICNMHN), Karsten Hartel
(MCZ), Vinicius Abilhoa (MHNCI), Claude Weber (MHNG), Pablo Lehmann (MHNUC),
Paulo Buckup (MNRJ), Osvaldo Oyakawa (MZUSP), Helmut Wellendorf (NMW), Wilson
Costa (UFRJ), Susan Jewett (USNM).
Ao Osvaldo Oyakawa, por coletar e fotografar os Hollandichthys da Estação
Ecológica Juréia-Itatins, São Paulo.
Ao Antony Harold, pelo envio de parte do material comparativo.
Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) pela
concessão da bolsa de mestrado, e auxílio financeiro para o custeio de parte do projeto (Proc.
464545/00-5).
A Pontifícia Universidade Católica do Rio Grande do Sul e Museu de Ciências e
Tecnologia pela infra-estrutura e recursos oferecidos, sem os quais não seria possível a
realização deste trabalho.
Ao Programa de Pós-Graduação em Biociências desta universidade, pela oportunidade
de realizar o curso.
Ao Centro de Microscopia e Microanálises (CEMM) da PUCRS, pelas imagens de
microscopia eletrônica de varredura dos dentes e ossos da boca de alguns espécimes de
Hollandichthys.
Ao PRONEX, Projeto de "Conhecimento, conservação e utilização racional da
diversidade da fauna de peixes do Brasil", coordenado pelo Dr. Naércio A. Menezes
(MZUSP), por disponibilizar recursos financeiros para a realização das expedições de coletas
de Hollandichthys.
A todos que me ajudaram direta ou indiretamente para a concretização deste estudo, o
meu Muito Obrigado!

vii
APRESENTAÇÃO
Esta dissertação está sendo apresentada conforme as Normas do Programa de Pós-
Graduação em Biociências da Pontifícia Universidade Católica do Rio Grande do Sul -
PUCRS.
A dissertação está estruturada em forma de um artigo científico redigido em inglês. É
apresentado um resumo em português. As tabelas e figuras estão no final do texto.
O periódico escolhido para a publicação da dissertação intitulada "Taxonomy and
Phylogeny of Hollandichthys Eigenmann, 1909 (Teleostei: Characidae) from South and
Southeastern Brazil" foi a revista "Neotropical Ichthyology", revista nova da Sociedade
Brasileira de Ictiologia com publicação semestral.
O artigo científico foi redigido conforme as normas que constam nas instruções para
os autores do periódico escolhido. Estas normas estão em Anexo (Appendix).
Neste estudo foi realizada uma revisão taxonômica das diversas populações de
Hollandichthys Eigenmann encontradas nos rios costeiros e ilhas do sul e sudeste do Brasil e
rio Tietê superior, endêmico da Mata Atlântica, e uma análise filogenética das relações de
Hollandichthys com Pseudochalceus Kner e outros gêneros de Characidae. O gênero
Hollandichthys é diagnosticado com base em sinapomorfias relacionadas a caracteres
osteológicos e de morfologia externa. É considerado grupo irmão de Pseudochalceus e mais
intimamente relacionado com Nematobrycon Eigenmann, Rachoviscus Myers, Nematocharax
Weitzman, Menezes & Britski e Aphyocharax Günther do que com outros representantes de
Characidae examinados. A espécie-tipo H. multifasciatus (Eigenmann & Norris), e as
espécies nominais H. affinis (Steindachner) e H. perstriatus (Miranda-Ribeiro) são redescritas
e revalidadas com base em novos espécimes. Cinco novas espécies são descritas para os rios
costeiros e ilhas entre o sul e sudeste brasileiro. Essas espécies distinguem-se pelo número de
raios procorrentes dorsais e ventrais da nadadeira caudal, número de escamas na série
longitudinal, diâmetro orbital e colorido padrão. Neótipo e Lectótipo são designados para H.
perstriatus e H. affinis, respectivamente. É fornecida uma chave para as oito espécies de
Hollandichthys. Com base na filogenia proposta para as espécies de Hollandichthys, discute-
se as relações históricas entre as drenagens costeiras do Sul e Sudeste do Brasil.

viii
 Taxonomy and Phylogeny of Hollandichthys Eigenmann, 1909 (Teleostei: Characidae)

from South and Southeastern Brazil

Vinicius Araújo Bertaco and Luiz Roberto Malabarba

(VAB) Laboratório de Ictiologia, Museu de Ciências e Tecnologia e Programa de Pós-

Graduação em Zoologia, Pontifícia Universidade Católica do Rio Grande do Sul, PUCRS,

Av. Ipiranga 6681, 90619-900, Porto Alegre, RS, Brasil. ubertaco@pucrs.br

(LRM) Laboratório de Ictiologia, Museu de Ciências e Tecnologia, Pontifícia Universidade

Católica do Rio Grande do Sul, PUCRS, Av. Ipiranga 6681, 90619-900, Porto Alegre, RS,

Brasil; and Departamento de Zoologia, Universidade Federal do Rio Grande do Sul, UFRGS,

Av. Bento Gonçalves, 9500, 91501-970, Porto Alegre, RS, Brasil. malabarb@pucrs.br

1
ABSTRACT

The genus Hollandichthys Eigenmann (1909) is revised and diagnosed based on

synapomorphies described from osteological and external morphology analysis. The type-

species H. multifasciatus (Eigenmann & Norris), and the nominal species H. affinis

(Steindachner) and H. perstriatus (Miranda-Ribeiro) are revalidated and redescribed, based on

new specimens. Five new species are described from the coastal rivers and islands between

south and southeastern Brazil. These are distinguished among themselves manly by number

of dorsal and ventral procurrent caudal-fin rays, number of scales in a longitudinal series

(same of the lateral line), orbital diameter, color pattern and color in life. The distribution area

of the genus is enlarged reaching the rio Tramandaí drainage, Rio Grande do Sul State, South

Brazil. Hollandichthys is hypothesized to be the sister group of Pseudochalceus, and this

clade more closely related to the clade including Nematobrycon, Rachoviscus, Nematocharax

and Aphyocharax than to other examined representatives of the Characidae. Neotype and

Lectotype are designated for H. perstriatus and H. affinis, respectively. A key to the eight

species of Hollandichthys is provided.

Keywords: Neotropical Fishes, Atlantic Forest, Hollandichthys, Phylogeny,

Characidae.

2
RESUMO

O gênero Hollandichthys Eigenmann (1909) é revisado e diagnosticado com base em

sinapomorfias relacionadas a caracteres osteológicos e de morfologia externa. A espécie-tipo,

H. multifasciatus (Eigenmann & Norris), e as espécies nominais H. affinis (Steindachner) e H.

perstriatus (Miranda-Ribeiro) são redescritas e revalidadas com base no exame de novos

espécimes. Cinco espécies novas são descritas para os rios costeiros e ilhas do sul e sudeste

brasileiro. Essas espécies distinguem-se, principalmente, pelo número de raios procorrentes

dorsais e ventrais da nadadeira caudal, número de escamas na série longitudinal, diâmetro

orbital e colorido padrão. A área de distribuição de Hollandichthys é ampliada ao Sul para o

sistema do rio Tramandaí, Rio Grande do Sul, sul do Brasil. Hollandichthys é considerado

grupo irmão de Pseudochalceus e este clado mais intimamente relacionado com o clado

formado por Nematobrycon, Rachoviscus, Nematocharax e Aphyocharax do que com outros

representantes de Characidae examinados. Neótipo e Lectótipo são designados para H.

perstriatus e H. affinis, respectivamente. É fornecida uma chave de identificação para as oito

espécies de Hollandichthys.

Palavras-chave: Peixes Neotropicais, Mata Atlântica, Hollandichthys, Filogenia,

Characidae.

3
INTRODUCTION

The genus Hollandichthys was initially proposed by Eigenmann (1909:257), and

diagnosed in key by the presence of “maxillary with teeth along its entire edge; lateral line

very short; species with dark lines along the sides”. In this paper, Eigenmann listed one

species, Hollandichthys multifasciatus (Eigenmann & Norris, 1900), and considered

Pseudochalceus affinis Steindachner, 1908, and P. perstriatus Miranda-Ribeiro, 1908 as

junior synonyms. Latter, Eigenmann (1910:432) listed Tetragonopterus multifasciatus

Eigenmann & Norris, 1900 as the type-species of the genus, and maintained the synonyms

previously proposed.

Among the nominal species listed by Eigenmann to Hollandichthys, Tetragonopterus

multifasciatus was briefly described by Eigenmann & Norris (1900) based in three specimens

collected in Cubatão [São Paulo]. Pseudochalceus affinis was described by Steindachner

(1908) to Joinvile, Santa Catarina, and P. perstriatus to “Corregos de Iporanga” [São Paulo],

by Miranda-Ribeiro (1908), all localities in the coastal drainages of Southeastern Brazil (Fig.

1).

Eigenmann (1917), discussing variability of the Tetragonopterinae pointed out the

common presence of dark lines following the spaces between successive rows of scales in

Astyanax lineatus (Perugia, 1891), A. steindachneri [= Bario steindachneri (Eigenmann,

1893)], Moenkhausia latissimus [= Gymnocorymbus latissimus (Eigenmann, 1908)],

Hollandichthys and Pseudochalceus Kner (1863). While discussing polyphyletic characters

diagnosing Tetragonopterinae genera, Eigenmann considered the complete dentition in the

maxillaries observed in Pristella Eigenmann (1908), Hemibrycon Günther (1864),

Nematobrycon Eigenmann (1911), Hollandichthys, and Pseudochalceus, versus the presence

of a few teeth or none, a derived character, acquired in at least three independent groups

among these genera. One of these groups seems to be formed by Pseudochalceus plus

4
Hollandichthys, indicated in the Fig. 1 of Eigenmann (1917:49) as a single lineage derived

from the "central" genus Astyanax Baird & Girard (1854), from which "many lines have

radiated" [sic].

Although proposed in 1909, only in 1921 Eigenmann presented a brief description for

the monotypic genus Hollandichthys. In this paper, Eigenmann redescribed the type-species

H. multifasciatus, and noted a similarity with the genus Pseudochalceus.

In 1966, Schultz described one new species of the Pseudochalceus from Colombia and

considered Hollandichthys a junior synonym of Pseudochalceus due to the lack of

"important" anatomical differences between the species of the two genera.

Géry (1972a, b), studding the fishes from Ecuador, compared the type-species of the

genera Pseudochalceus and Hollandichthys and found out several interspecific differences,

and decided to use Hollandichthys as a subgenus of Pseudochalceus.

Considering the differences between the species of the two genera and the distribution

of H. multifasciatus (Atlantic coastal streams between São Paulo and Santa Catarina states)

and of the species of Pseudochalceus (western Ecuador and Colombia), Weitzman et al.

(1986) decided to revalidate Hollandichthys until a phylogenetic analysis to demonstrate

possible relationships between these genera become available.

The genus Hollandichthys is herein phylogenetically diagnosed and its relationships

with other characid genera discussed. The type-species H. multifasciatus, and the nominal

species H. affinis and H. perstriatus are revalidated and redescribed, based on new specimens.

Five new species are described from the coastal rivers and island between south and

southeastern Brazil.

5
METHODS AND MATERIALS

Counts were taken as described by Fink & Weitzman (1974), with the exception of the

number of scale rows below lateral line which were counted from the scale row ventral to

lateral line to the scale row nearest the first pelvic-fin ray. Total vertebral counts include the

four vertebrae of the Weberian apparatus. Supraneural, gill-rakers of first arch, teeth and

procurrent caudal-fin-ray counts were taken from cleared and stained specimens prepared

according to the method of Taylor & Van Dyke (1985). Procurrent caudal-fin-ray counts were

also taken in some alcohol preserved specimens. Osteological terminology is that used by

Weitzman (1962). Teeth and bone SEM (scanning electronic microscope) photos were taken

from dissected cleared and stained specimens.

Measurements were taken point to point with dial calipers, and made in the left side of

specimens when possible. All measurements other than standard length (SL) are expressed as

a percentage of SL except subunits of the head, which are recorded as a percentage of head

length (HL). Basic descriptive statistics and statistical tests for meristic data were computed

using SigmaStat for Windows 2.0, Jandel Scientific, and SigmaPlot for Windows 4.0, Jandel

Scientific, and follow the procedures described in Weitzman & Malabarba (1999:3).

Morphometric data for the type series of each species and non-type specimens are

presented separately in tables. Data for primer types are given between parenthesis along with

the mean and number of specimens counted in the description of each species.

The several geographically isolated populations of Hollandichthys were individually

compared to each other and differences in counts were tested through a Kruskal-Wallis One

Way Analysis of Variance on Ranks, mostly applied to compare dorsal and ventral procurrent

rays counts among all populations. Tests of differences between the sexes of a given species

were performed using original measured lengths. These tests, when indicate differences, are

represented by linear regressions.

6
 Proposed hypotheses of phylogenetic relationship among studied taxa followed the

phylogenetic method of Hennig (1966), as discussed by Wiley (1981). Outgroup comparison

are used to hypothesize ingroup character polarity. Phylogenetic analyses were performed

with Hennig86 (Farris, 1988) coupled with Tree Gardener (Ramos, 1997). All transformation

series were considered unordered, except when noted. Maximum parsimony analyses were

undertaken for each outgroup topology using the ie* algorithm of Hennig86. The hypothesis

was also tested using PAUP (version 3.1.1; Swofford, 1993) and using the Branch and Bound

algorithm. MacClade version 3.1 (Maddison & Maddison, 1992) was used to trace character

evolution. The matrix of character distribution (Table 1) included data from all known species

of Hollandichthys and outgroup characids. A list of comparative material is given at the end

of the text.

The ingroup embrace all Hollandichthys species, representatives of genera previously

proposed as possibly related to Hollandichthys as Nematocharax (in Weitzman et al., 1986)

and Pseudochalceus (in Schultz, 1966; Géry, 1972a, b); and species that share with

Hollandichthys some unusual characters in Characidae (e.g. the longitudinal black stripes).

Additionally, representatives of some putative monophyletic characid Clades were included,

as Aphyocharax Günther (1868) of the Aphyocharacinae, Odontostilbe Cope (1870) of the

Cheirodontinae, Diapoma Cope (1894) of the Glandulocaudinae, and Poptella Eigenmann

(1908) of the Stethaprioninae. Astyanax, Hemibrycon, and Nematobrycon were chosen as

representatives of the Tetragonopterinae. The outgroup incorporated Brycon Müller &

Troschel (1844) and Bryconops Kner (1858), considered hypothetically basal species of the

Characidae that retain a supraorbital bone (Weitzman & Malabarba, 1998).

Ecological and habitat information provided for some species in this paper are based

in field observations, literature information, stomach content analyses conducted during this

study, and personal communications.

7
 Specimens examined belong to the following institutions: ANSP, Academy of Natural

Sciences, Philadelphia; CAS, California Academy of Sciences, San Francisco; DZSJRP,

Departamento de Zoologia da Universidade São José do Rio Preto, São Paulo; FMNH, Field

Museum of Natural History, Chicago; ICNMHN, Instituto de Ciencias Naturales, Museo de

Historia Natural, Universidad Nacional de Colombia, Bogotá; INPA, Instituto Nacional de

Pesquisa da Amazônia, Manaus; MCP, Museu de Ciências e Tecnologia, Pontifícia

Universidade Católica do Rio Grande do Sul, Porto Alegre; MCZ, Museum of Comparative

Zoology, Cambridge; MHNCI, Museu de História Natural de Capão da Imbuia, Curitiba;

MHNG, Muséum d’Histoire Naturelle, Geneva; MHNUC, Museo de Historia Natural de la

Universidad del Cauca, Popayán; MNRJ, Museu Nacional, Rio de Janeiro; MZUSP, Museu

de Zoologia, Universidade de São Paulo, São Paulo; NMW, Naturhistorisches Museum,

Viena; UFRJ, Universidade Federal do Rio de Janeiro, Rio de Janeiro; USNM, National

Museum of Natural History, Smithsonian Institution, Washington D.C. List of examined

specimens of each species includes in this order: catalog number; total number of specimens

of each lot, followed by the number of specimens cleared and stained (c&s) in parentheses,

standard length range, collecting locality, coordinates, and date. Collectors are given only for

type series of each species. List of comparative material includes in this order: catalog

number; total number of specimens of each lot, standard length range, and collecting locality.

RESULTS

Character descriptions

Characters are grouped under separate heading according to the region of the body to

which they are associated, and listed from anterior to posterior regions of body. Consistency

index obtained for each character is given between parenthesis along with their descriptions.

All characters are treated as unordered in the analysis, except for characters 3 and 83.

8
 HEAD SKELETON

1 - Maxillary shape (CI = 1.00). 0 = Different of state 1 (Fig. 2b). 1 = Median toothed portion,

between the fifth and ninth tooth narrower than distal and proximal tips of the maxilla (Fig.

2a).

2 - Posterior tip of maxilla (CI = 1.00). 0 = Narrow. 1 = Enlarged (Fig. 2a).

3 - Maxillary profile (CI = 1.00). 0 = Not curved, maxillary approximately straight (Fig. 2a-

b). 1 = Curved in anterior portion (near toothed portion), with dorsal margin concave and

ventral margin convex. This character was ordered in the analysis.

State 1 is found only in Bryconops species (see Fig. 9, B. inpai Knöppel, Junk & Géry,

1968, in Machado-Allison et al., 1993:3-4 and 9).

4 - Maxillary shape (CI = 1.00). 0 = Different of state 1 (Fig. 2b). 1 = Elongate, anterodorsal

portion convex and posterodorsal portion concave, and all toothed portion smoothly concave

(see species descriptions; Fig. 2a).

5 - Length of maxilla (CI = 0.25). 0 = Very long, reaching posterior tip of infraorbital 2. 1 =

Long, reaching up to 2/3 of infraorbital 2 length.

6 - Anterior portion of maxilla that articulates with premaxilla (CI = 1.00). 0 = Short, broad,

and forming an internal angle of approximately 50 degrees to longitudinal maxillary axis. 1 =

Very long, narrow and forming an internal angle of approximately 25 degrees to longitudinal

maxillary axis.

7 - Dentary-quadrate joint (CI = 0.67). 0 = Situated approximately on the vertical passing

through the center of orbital opening. 1 = Situated anterior to a vertical passing through the

center of orbital opening. 2 = Situated posterior to vertical passing through the center of

orbital opening.

8 - Contact area of metapterygoid and quadrate (CI = 0.29). 0 = Anteroventral region of

metapterygoid and posterodorsal region of quadrate distant from each other (Fig. 3c). 1 =

9
Anteroventral region of metapterygoid and posterodorsal region of quadrate without contact,

with a small space between (Fig. 3b). 2 = Anteroventral region of metapterygoid and

posterodorsal region of quadrate in contact (Fig. 3a).

9 - Shape of the fenestra located between quadrate and metapterygoid bones (CI = 0.25). 0 =

Oval, horizontally elongated (Fig. 3b). 1 = Rounded, approximately circular (Fig. 3a).

10 - Posteroventral region of metapterygoid (CI = 0.25). 0 = With developed concavity in

margin of the posteroventral region of metapterygoid (Fig. 3c). 1 = With an opening almost

completely surrounded by bone or with a foramen, completely surrounded by bone in the

posteroventral region of metapterygoid (Fig. 3a-b).

11 - Dorsal stem of quadrate (CI = 0.50). 0 = Short and broad, smaller than posterior stem

(Fig. 3b). 1 = Elongate, dorsal and posterior branches approximately of same size (Fig. 3a).

12 - Shape of antorbital (CI = 0.67). 0 = Short, anterior portion enlarged, broader than

posterior portion (Fig. 4a). 1 = Elongate, anterior portion broad and posterior portion narrow

or anterior and posterior portions approximately same size (Fig. 4b). 2 = Very short (Fig. 4c).

13 - Supraorbital bone (CI = 1.00). 0 = Present. 1 = Absent.

The presence of the supraorbital is a primitive condition in Characidae, and is found in

species of Brycon, Bryconops, Salminus Agassiz (1829), Triportheus Cope (1872),

Lignobrycon Eigenmann & Myers (1929), Clupeocharax Pearson (1924) and Engraulisoma

Castro (1981). Among examined characids the supraorbital bone is present only in outgroup

species of Brycon and Bryconops. All species of the ingroup lack the supraorbital bone.

14 - Number of infraorbital bones (CI = 1.00). 0 = 6 infraorbitals. 1 = One infraorbital in the

position of the infraorbitals 3 and 4, resulting in a total number of 5 infraorbitals (Fig. 5a-b)

(or 4 infraorbitals in Rachoviscus Myers (1926), that also lack the infraorbital 6, see Figure 5c

and character 15).

10
 In the state 1, presence of 5 infraorbitals can be due to the fusion of third and fourth

infraorbitals, or loss of one of these bones. The hypothesis of fusion (IO3 + IO4) is based in

the position of the third infraorbital, situated in all posteroventral portion of the orbits,

originally occupied by the IO3 and IO4.

15 - Infraorbital 6 (CI = 1.00). 0 = Present (Fig. 5a-b). 1 = Absent (Fig. 5c).

The lost of the IO6 is a derived condition in Characidae, and occurs only in

Rachoviscus and A. anisitsi Eigenmann & Kennedy (1903) (state 1). The sixth infraorbital is

an independent bone located in lateral surface of cranium (just below the frontal) above the

sphenotic spine or pterosphenotic. The infraorbital 6 possess a laterosensory canal that is

connected with the laterosensory canal of the parietal bone.

16 - Laterosensory canal of infraorbital 1 (CI = 0.67). 0 = Present in all extension of

infraorbital 1 (see Fig. 8, B. meeki Eigenmann & Hildebrand, 1918, in Weitzman, 1962:64). 1

= Present only in posterior portion of infraorbital 1 (Fig. 5a). 2 = Absent (Fig. 5b-c).

Species of Hollandichthys have the laterosensory canal of infraorbital 1 located only in

the posterior portion this bone (state 1). The state 0 is found in outgroup, N. venustus

Weitzman, Menezes & Britski (1986), H. dariensis Meek & Hildebrand (1916), A. lineatus,

D. terofali (Géry, 1964), A. anisitsi, P. paraguayensis (Eigenmann, 1907) and O. fugitiva

Cope (1870). State 2 occurs in Pseudochalceus, Rachoviscus, and N. palmeri Eigenmann

(1911).

17 - Laterosensory canal of infraorbital 2 (CI = 1.00). 0 = Present in all extension of

infraorbital 2 (Fig. 5a-b). 1 = Laterosensory canal not closed and present only in posterodorsal

portion of infraorbital 2 (Fig. 5c).

In Rachoviscus species the laterosensory canal of infraorbital 2 is opened along its

length, not forming a tube (state 1).

11
18 - Disposition of infraorbitals 1, 2 and 3 (CI = 1.00). 0 = Infraorbitals 1, 2 and 3 base not

aligned; ventral margins of infraorbitals 1, 2 and 3 forming a curve (Fig. 5a-c). 1 =

Infraorbital 1, 2 and anteroventral portion of infraorbital 3 aligned; ventral margins of

infraorbitals 1, 2 and 3 nearly in a straight line (Fig. 5b).

19 - Shape of infraorbital 2 (CI = 1.00). 0 = Different of state 1 (shape variation considerable

and not triangular as state 1) (Fig. 5a-c). 1 = Triangular (Fig. 5b).

The infraorbital 2 in species of Pseudochalceus is approximately an isosceles triangle.

In other examined characids and in most Characidae the infraorbital 2 has a variable shape.

20 - Rhinosphenoid (CI = 0.33). 0 = Present. 1 = Absent.

Absence of rhinosphenoid was observed in Hollandichthys, Rachoviscus, A. lineatus

and A. anisitsi. Chernoff & Machado-Allison (1990:266) considered the loss of the

rhinosphenoid a potentially derived condition in Ceratobranchia Eigenmann (1914).

21 - Bony projection in anteroventral region of orbitosphenoid (CI = 0.50). 0 = Expanded

(enlarged). 1 = Not expanded (distal tip pointed).

22 - Posteroventral region of orbitosphenoid (CI = 0.33). 0 = Without bony projection and

without a concavity in the ventral profile. 1 = With a bony projection and with a developed

concavity in the ventral profile.

23 - Shape of parasphenoid (CI = 0.50). 0 = Curved (ventral region convex), and crossing

along horizontal orbital diameter. 1 = Nearly straight (ventral region smooth convex), and

crossing below horizontal orbital diameter.

24 - Length of supraoccipital process (CI = 0.50). 0 = Very short, not reaching anterodorsal

region of neural complex. 1 = Short, reaching and/or slightly surpassing the anterodorsal

region of neural complex. 2 = Very long, exceeding the anterodorsal region of neural

complex.

12
25 - Shape of ectopterygoid (dorsal view) (CI = 0.33). 0 = Elongate (Fig. 6). 1 = Short, broad

posterior portion.

26 - Ectopterygoid (dorsal view) (CI = 1.00). 0 = Without lateral bony projection on

mediadistal portion (Fig. 6b). 1 = With lateral bony projection on mediadistal portion (Fig.

6a).

27 - Pterotic spine (CI = 0.25). 0 = Absent or reduced. 1 = Developed.

28 - Anteroventral bony projection of sphenotic (CI = 1.00). 0 = Absent or very short. 1 =

Present.

29 - Shape of the anteroventral bony projection of sphenotic (CI = 0.33). 0 = Short, not

surpassing anterior portion of the hyomandibular-skull joint. 1 = Long, surpassing the anterior

portion of the hyomandibular-skull joint.

30 - Shape of opercle (CI = 0.50). 0 = Opercle usually expanded posteriorly near its median

and lower portion (Fig. 7a). 1 = Opercle vertically elongate and narrow; lower half usually as

wide as upper half (Fig. 7b).

31 - Posterodorsal margin of opercle (CI = 0.50). 0 = Slightly straight or smoothly concave. 1

= Concave.

32 - Space between gill-rakers of first gill arch (CI = 0.50). 0 = Small, approximately equal to

length of gill-rakers base. 1 = Large, 2 to 3 times larger than length of gill-rakers base.

33 - Shape of distal tip of basihyal (CI = 0.33). 0 = Very broad, and basihyal short (Fig. 8a). 1

= narrow, and basihyal long (Fig. 8b).

34 - Number of gill-rakers of first hypobranquial (CI = 0.33). 0 = 3 gill-rakers. 1 = 1 to 2 gill-

rakers.

DORSAL FIN

35 - Posterior profile of the dorsal fin (CI = 1.00). 0 = Not rounded. 1 = Rounded, distal tip

dorsal-fin rays forming a semi circle (see Figs. 23-30).

13
 Specimens of Hollandichthys have the distal tip dorsal fin rays forming a semi circle

(state 1). This character is a synapomorphy for genus, and not found in all taxa of comparative

material. The generalized condition in characids is the occurrence of acute dorsal fin profile.

36 - Anterior dorsal-fin rays in males (CI = 0.50). 0 = Not elongate. 1 = Elongated.

State 1 is found in the larger specimens of all Pseudochalceus species (unidentified

sex; possibly males) and in males of N. venustus (Weitzman et al., 1986).

37 - Small spines on anterior branched dorsal-fin rays of males (CI = 1.00). 0 = Absent. 1 =

Present.

The state 1 is only present in N. venustus (Weitzman et al., 1986:338), dorsal fin with

small bony spinules scattered distally on branched rays two to six. The presence of spinules in

dorsal-fin rays is a derived condition in Characidae.

38 - Number of dorsal-fin pterygiophores and rays (CI = 1.00). 0 = 10 dorsal-fin

pterygiophores and 11 dorsal-fin rays. 1 = 9 dorsal-fin pterygiophores and 9-10 dorsal-fin

rays.

The number of elements in the dorsal-fin skeleton of characids rarely varies. The usual

characid fin-ray count is two anterior unbranched rays plus nine branched rays. This putative

primitive number (ii, 9; state 0) is found among all the hypothetically basal species of the

Characidae (Brycon, Bryconops, Lignobrycon and Triportheus) and majority of the characid

fishes (Malabarba & Weitzman, in press). The derived condition (ii, 8; state 1) is found in H.

dariensis and in D. terofali.

ADIPOSE FIN

39 - Adipose fin (CI = 1.00). 0 = Present. 1 = Absent.

Absence of adipose fin is found only in N. palmeri.

40 - Shape of adipose fin in larger specimens (CI = 1.00). 0 = Elongate (see Figs. 23-28 and

30). 1 = Rounded (see Fig. 29).

14
 Nearly all species of Hollandichthys have adipose fin elongate except in

Hollandichthys sp. n. D that present adipose fin approximately rounded and deep

(autapomorphy of this species). Among comparative material specimens of P. lineatus Kner

(1863) have adipose fin elongate (ANSP 75906), or slightly rounded (MCZ 48730), and R.

crassiceps Myers (1926) have adipose fin slightly rounded (see Fig. 3-4 in Weitzman & Cruz,

1981), but different of the state assigned for Hollandichthys sp. n. D.

PECTORAL FIN

41 - Shape of cartilaginous flap in external lateral surface of first pectoral- and pelvic-fin rays

(CI = 1.00). 0 = Smooth. 1 = Serrated.

The state 1 is found only in Hollandichthys and Pseudochalceus species, and is more

clearly discernible in large specimens.

42 - Laminar bony flange on postcleithrum 3 (CI = 0.50). 0 = Absent (Fig. 9c). 1 = Present

(Figs. 9a-b).

43 - Shape of laminar bony flange on postcleithrum 3 (CI = 1.00). 0 = Smaller than half-

length of postcleithrum 3 (Fig. 9b). 1 = Larger than half-length of postcleithrum 3 (Fig. 9a).

State 1 is present only in Hollandichthys. All species of Hollandichthys have a large

laminar bony flange on postcleithrum 3, and usually located in median portion this bone (state

1). Among comparative material the laminar bony flange can be absent (B. pesu Müller &

Troschel (1845) and B. melanurus (Bloch, 1794)) or with a small bony flange (remaining

taxa).

44 - Anterodorsal bony projection of postcleithrum 1 (CI = 0.25). 0 = Postcleithrum 1

elongate and with bony projection acute (Fig. 10a). 1 = Postcleithrum 1 short, nearly rounded

and with a shorter bony projection (Fig. 10b).

Postcleithrum 1 absent in D. terofali.

15
45 - Coracoid bone (CI = 0.50). 0 = Elongate, longer than deep (bone depth) (Fig. 11b). 1 =

Short and high, and reduced in length (Fig. 11a).

46 - Coracoid foramen located near mesocoracoid and coracoid joint (CI = 1.00). 0 =

Developed, wide (Fig. 11b). 1 = Reduced, small (Fig. 11a).

47 - Shape of posterior laminar projection of cleithrum (lateral view) (CI = 1.00). 0 =

Elongate and narrow (Fig. 12b). 1 = Wide, square approximately (Fig. 12a). 2 = Small and

with acute bony projection (present only in D. terofali).

48 - Posteroventral margin of cleithrum (lateral view) (CI = 0.50). 0 = Reaching and/or

surpassing the base of first pectoral-fin ray (Fig. 12b-c). 1 = Not reaching and distant from

base of first pectoral-fin ray (separate by a gap) (Fig. 12a).

49 - Bony projection from the posterodorsal region of scapula (CI = 0.33). 0 = Short and

wide, not reaching the postcleithrum 2 (Fig. 12b). 1 = Elongate and reaching or near the

postcleithrum 2 (Fig. 12a).

50 - Bony process (second spine) of posttemporal (CI = 0.50). 0 = Narrow and very

elongated. 1 = Short. 2 = Reduced or absent.

51 - Laterosensory canal of extrascapular bone (CI = 1.00). 0 = Presence of one or several

branches in the laterosensory canal. 1 = Extrascapular reduced with an unbranched

laterosensory canal or without laterosensory canal.

52 - Position of the postcleithrum 3 (CI = 1.00). 0 = Postcleithrum 3 obliquous, not

perpendicular to body axis, and distal tip inclined to the anterior region of body. 1 =

Postcleithrum 3 perpendicular to body axis, and distal tip directed to the dorsal region of

body.

In the specimens examined of R. crassiceps the postcleithrum 3 is very elongate,

reaching of postcleithrum 1 base.

16
53 - Coracoid bones (CI = 0.75). 0 = Ventral border longer than dorsal border, with a bony

projection on posteroventral region. 1 = Dorsal and ventral borders nearly equal in length. 2 =

Dorsal border longer than ventral border, with or without a bony projection on posterodorsal

region. 3 = Ventral border extremely expanded (P. paraguayensis).

PELVIC FIN

54 - Anterior pelvic-fin rays of males (CI = 1.00). 0 = Normal, not elongate. 1 = Very

elongated (filamentous rays).

Only N. venustus have anterior pelvic-fin rays greatly elongated, a sexually dimorphic

character used by Weitzman et al. (1986) to recognize males.

55 - First pelvic-fin ray (CI = 1.00). 0 = Unbranched. 1 = Branched (Fig. 13).

56 - Anterior portion of pelvic bone (CI = 0.50). 0 = Narrow, without lateral expansion on the

inner surface of pelvic bone. 1 = Broad, with lateral expansion on the inner surface of pelvic

bone.

57 - Total number of pelvic-fin rays (CI = 0.50). 0 = 8 rays. 1 = 7 rays. 2 = 6 rays.

58 - Bony hooks on anal and pelvic -fins rays of males (CI = 0.50). 0 = Absent. 1 = Present

(Fig. 13).

Bony hooks on anal and pelvic fins rays represent a secondary sexual character usually

found in mature males of many characids (see Azpelicueta & Garcia, 2000:252). Among

comparative material only N. palmeri, A. lineatus, H. dariensis and Pseudochalceus species

have no hooks in anal and pelvic fin rays.

The specimens examined of B. pesu had no hooks on pelvic fins, but these are present

in other Brycon species, according to Lima (2001). Since presence of hooks depends on the

examination of mature specimens, we prefer to assign the state 0 for B. pesu. Also, the only

specimen examined of Bryconops sp. had no hooks, but all other species we have examined

present hooks in unbranched and branched pelvic-fin rays of males. We also assigned state 0

17
in the matrix for this species. Males examined of A. anisitsi have hooks in all pelvic-fin rays.

See more detailed discussion in characters 59 and 64.

59 - Distribution of hooks on pelvic-fin rays of males (CI = 0.80). 0 = Present, starting from

the first branched ray. 1 = Present, starting from the second branched ray. 2 = Present, starting

from the third branched ray (Fig. 13).

Presence of hooks in the pelvic fin is a secondary sexual character useful for

recognition of males in Characidae. The generalized condition in characids is presence of

hooks in the unbranched and branched pelvic-fin rays. All examined males of Hollandichthys

possess hooks from the third to fifth branched rays (usually 3-4 rays). The absence of hooks

in the first and second branched pelvic-fin ray is unique and represents a synapomorphy for

the genus.

Males of N. venustus have hooks on the second to fifth branched pelvic-fin rays

(Weitzman et al., 1986:339). Males of R. crassiceps and R. graciliceps Weitzman & Cruz

(1981) have hooks on the second and third branched pelvic-fin rays (see Weitzman & Cruz,

1981).

Other taxa of the comparative material, including Pseudochalceus species, N. palmeri,

A. lineatus and H. dariensis have no hooks on fins. The examination of adult specimens of

Pseudochalceus species, with well-developed elongated dorsal-, anal-, and pelvic-fin rays,

allows the possible recognition of mature males and the recognition of the putative absence of

hooks in these species. Males of the N. palmeri also have principal and median caudal-fin rays

elongate, but no hooks in the anal and pelvic fin rays, also suggesting the absence of hooks in

this species. We do not have information to analyze if the absence of fin hooks in Astyanax

lineatus and H. dariensis are also characteristic for those species or if it is due to the lack of

mature males among examined specimens.

18
 ANAL FIN

60 - Shape of the last anal-fin rays of males (CI = 0.50). 0 = Not elongate. 1 = Last anal-fin

rays elongate, the penultimate and 2 or 3 immediately anterior rays larger (see Figs. 23-30).

Males of Hollandichthys have 3-4 last anal-fin rays elongate, with the penultimate ray

larger. Males of Pseudochalceus sp. and P. longianalis Géry (1972) have the 5-6 last anal-fin

rays elongate, and penultimate plus 2 or 3 immediately anterior rays larger. In P. lineatus and

P. kyburzi Schultz (1966) it was not possible to observe this character due to the small

number of examined specimens. Mature males of N. palmeri also have 5-7 last anal-fin rays

elongate, but the last three anal-fin rays are smaller than the remaining ones. In other

characids, a similar condition is found in males of Mimagoniates microlepis (Steindachner,

1877).

61 - Anterior anal-fin rays of males (CI = 1.00). 0 = Not elongate. 1 = Elongate, rays

filamentous.

The state 1 is found only mature males of N. venustus. Anal-fin rays of anterior lobe

longer in males than females, reaching posteriorly beyond caudal-fin base (see Fig. 1 in

Weitzman et al., 1986:338).

62 - Skin flap in proximal portion of first anal-fin rays (unbranched to 5-6 branched rays in

larger specimens) (CI = 0.33). 0 = Absent. 1 = Present, little or very developed (Fig. 14).

Larger examined specimens of Hollandichthys possess very developed skin flaps

located in the proximal portion of first anal-fin rays. Pseudochalceus species have little

developed skin flaps. The state 1 also is found in outgroup, N. venustus and A. lineatus.

63 - Profile of anal fin in males (CI = 1.00). 0 = Without concavity in the distal margin

between 20th-25th branched anal-fin rays. 1 = With concavity in the distal margin, formed by

a reduction in length of branched anal-fin rays 20-25 (see Figs. 23-30).

19
 Both Hollandichthys and Pseudochalceus species presents an elongation of the

posterior anal-fin rays, but only males of Hollandichthys have a concavity in the posterior

profile of the anal fin. This character can be associated with a reduction in the length of the

branched rays immediately anterior, to the elongate ones.

64 - Distribution of hooks in the anal-fin rays of males (CI = 1.00). 0 = Present, starting in the

last unbranched and first branched rays along a variable number of branched rays. 1 =

Present, starting in the second branched ray along a variable number of branched rays. 2 =

Present, starting in the third branched ray along a variable number of branched rays.

Presence of hooks in the anal fin is a character widespread in Characidae, showing

some specializations that have been useful in diagnosing some included taxa. The putative

primitive condition regarding their distribution along the anal fin is its presence starting in the

last unbranched and/or first branched anal-fin rays (Malabarba, 1998). Rachoviscus has anal-

fin hooks starting in the second branched ray (Weitzman & Cruz, 1981) (state 1), and

Hollandichthys starting in the third branched ray (state 2), both derived conditions in

Characidae. Malabarba (1998) used the presence of hooks in the anal-fin starting in the

second branched ray as a synapomorphy for Nanocheirodon Malabarba (1998), but this is

clearly homoplastic regarding the condition observed in Rachoviscus. Males of N. venustus

have anal-fin hooks on 3-5 unbranched and on 1-11 branched rays (Weitzman et al.,

1986:338).

The specimens examined of B. pesu had no hooks on fins, but these are present in

other Brycon species, according to Lima (2001). Since presence of hooks depends on the

examination of mature specimens, we prefer to assign the state 0 for B. pesu. Also, the only

specimen examined of Bryconops sp. had no hooks, but all other species we have examined

present hooks in the anal fin of males. We also assigned state 0 in the matrix for this species.

20
 Pseudochalceus species, among examined material, had no hooks on fins. The

presence of apparently adult specimens, however, with well-developed dorsal-, anal-, and

pelvic-fin rays, allows the possible recognition of mature males and the recognition of the

putative absence of hooks in these species. Also it was not observed hooks on pelvic and anal

fins of specimens of N. palmeri, H. dariensis and A. lineatus.

CAUDAL FIN

65 - Small bony spines in the dorsal surface of the proximal portion of dorsal unbranched

principal caudal-fin ray and/or last dorsal procurrent caudal-fin ray; and in the ventral surface

of the proximal portion of ventral unbranched principal caudal-fin ray and/or last ventral

procurrent caudal-fin ray (specimens larger than 60 mm SL) (CI = 1.00). 0 = Absent. 1 =

Present (Fig. 15).

State 1 represents a synapomorphy for Hollandichthys. It is found usually in both

sexes and in specimens larger than 60 mm SL. These spines are more developed in two

populations (MNRJ 20247; MNRJ 10952) of Hollandichthys sp. n. E. Spines were not

observed in the procurrent fin rays of Hollandichthys sp. n. A, Hollandichthys sp. n. B and H.

affinis, and in the principal caudal fin rays of H. affinis.

The state 0 is putatively a primitive condition in Characidae. Presence of spines or

hooks in the caudal fin has been useful in diagnosing some taxa in Glandulocaudinae

(Menezes & Weitzman, 1990; Weitzman & Menezes, 1998) and in the tribe Compsurini of

the Cheirodontinae (Malabarba, 1998; Malabarba & Weitzman, 1999; Malabarba &

Weitzman, 2000). These hooks, however, are usually present in the median caudal-fin rays

and are clearly non-homologous to those observed in Hollandichthys.

66 - Shape of cartilaginous flap in the dorsal surface of dorsal principal unbranched caudal-fin

ray and in the ventral surface of ventral principal unbranched caudal-fin ray (CI = 1.00). 0 =

Smooth. 1 = Serrated (Fig. 16).

21
 The state 1 is found only in species of Hollandichthys, except in Hollandichthys sp. n.

A. In the examined comparative material it is found only a smooth cartilaginous flap in the

dorsal surface of dorsal principal unbranched caudal-fin ray and in the ventral surface of

ventral principal unbranched caudal-fin ray.

67 - Caudal-fin shape (CI = 0.33). 0 = Distal tip of caudal lobes acute. 1 = Distal tip of caudal

lobes rounded (see Figs. 23-30).

Males and females of the Hollandichthys have distal tip of caudal lobes slightly

rounded (state 1). This character also is found in P. kyburzi and R. crassiceps. Males of the N.

palmeri have principal caudal-fin rays elongate, but females and juvenile present caudal lobes

acute (state 0).

68 - Length of the principal caudal-fin rays of males (CI = 1.00). 0 = Central caudal-fin rays

short, smaller than remaining principal caudal-fin rays; dorsal and ventral caudal-fin rays not

excessively elongated. 1 = Central caudal-fin rays elongated, larger than adjacent principal

rays; dorsal and ventral caudal-fin rays elongated, almost twice the length of adjacent rays.

Males of the N. palmeri have principal caudal-fin rays elongate, but females and

juvenile present caudal lobes acute (state 0). This character is an autapomorphy for N.

palmeri.

PROCURRENT CAUDAL-FIN RAYS

69 - Shape of distal tip of the anterior dorsal and ventral procurrent caudal-fin rays (CI =

1.00). 0 = Without ramifications. 1 = With ramifications (Fig. 17).

The ramifications of the distal tip of anterior dorsal and ventral procurrent caudal-fin

rays (usually in 1-6 rays) are found only in the species of Hollandichthys, except in

Hollandichthys sp. n. A. All specimen of the comparative material had no ramifications in the

distal tip of the procurrent caudal-fin rays.

22
70 - Position of the anterior dorsal procurrent caudal-fin rays (CI = 1.00). 0 = Main axis of the

anterior dorsal procurrent caudal-fin rays obliquous to body axis, not perpendicular. 1 = Main

axis of the anterior dorsal procurrent caudal-fin rays perpendicular to body axis (Fig. 17).

The state 1 is found only Hollandichthys, except in species Hollandichthys sp. n. A.

This character is present in both sexes, but is more distinct in mature males. The anterior

dorsal procurrent caudal-fin rays of characids are angled relative to body axis and posteriorly

directed.

71 - Shape of anterior dorsal procurrent caudal-fin rays in males larger than 50 mm SL (CI =

0.50). 0 = Approximately linear or slightly curved. 1 = Sinuous, "S" shaped (Fig. 17).

The state 1 is present in almost all species of Hollandichthys, except Hollandichthys

sp. n. A. In the comparative material only N. palmeri possess anterior dorsal procurrent

caudal-fin rays sinuous.

72 - Procurrent caudal-fin rays in the dorsal and ventral profiles of the caudal peduncle in

large specimens (CI = 0.33). 0 = Not pronounced. 1 = Pronounced (see Fig. 23).

The state 1 is more clearly discernible in larger males of Hollandichthys. It is also

found in P. lineatus, and Rachoviscus species.

73 - Number of dorsal procurrent caudal-fin rays (CI = 1.00) (Fig. 18). 0 = 9-11. 1 = 10-12. 2

= 11-14.

States of meristic characters were treated according to the statistical analyses of its

variation among populations. Data were first analyzed through boxplot graphs (SigmaPlot)

showing the distribution of data according to the 10th, 25th, 75th, 90th percentiles and median of

each analyzed population. Differences among data were tested using Kruskal-Wallis One

Way Analysis of Variance on Ranks (SigmaStat) and considered only for P ≤ 0.001. For those

populations showing no statistical differences, a single state character was assigned. Statistical

differences among groups of geographically continuous populations formed (e.g. rio Maquiné,

23
rio Três Forquilhas and rio Mampituba) were further tested again with the same statistical

procedure. To isolate the group or groups that differ from the others we use All Pairwise

Multiple Comparison Procedures (Dunn's Method).

Sates of this character are analyzed the only among the species of Hollandichthys,

since no information is available regarding its variability in the comparative material.

Although partially overlapping, the dorsal procurrent caudal-fin ray counts are statistically

different when submitted to a Mann-Whitney Ram Sum Test for non-parametric data (P ≤

0.001), as follows:

State 0 (9-11) = [8-10, x = 9.4, n = 20], Hollandichthys sp. n. A.

State 1 (10-12) = [10-12, x = 11.4, n = 32], H. affinis.

State 1 (11-14) = [11-14, x ≥ 12.4], remaining species of Hollandichthys.

74 - Number of ventral procurrent caudal-fin rays (CI = 1.00) (Fig. 19). 0 = 7-9. 1 = 9-12.

This character was used only to analyze the relationships among the species of

Hollandichthys, since no information was available regarding its variability in the

comparative material. See discussion under character 73. States were assigned as follows:

State 0 (7-9) = [7-8, x = 7.7, n = 20], Hollandichthys sp. n. A.

State 1 (9-12) = [9-12, x ≥ 9.4], remaining species of Hollandichthys.

TEETH

75 - Number of premaxillary tooth rows (CI = 0.67). 0 = 3 rows. 1 = 2 rows (Fig. 20b). 2 = 1

row.

Zanata (2000) considered the presence of three tooth rows in premaxilla as one of the

synapomorphies that corroborate the monophyly of Brycon.

76 - Number of teeth in the inner series of premaxilla (CI = 0.50). 0 = 4 teeth. 1 = 5-6 teeth

(Fig. 20b).

24
77 - Symphyseal tooth of inner row of premaxilla (CI = 1.00). 0 = Symphyseal tooth

approximately the same length of adjacent lateral teeth. 1 = Symphyseal tooth larger than

adjacent lateral teeth (Fig. 20b).

In all Hollandichthys species the symphysean tooth is larger than adjacent lateral teeth

(state 1), with its length one and a half times longer than adjacent lateral teeth. This tooth

shape is unique to Hollandichthys species among examined characids and represents a

synapomorphy for Hollandichthys (character not informed for the taxa with one tooth row in

premaxilla).

78 - Tooth shape (CI = 0.50). 0 = Different of state 1 (tooth base enlarged, distal tip not

expanded, with conical cusps; Fig. 20). 1 = Teeth pedunculate, largely expanded, and

compressed distally.

The state 1 represents a derived condition in characids present in cheirodontines.

Nematocharax venustus has teeth pedunculate in inner row of the premaxilla (see Fig. 4 in

Weitzman et al., 1986:341) and O. fugitiva has teeth pedunculate in a single regular tooth row

of premaxilla (see character 55.1 in Malabarba, 1998:228). Both species have teeth

pedunculate in the dentary.

79 - Distribution of maxillary teeth (CI = 0.20). 0 = Teeth only in the anterior portion of

maxilla (less than half-length). 1 = Nearly fully toothed maxilla (more than two thirds of bone

length) (Figs. 2 and 20a).

Species of Hollandichthys and Pseudochalceus, B. pesu, N. venustus, R. crassiceps, H.

dariensis and N. palmeri present an elevated number of maxillary teeth (10 to 23 teeth).

Weitzman et al. (1986:344) considered that a high number of teeth in the maxillary

bone might relate N. venustus, H. multifasciatus and Rachoviscus, but these authors did not

found other possible synapomorphies for these genera. Because of its nearly fully toothed

maxilla H. multifasciatus was considered a species of Pseudochalceus by Schultz (1966:26)

25
and Géry (1972:30). The high maxillary tooth counts seems to have appeared a number of

times in characids. In our analysis, for example, it seems to be independently acquired in our

Clade A, H. dariensis and in the basal species of Brycon.

80 - First maxillary tooth in larger specimens (CI = 1.00). 0 = Tooth with approximately the

same length of remaining teeth (Figs. 2a and 20a). 1 = Tooth much larger than three

subsequent teeth (Fig. 2b).

State 1 represents a synapomorphy for Pseudochalceus. Larger specimens of

Pseudochalceus species have first maxillary tooth much larger than three subsequent teeth.

Maxillary first to fourth tooth in larger specimens of Pseudochalceus decreasing abruptly in

length. In Hollandichthys and in comparative material the anteriormost maxillary tooth is

slightly larger than or has the same size of remaining teeth.

81 - Number of the cusps of maxillary teeth in larger specimens (CI = 0.50). 0 = All teeth

conical. 1 = 1-3 cusps. 2 = Near all teeth tricuspidate (except ultimate and penultimate teeth)

(Figs. 2a and 20a). 3 = More than 3 cusps (tooth with 3-5 cusps, except 2-3 last teeth) (Fig.

2b).

Larger specimens of Hollandichthys have nearly all maxillary teeth tricuspid (state 2),

except 1-2 last teeth (conical or bicuspid). The state 2 also occurs in B. pesu and P.

paraguayensis. Teeth conical are presents only A. anisitsi and Rachoviscus species (state 0).

H. dariensis and N. palmeri possess maxillary teeth with 1-3 cusps (state 1). The state 3 in

found A. lineatus, B. melanurus, Bryconops sp., D. terofali, N. venustus, O. fugitiva and

Pseudochalceus species (see discussion of character 72 in Malabarba, 1998:220)

82 - Disposition of teeth on maxillary (CI = 1.00). 0 = Different of state 1 (teeth of maxillary

not decreasing from 1 to 7) (Fig. 2b). 1 = Teeth gradually decreasing in length from first to 5-

7, and little spaced from each other (Figs. 2a and 20a).

26
 The state 1 is a synapomorphy for Hollandichthys. Specimens of Hollandichthys have

maxillary teeth gradually decreasing in length from first to 5-7 (state 1); remaining teeth are

approximately same size. Larger specimens of Pseudochalceus species have first maxillary

tooth much larger than the three subsequent teeth. Maxillary teeth 1-4 in larger specimens of

Pseudochalceus decreasing abruptly in length. (A. lineatus, Bryconops sp., P. paraguayensis

and O. fugitiva have little teeth in maxillary (state 0)).

83 - Disposition of maxillary teeth in relation of premaxillary teeth (CI = 1.00). 0 = Anterior

maxillary teeth approximately perpendicular to premaxillary teeth. 1 = Anterior maxillary

teeth slightly aligned to premaxillary teeth. This character was ordered in the analysis.

The state 0 is found only in B. melanurus and Bryconops sp. In the species of the

genus Bryconops the articulation of the premaxilla to the maxilla form an angle of 90 degrees

(synapomorphy for genus, Machado-Allison et al., 1993:3-4).

SCALES

84 - Posterior margin of scales from belly region (CI = 1.00). 0 = Rounded or convex margin.

1 = Truncate or concave margin (Fig. 21).

In Characidae scales from belly region possess posterior margin rounded or convex

(state 0). Species of Hollandichthys have body scales laterally covering the belly with

posterior margin truncate or concave (state 1).

COLOR PATTERN

85 - Longitudinal stripes (CI = 0.50). 0 = Absent. 1 = Longitudinal stripes strongly dark

pigmented and clearly defined; scales ventral to the longitudinal scale series that bears the

lateral line not pigmented giving a high contrast to the longitudinal stripes (see Figs. 23-25

and 27-30). 2 = Longitudinal stripes faint pigmented; scales ventral to the longitudinal scale

series that bears the lateral line pigmented along their borders; longitudinal stripes not

conspicuous as in state “1” (Fig. 26).

27
86 - Ontogenetic variation in the number of humeral spots (CI = 1.00) (Table 2). 0 = Absence

of ontogenetic changes in the number of humeral spots. 1 = Presence of two humeral spots in

young specimens and loss of the anterior humeral spot in specimens larger than 60 mm SL. 2

= Presence of two humeral spots in young specimens and loss of both humeral spots in

specimens larger than 60 mm SL.

Specimens of all Hollandichthys species smaller than 50 mm SL have two humeral

spots. Hollandichthys sp. n. A, Hollandichthys sp. n. B and H. affinis have no humeral spot in

specimens larger than 50 mm SL, while the remaining Hollandichthys species maintain the

second humeral spot. In all species of the comparative material it was observed no change in

the number of humeral spots during the ontogenetic development.

87 - Shape of humeral spot in adults (larger than 60 mm SL) (CI = 1.00). 0 = Different of state

1 (Fig. 23). 1 = Large black humeral spot located over 7 to 13-14 lateral line scales and

extending over first longitudinal series of scales above and below lateral line (see Fig. 28).

The state 1 is autapomorphy of Hollandichthys sp. n. C. Others species of

Hollandichthys that maintain the second humeral spot in specimens larger than 60 mm SL

possess humeral spot located over 7 to 9-10 lateral line scales and extending over first

longitudinal series of scales above and below lateral line.

88 - Adipose-fin color pattern of larger specimens (CI = 1.00). 0 = Adipose-fin bordered with

black pigments, usually more intense at adipose-fin base and extending to the dorsal contour

(see Fig. 25). 1 = A conspicuous black spot extending posteriorly from adipose-fin base

through half or 2/3 of the adipose-fin length; absence of a pigmented contour (see Fig. 23).

State 1 is found only in the type-species of Hollandichthys, H. multifasciatus. In the

other species of the genus the adipose fin is pigmented only near its base and along its border.

Among the comparative material, some P. lineatus specimens have a pigmented adipose fin,

28
but it is grayish, showing a degrade pattern from its base to distal portion, lacking a black

pigmented and clearly delimited spot as described in state 1.

89 - Color pattern above anal fin (CI = 1.00). 0 = Body just above anal-fin base weakly

pigmented, usually with faint longitudinal irregular zigzag stripes (see Figs. 23-28 and 30). 1

= Body just above anal-fin base with conspicuous black spots (see Fig. 29).

The state 1 is autapomorphy of Hollandichthys sp. n. D. In other species of

Hollandichthys occur faint longitudinal irregular zigzag stripes above anal-fin base (state 0).

Among the comparative material, Pseudochalceus sp., P. longianalis and P. kyburzi, have

black spots over the lateral body surface, but it is considered a distinct condition of that

described as state 1.

90 - Pigmentation of caudal-fin base (CI = 1.00). 0 = Unpigmented (or different of state 1)

(see Figs. 23-25 and 27-30). 1 = Presence of a faint black spot covering the basal portion of

the median caudal-fin rays (see Fig. 26).

The state 1 is autapomorphy for Hollandichthys sp. n. A. Among the comparative

material several taxa possess pigmented caudal peduncle, but not as described in state 1.

OTHER CHARACTERS

91 - Position of the first supraneural (CI = 1.00). 0 = Positioned before the neural spine of

fourth vertebra. 1 = Positioned after the neural spine of fourth vertebra.

INSEMINATION

92 - Insemination (CI = 0.50). 0 = Absent. 1 = Present.

Most characiform fishes appear to be externally fertilized (Burns et al., 1998). In

Characidae, insemination occurs in Glandulocaudinae (here represented by D. terofali), in a

tribe of Cheirodontinae (Compsurini), and some other characid genera (e.g. Attonitus Vari &

Ortega (2000), Creagrutus Gunther (1864), Knodus Eigenmann (1911), Bryconamericus

pectinatus Vari & Siebert (1990) and Brittanichthys axelrodi Géry (1965)) (Burns et al.,

29
1995; Burns & Weitzman, in press). Within Cheirodontinae, the presence of insemination is a

derived condition and constitutes a putative synapomorphy of the Compsurini (Malabarba,

1998:216). This character is also one of four unequivocal synapomorphies for

Glandulocaudinae (Weitzman & Menezes, 1998:178).

Histological analysis of ovary of some populations of Hollandichthys demonstrates the

presence of spermatozoa within the ovary, and confirm the presence of insemination among

Hollandichthys species (Azevedo, Bertaco & Malabarba, 2002; Burns & Weitzman, in press).

93 - Sperm storage area in testes (CI = 0.50). 0 = Absent. 1 = Present.

Hollandichthys presents the development of a region of the testis for use in sperm

storage (state 1), thus as D. terofali and other glandulocaudines (Weitzman & Menezes,

1998). This modification is found in inseminating species, but also may be present in some

externally fertilizing teleosts (Burns & Weitzman, in press).

94 - Sperm nuclei (CI = 0.50). 0 = Spherical. 1 = Elongate.

Sperm with spherical or slightly ovoid nuclei are often referred to as "aquasperm" and

considered to be the plesiomorphic sperm type for teleosts (Jamieson, 1991). In Characidae

most inseminating species have modification in the spermatic cells, with sperm nuclei

elongate (Burns & Weitzman, in press). The sperm nuclei is elongate in Hollandichthys

(Azevedo et al., 2002), in glandulocaudines (Burns et al., 1995) and inseminating

cheirodontines (Burns et al., 1997; 1998).

95 - Position of flagellum (CI = 1.00). 0 = Flagellum in posterior region. 1 = Flagellum in

anterior region, and folded over nuclei.

Hollandichthys present flagellum in posterior region of nuclei, as most externally

fertilizing species (state 0). Glandulocaudines have flagellum in anterior region, and folded

over nuclei (state 1) (Burns et al., 1998).

30
Phylogenetic relationships of Hollandichthys

The Cladogram of Figure 22 shows the strict consensus tree obtained from six equally

parsimonious trees (CI = 0.55; RI = 0.82) achieved through the analysis of 95 characters from

25 taxa (Table 1). The analysis allows the recognition of Hollandichthys as monophyletic,

including 8 species. Pseudochalceus is also proposed as monophyletic, including 4 species,

and both genera forming a monophyletic Clade. Relationships of the Clade Hollandichthys +

Pseudochalceus lies with another hypothesized Clade formed by Nematobrycon +

(Aphyocharax + Rachoviscus), and the group formed by these two Clades is hypothesized as

sister group to Nematocharax.

Although the relationships above are strongly corroborated by several characters, the

relationships of this large Clade (Nematocharax + ((Nematobrycon + (Aphyocharax +

Rachoviscus)) + (Hollandichthys + Pseudochalceus))) with the remaining comparative

material are weakly supported by several ambiguous characters.

We deserve the discussion here to the relationships of the Clade specified above and

named Clade A in our cladogram. Characters and possible states of the remaining portions of

the cladogram are briefly presented in Table 3.

The six equally parsimonious trees obtained grouped Bryconops melanurus and

Brycon pesu as a basal clade, forming a politomy with Bryconops sp. and with a clade

containing all remaining taxa. Such a hypothesis was supported by several ambiguous

characters at this level of analysis (see Table 3) that do not reflect true relationships among

Bryconops species; it was not made also an effective effort to find characters to solve the

relationships at this level of analysis. We tested an alternative hypothesis, presented in Fig.

22, grouping the two Bryconops species and found a tree just one step longer than the trees

obtained through the analysis. Such a hypothesis changed only ambiguous character

transformations at the most basal portions of the obtained tree, and did not change characters

31
defining Clade A and included clades or taxa. We preferred to present this one step longer

alternative hypothesis, recognizing Bryconops as monophyletic.

CLADE A

(Nematocharax + ((Nematobrycon + (Aphyocharax + Rachoviscus))

+ (Hollandichthys + Pseudochalceus)))

Synapomorphies:

- Posteroventral region of orbitosphenoid with a bony projection and developed concavity in

the ventral profile (Ch. 22.1). This condition was acquired independently by parsimony in D.

terofali and O. fugitiva. The absence of bony projection and concavity (state 0) in N. palmeri

means a reversal.

- Space between gill-rakers of first gill arch large, 2 to 3 times larger than length of gill-rakers

(Ch. 32.1). Acquired independently by parsimony in H. dariensis.

- Presence of 1 to 2 gill-rakers on first hypobranquial (state 1). Acquired independently by

parsimony in H. dariensis, O. fugitiva and Bryconops sp.

- Anterodorsal bony projection of the postcleithrum 1 elongate and acute (Ch. 44.0). Acquired

independently by parsimony in O. fugitiva and B. pesu. The presence of a short, nearly

rounded bony projection (state 1) in the Postcleithrum 1 in Pseudochalceus sp. means a

reversal.

- Maxillary nearly fully toothed (more than 2/3 of maxillary length) (Ch. 79.1). Acquired

independently by parsimony in H. dariensis. The teeth restricted to the anterior portion of the

maxilla (less than half-length) (state 0) in A. anisitsi and R. graciliceps means a reversal.

Ambiguous synapomorphies:

- Applying the Acctran algorithm, presence of bony hooks in the pelvic-fin of males, starting

in the second branched ray (Ch. 59.1) means a synapomorphy for Clade A, with a reversal

32
(Ch. 59.0) in A. anisitsi. In the alternative hypothesis, state 0 is assigned to Clade A and state

1 is assigned as independently derived in N. venustus and Rachoviscus.

Nematocharax venustus Weitzman, Menezes & Britski, 1986

Autapomorphies:

- Anteroventral region of metapterygoid and posterodorsal region of quadrate without contact,

small space between the regions (Ch. 8.1). By parsimony, this character state is assigned as an

autapomorphy for Nematocharax, but is also assigned as independently derived three

additional times in P. lineatus + P. longianalis, in A. lineatus, and Bryconops sp.

- Anterior dorsal-fin rays elongate in males (Ch. 36.1). By parsimony, this condition is also a

synapomorphy for Pseudochalceus Clade.

- Presence of small spines on anterior branched dorsal-fin rays of males (Ch. 37.1).

- Anterior pelvic-fin rays of males great elongated (Ch. 54.1).

- Anterior anal-fin rays elongate in males (Ch. 61.1).

- Presence of teeth pedunculate, largely expanded, and compressed distally on premaxilla

(Ch. 78.1). Acquired independently in O. fugitiva.

CLADE B

((Nematobrycon + (Aphyocharax + Rachoviscus))

+ (Hollandichthys + Pseudochalceus)))

Synapomorphies:

- Number of infraorbital bones. One infraorbital in the position of the infraorbitals 3 and 4,

resulting in a total number of 5 infraorbitals or 4 infraorbitals (in Rachoviscus only, that also

lack the infraorbital 6) (Ch. 14.1). See comments in character description.

- Absence of the laterosensory canal of infraorbital 1 (Ch. 16.2). By parsimony, presence of

33
the laterosensory canal in all extension of infraorbital 1 (state 0) in A. anisitsi a means

reversal, and presence only in posterior portion of infraorbital 1 (state 1) a secondary

condition acquired independently in Hollandichthys Clade.

- Anteroventral bony projection of sphenotic short, not surpassing anterior portion of the

hyomandibular-skull joint (Ch. 29.0), acquired independently in H. dariensis, D. terofali and

O. fugitiva. By parsimony, bony projection of sphenotic long (state 1) means a reversal in

Pseudochalceus sp. + P. kyburzi.

- Coracoid bone with dorsal border longer than ventral border, with or without a bony

projection on posterodorsal region (Ch. 53.2). Coracoid bone with ventral border longer than

dorsal border, and with a bony projection on posteroventral region (state 0) in A. anisitsi a

means reversal.

Ambiguous synapomorphies:

- Applying the Acctran algorithm, 7 rays in the pelvic-fin (Ch. 57.1) means a synapomorphy

for Clade B, with a reversal (Ch. 57.0) in Pseudochalceus. Otherwise, state 0 is assigned to

this Clade and state 1 is assigned as independently derived in Clade C and Hollandichthys.

- Applying the Acctran algorithm, last anal-fin rays elongate, the penultimate and 2 or 3

anterior rays largest (Ch. 60.1) means a synapomorphy for Clade B, with a reversal (Ch.

60.0) in Clade D. Otherwise, state 0 is assigned to this Clade and state 1 is assigned as

independently derived in Clade E and N. palmeri.

CLADE C

(Nematobrycon + (Aphyocharax + Rachoviscus)

Synapomorphies:

- Supraoccipital process short, reaching and/or slightly surpassing the anterodorsal region of

neural complex (Ch. 24.1). Supraoccipital very short, not reaching anterodorsal region of

34
neural complex (state 0) in Rachoviscus Clade means reversal.

- Absence of the skin flap in proximal portion of first anal-fin rays (unbranched to 5-6

branched rays in larger specimens) (Ch. 62.0). This derived condition is also assigned as

acquired independently in P. paraguayensis, D. terofali and O. fugitiva.

Ambiguous synapomorphies:

- Metapterygoid bone with developed concavity in margin of the posteroventral region (Ch.

10.0). It is equally parsimonious to accept a reversal to state 0 as a synapomorphy of Clade C

with the re-acquisition of state 1 in A. anisitsi, or the independent reversals of this character to

state 0 in N. palmeri and Rachoviscus Clade.

- Bony process of posttemporal reduced or absent. (Ch. 50.2) is found exclusively in Clade C,

but multiple equally parsimonious hypothesis of character transformation can be assumed in

this case. States 0 or 1 can be assumed as a synapomorphy for Clade B and state 2 as an

exclusive synapomorphy of Clade C, or state 2 as a synapomorphy of Clade B, with a reversal

to state 0 in Clade E.

- Seven pelvic-fin rays (Ch. 57.1). See discussion under Clade B.

- Maxillary teeth conical (Ch. 81.0). It is equally parsimonious to accept a reversal to state 0

as a synapomorphy of Clade C with the acquisition of state 1 in N. palmeri, the acquisition of

state 1 as a synapomorphy of Clade C with the reversal to state 0 in Clade D; or the

independent reversal to state 0 in Clade D and acquisition of state 1 in N. palmeri.

Nematobrycon palmeri Eigenmann, 1911

Autapomorphies:

- Posteroventral region of orbitosphenoid without bony projection and without a concavity in

the ventral profile (Ch. 22.0). By parsimony, this character state is assigned as an

autapomorphy for Nematobrycon, but is also assigned as acquired independently in P.

35
paraguayensis, A. lineatus, H. dariensis and in outgroup.

- Adipose fin absent (Ch. 39.1).

- Absence of bony hooks on anal and pelvic -fins rays of males (Ch. 58.0). This derived

condition is also assigned as a synapomorphy for Pseudochalceus Clade.

- Length of the principal caudal-fin rays of males. Central caudal-fin rays elongated, larger

than adjacent principal rays; dorsal and ventral caudal-fin rays elongated, almost twice the

length of adjacent rays (Ch. 68.1).

- Anterior dorsal procurrent caudal-fin rays in males sinuous, "S" shaped (Ch. 71.1). This

derived condition also found in most species of Hollandichthys, except in Hollandichthys sp.

n. A.

- Four teeth in the inner row of premaxilla (Ch. 76.0). Acquired independently in H.

dariensis and D. terofali.

Ambiguous autapomorphies:

- Metapterygoid bone with developed concavity in margin of the posteroventral region (Ch.

10.0). See discussion under Clade C.

- Five and six last anal-fin rays elongate, larger penultimate and 2 or 3 anterior rays (Ch.

60.1). See discussion under Clade B.

- Maxillary teeth 1-3 cusps (Ch. 81.1). See discussion under Clade C.

CLADE D

(Aphyocharax + Rachoviscus)

Synapomorphies:

- Infraorbital 6 absent (Ch. 15.1).

- Rhinosphenoid bone absent (Ch. 20.1).

Ambiguous synapomorphies:

36
- Last anal-fin rays of males not elongate (Ch. 60.0). See discussion under Clade B.

- Maxillary with teeth only in anterior portion (-1/2 of length) (Ch. 79.0). It is equally

parsimonious to accept a reversal to state 0 as a synapomorphy of Clade D with the re-

acquisition of state 1 in R. crassiceps, or the independent reversals of this character to state 0

in A. anisitsi and R. graciliceps.

- Maxillary teeth conical (Ch. 81.0). See discussion under Clade C.

Aphyocharax anisitsi Eigenmann & Kennedy, 1903

Autapomorphies:

- Laterosensory canal present in all extension of infraorbital 1 (Ch. 16.0). This character state

represents a reversion in A. anisitsi.

- Parasphenoid bone nearly straight (ventral region smooth convex), and crossing below

horizontal orbital diameter (Ch. 23.1). Acquired independently in Hollandichthys.

- Coracoid bone elongated, longer than deep (Ch. 45.0).

- Coracoid bones with ventral border longer than dorsal border, and with a bony projection on

posteroventral region (Ch. 53.0). By parsimony acquired independently in H. dariensis, D.

terofali, O. fugitiva and outgroup.

- Presence of 1 teeth row in premaxilla (Ch. 75.2). Acquired independently in O. fugitiva.

Ambiguous autapomorphies:

- Posteroventral region of metapterygoid with an opening completely surrounded by bone

(foramen) (Ch. 10.1). See discussion under Clade C.

- Applying the Acctran algorithm, presence of bony hooks starting in the first branched

pelvic-fin rays of males (Ch. 59.0) means a reversal in A. anisitsi. See discussion under Clade

A.

37
- Maxillary with teeth only in the anterior portion (-1/2 of length) (Ch. 79.0). See discussion

under Clade D.

Rachoviscus Clade

Synapomorphies:

- Antorbital bone very short (Ch. 12.2).

- Laterosensory canal not closed and situated only in posterodorsal portion of infraorbital 2

(Ch. 17.1).

- Supraoccipital process very short, not reaching anterodorsal region of neural complex (Ch.

24.0). Acquired independently in B. pesu.

- Ectopterygoid bone short, posterior portion broad (Ch. 25.1). Acquired independently in A.

lineatus and D. terofali.

- Posteroventral margin of cleithrum not reaching and distant from base of first pectoral-fin

ray (separate by a gap) (Ch. 48.1). Acquired independently in Hollandichthys.

- Bony projection from the posterodorsal region of scapula elongate and reaching incontactor

near the postcleithrum 2 (Ch. 49.1). Acquired independently in H. dariensis, O. fugitiva and

Hollandichthys.

- Laterosensory canal of extrascapular bone reduced with an unbranched laterosensory canal

or without laterosensory canal (Ch. 51.1).

- Bony hooks starting in the second branched anal-fin ray along a variable number of

branched anal-fin rays (Ch. 64.1).

- Procurrent caudal-fin rays in the dorsal and ventral profiles of the caudal peduncle

pronounced in large specimens (Ch. 72.1). Acquired independently in P. lineatus and

Hollandichthys.

Ambiguous autapomorphies:

38
- Metapterygoid bone with developed concavity in margin of the posteroventral region (Ch.

10.0). See discussion under Clade C.

Rachoviscus crassiceps Myers, 1926

Autapomorphies:

- Maxillary very long, reaching posterior tip of infraorbital 2 (Ch. 5.0). This character state is

also assigned as acquired independently four additional times in Hollandichthys Clade, H.

dariensis, B. melanurus and B. pesu.

- Six pelvic-fin rays (Ch. 57.2).

- Distal tip of caudal fin lobes rounded (Ch. 67.1). Acquired independently in P. kyburzi and

Hollandichthys Clade.

- Nearly fully toothed maxillary (+ 2/3 of length) (Ch. 79.1). See discussion under Clade D.

Rachoviscus graciliceps Weitzman & Cruz, 1981

Autapomorphies:

- Maxillary with teeth only in the anterior portion (-1/2 of length) (Ch. 79.0). See discussion

under Clade D.

CLADE E

(Hollandichthys + Pseudochalceus)

Synapomorphies:

- Maxilla very long, reaching posterior tip of infraorbital 2 (Ch. 5.0). This character state is

also assigned as acquired independently four additional times in R. crassiceps, H. dariensis,

B. melanurus and B. pesu.

- Dentary-quadrate joint situated posterior to vertical passing through vertical orbital diameter

39
(Ch. 7.2).

- Anteroventral region of metapterygoid and posterodorsal region of quadrate in contact (Ch.

8.2). These regions without contact (state 1) represent a reversal in P. lineatus + P.

longianalis.

- Fenestra between quadrate and metapterygoid bones rounded, approximately circular (Ch.

9.1). By parsimony acquired independently in A. lineatus and B. melanurus. The presence of

an oval fenestra horizontally elongated (state 0) in P. kyburzi represent a reversion.

- Dorsal stem of quadrate elongated, dorsal and posterior branches approximately of same size

(Ch. 11.1). Acquired independently in A. lineatus.

- Cartilaginous flap serrated in external lateral surface of first pectoral- and pelvic- fin rays

(Ch. 41.1).

Ambiguous synapomorphies:

- Bony process of posttemporal narrow and very elongated (Ch. 50.0). See discussion under

Clade C.

- Five to six last anal-fin rays elongate, larger penultimate and 2 or 3 anterior rays (Ch. 60.1).

See discussion under Clade B.

Pseudochalceus Clade

Synapomorphies:

- Disposition of infraorbitals 1, 2 and 3. Infraorbital 1, 2 and anteroventral portion of

infraorbital 3 aligned, ventral margins of infraorbitals 1, 2 and 3 nearly in a straight line (Ch.

18.1).

- Infraorbital 2 approximately triangular (Ch. 19.1).

- Opercle vertically elongate and narrow; lower half usually as wide as upper half (Ch. 30.1).

Independently acquired in P. paraguayensis.

40
- Anterior of dorsal-fin rays in males elongated (Ch. 36.1). Independently acquired in N.

venustus. See character 36 description.

- Absence the bony hooks on anal and pelvic -fins rays of males (Ch. 58.0). Acquired

independently in N. venustus.

- First maxillary tooth larger than three subsequent teeth in larger specimens (Ch. 80.1).

Ambiguous synapomorphies:

- Eight pelvic-fin rays (Ch. 57.0). See discussion under Clade B.

P. lineatus + P. longianalis

Synapomorphy:

- Anteroventral region of metapterygoid and posterodorsal region of quadrate without contact,

and with small space between the regions (Ch. 8.1). Independently acquired in N. venustus, A.

lineatus and Bryconops sp.

Pseudochalceus sp. + P. kyburzi

Synapomorphy:

- Anteroventral bony projection of sphenotic long, surpassing the anterior portion of the

hyomandibular-skull joint (Ch. 29.1). See discussion under Clade B.

Pseudochalceus lineatus Kner & Steindachner, 1863

Autapomorphies:

- Procurrent caudal-fin rays in the dorsal and ventral profiles of the caudal peduncle in large

specimens pronounced (Ch. 72.1). By parsimony is also synapomorphies for Hollandichthys

Clade and Rachoviscus Clade.

- Longitudinal stripes strongly dark pigmented and clearly defined; scales ventral to the

41
longitudinal scale series that bears the lateral line not pigmented giving a high contrast to the

longitudinal stripes (Ch. 85.1). Independently acquired in A. lineatus and Hollandichthys

Clade.

Pseudochalceus longianalis Géry, 1972

- No autapomorphies found.

Pseudochalceus sp.

Autapomorphy:

- Postcleithrum 1 short, nearly rounded and with a shorter bony projection in the anterodorsal

portion (Ch. 44.1). This character state is acquired independently in P. paraguayensis, A.

lineatus, H. dariensis and Bryconops species.

Pseudochalceus kyburzi Schultz, 1966

Autapomorphies:

- Fenestra quadrate-metapterygoid oval, horizontally elongated (Ch. 9.0). See discussion

under Clade E.

- Pterotic spine developed (Ch. 27.1). By parsimony, this character state was acquired

independently in Hollandichthys, A. lineatus, H. dariensis, D. terofali and outgroup.

- Distal tip of caudal-fin lobes rounded (Ch. 67.1). This derived condition is independent

acquisition in Hollandichthys and R. crassiceps.

Hollandichthys Clade

Synapomorphies:

- Median toothed portion of the maxilla, between the fifth and ninth teeth narrower than distal

and proximal tips of the maxilla (Ch. 1.1; Figs. 2a and 20a).

42
- Maxillary elongated, anterodorsal portion convex and posterodorsal portion concave, and all

toothed portion smoothly concave (Ch. 4.1; Figs. 2a and 20a).

- Laterosensory canal of infraorbital 1 present only in posterior portion of infraorbital 1 (Ch.

16.1; Fig. 5a).

- Rhinosphenoid bone absent (Ch. 20.1). Acquired independently in Clade D and A. lineatus.

- Parasphenoid bone nearly straight (ventral region smooth convex), and crossing below

horizontal orbital diameter (Ch. 23.1). Acquired independently in A. anisitsi.

- Ectopterygoid bone with lateral bony projection on mediadistal portion (Ch. 26.1; Fig. 6a).

- Pterotic spine developed (Ch. 27.1). Acquired independently in P. kyburzi, A. lineatus, H.

dariensis, D. terofali and outgroup.

- Distal tip of basihyal very broad, and basihyal short (Ch. 33.0; Fig. 8a). Acquired

independently in O. fugitiva and Bryconops sp.

- Posterior profile of the dorsal fin rounded, distal tip of dorsal-fin rays forming a semi circle

(Ch. 35.1; Figs. 23-30).

- Laminar bony flange of postcleithrum 3 larger than half-length of postcleithrum 3 (Ch. 43.1;

Fig. 9a).

- Posterior laminar projection of cleithrum wide, square approximately (Ch. 47.1; Fig. 12a).

- Posteroventral margin of cleithrum not reaching and distant from base of first pectoral-fin

ray, separate by a gap (Ch. 48.1; Fig. 12a). Acquired independently in Rachoviscus Clade.

- Bony projection from the posterodorsal region of scapula elongate and reaching or near the

postcleithrum 2 (Ch. 49.1). Acquired independently in Rachoviscus Clade, H. dariensis and

O. fugitiva.

- First pelvic-fin ray branched (Ch. 55.1; Fig. 13).

- Anal fin of the males with concavity in the distal margin, formed by a reduction in length of

branched anal-fin rays 20-25 (Ch. 63.1; Figs. 23-30).

43
- Presence of small bony spines in the dorsal surface of the proximal portion of dorsal

unbranched principal caudal-fin ray and/or last dorsal procurrent caudal-fin rays and in the

ventral surface of the proximal portion of ventral unbranched principal caudal-fin ray and/or

last ventral procurrent caudal-fin rays (specimens larger than 60 mm SL) (Ch. 65.1; Fig. 15).

- Distal tip of caudal-fin lobes slightly rounded (Ch. 67.1; Figs. 23-30). Acquired

independently in P. kyburzi and R. crassiceps.

- Procurrent caudal-fin rays in the dorsal and ventral profiles of the caudal peduncle in large

specimens pronounced (Ch. 72.1; Fig. 23). Acquired independently in P. lineatus and

Rachoviscus Clade.

- Symphyseal tooth of inner row of premaxilla larger than subsequent teeth (Ch. 77.1; Fig.

20b).

- Near all maxillary teeth tricuspidate, except 1-2 teeth last (Ch. 81.2; Figs. 2a and 20a).

Acquired independently in P. paraguayensis and B. pesu.

- Teeth of the maxillary gradually decreasing in size from first to 5-7, and little spaced from

each other (Ch. 82.1; Figs. 2a and 20a).

- Posterior margin of scales from belly region truncate or concave (Ch. 84.1; Fig. 21).

- Ontogenetic variation in the number of humeral spots. Presence of two humeral spots in

young specimens and loss of both humeral spots in specimens larger than 60 mm SL (Ch.

86.2; Table 2).

- Insemination present (Ch. 92.1). Acquired independently in D. terofali.

- Present of sperm storage area in testes (Ch. 93.1). Acquired independently in D. terofali.

- Sperm nuclei elongate (Ch. 94.1). Acquired independently in D. terofali.

Ambiguous synapomorphies:

- Seven pelvic-fin rays (Ch. 57.1). See discussion under Clade B.

44
- Presence of hooks in pelvic fin starting in the third branched ray (Ch. 59.2; Fig. 13). Since

no hooks are present in Pseudochalceus species, this character is treated as no informed for

this genus. By parsimony, state 2 can be assumed as present in the ancestor of both genera

Hollandichthys and Pseudochalceus, or acquired only in the ancestor of Hollandichthys.

Since it is not observable in Pseudochalceus, we prefer to list it as a synapomorphy for

Hollandichthys.

- Bony hooks starting in the third branched anal-fin ray along a variable number of branched

anal-fin rays (Ch. 64.2). It is the same situation of the Character 59 above. Since hooks are

not observable in Pseudochalceus, we prefer to list it as a synapomorphy for Hollandichthys.

- Presence of longitudinal stripes is a synapomorphy for Hollandichthys and independently

derived from those observed in P. lineatus and A. lineatus (Ch. 85). Hollandichthys species,

however, presents two different states for this character that allows multiple equally

parsimonious hypothesis of character transformation among the species of the genus. States 1

or 2 can be assumed as a synapomorphy for Hollandichthys Clade, with subsequent changes

do states 2 and 1 respectively, in Hollandichthys sp. n. A and in Hollandichthys Clade H1, or

states 1 and 2 can be assumed as independently derived in Hollandichthys Clade H1 and

Hollandichthys sp. n. A, respectively. We reject the last hypothesis, since homology of the

longitudinal stripes present in Hollandichthys Clade H1 and Hollandichthys sp. n. A is

evident.

Phylogenetic relationships of Hollandichthys species

The analysis of the relationships among the recognized species of Hollandichthys

presents a partial resolution, with the recognition of Hollandichthys sp. n. A as sister group to

an internal large Clade including the remaining species, named Clade H1 (Fig. 22). This can

also be split in an internal Clade including 5 species, named Clade H2, forming a politomy

45
with H. affinis and Hollandichthys sp. n. B. No further resolution was found. There are

however unambiguous autapomorphies that allows the recognition of Hollandichthys sp. n. A,

Hollandichthys sp. n. C, H. multifasciatus and Hollandichthys sp. n. D.

Hollandichthys sp. n. A

Autapomorphy:

- Presence of a faint black spot covering the basal portion of the median caudal-fin rays (Ch.

90.1; Fig. 26).

Ambiguous autapomorphies:

- Presence of 9-11 dorsal procurrent caudal-fin rays (Ch. 73.0; Fig. 18). No states of this

character were assigned to the comparative material, outside Hollandichthys, due to the

examination of a smaller number of specimens of those taxa. By parsimony, all states

assigned to this character (0, 1 or 2), can be assumed as a synapomorphy of Hollandichthys,

with subsequent state changes in Hollandichthys sp. n. A, H. affinis, Hollandichthys sp. n. B,

Clade H1 and/or Clade H2.

- Presence of 7-9 ventral procurrent caudal-fin rays (Ch. 74.0; Fig. 19). No states of this

character were assigned to the comparative material, outside Hollandichthys, due to the

examination of a smaller number of specimens of those taxa. By parsimony, both states 0 or 1

can be assumed as a synapomorphy of Hollandichthys, with a subsequent change to state 0 in

Hollandichthys sp. n. A or to state 1 in Clade H1. It is also equally parsimonious to accept the

independent acquisition of state 0 and 1 in Hollandichthys sp. n. A and Clade H1.

- Presence of the longitudinal stripes faint pigmented; scales ventral to the longitudinal scale

series that bears the lateral line pigmented along their borders; longitudinal stripes not

conspicuous as in state “1” (Ch. 85.2; Fig. 26). See discussion under Hollandichthys.

46
 Hollandichthys Clade H1

Synapomorphies:

- Cartilaginous flap serrated in the dorsal surface of dorsal principal unbranched caudal-fin

ray and in the ventral surface of ventral principal unbranched caudal-fin ray (Ch. 66.1; Fig.

16).

- Distal tip of anterior dorsal and ventral procurrent caudal-fin rays with ramifications (Ch.

69.1; Fig. 17).

- Main axis of the anterior dorsal procurrent caudal-fin rays perpendicular to body axis (Ch.

70.1; Fig. 17).

- Anterior dorsal procurrent caudal-fin rays sinuous ("S" shaped) in males larger than 50 mm

SL (Ch. 71.1; Fig. 17).

Ambiguous synapomorphies:

- Both states 1 or 2 of Ch. 73 can be assumed as a synapomorphy for this Clade. See

discussion under Hollandichthys sp. n. A.

- Presence of 9-12 ventral procurrent caudal-fin rays (Ch. 74.1; Fig. 19). See discussion under

Hollandichthys sp. n. A.

- Presence of the longitudinal stripes strongly dark pigmented and clearly defined; scales

ventral to the longitudinal scale series that bears the lateral line not pigmented giving a high

contrast to the longitudinal stripes (Ch. 85.1). See discussion under Hollandichthys.

Hollandichthys sp. n. B

- No exclusive autapomorphy is found. See the diagnosis of this species.

Ambiguous autapomorphy:

- Presence of 11-14 dorsal procurrent caudal-fin rays (Ch. 73.2; Fig. 18). See discussion

under Hollandichthys sp. n. A.

47
 Hollandichthys affinis (Steindachner, 1908)

- No exclusive autapomorphy is found. See the diagnosis of this species.

Ambiguous autapomorphy:

- Presence of 10-12 dorsal procurrent caudal-fin rays (Ch. 73.1; Fig. 18). See discussion

under Hollandichthys sp. n. A.

Hollandichthys Clade H2

Synapomorphy:

- Ontogenetic variation in the number of humeral spots. Presence of two humeral spots in

young specimens and loss of the first humeral spot in specimens larger than 60 mm SL (Ch.

86.1; Table 2).

Ambiguous synapomorphy:

- Presence of 11-14 dorsal procurrent caudal-fin rays (Ch. 73.2; Fig. 18). See discussion

under Hollandichthys sp. n. A.

Hollandichthys sp. n. C

Autapomorphy:

- Presence of the large black humeral spot located over 7 to 13-14 lateral line scales and

extending over first longitudinal series of scales above and below lateral line (Ch. 87.1; Fig.

28).

Hollandichthys perstriatus (Miranda-Ribeiro, 1908)

- No autapomorphy is found. See the diagnosis of this species.

48
 Hollandichthys multifasciatus (Eigenmann & Norris, 1900)

Autapomorphy:

- Presence of the conspicuous black spot extending posteriorly from adipose-fin base through

half or 2/3 of the adipose-fin length; absence of a pigmented contour (Ch. 88.1; Fig. 23).

Hollandichthys sp. n. D

Autapomorphy:

- Color pattern above anal-fin base with conspicuous black spots (Ch. 89. 1; Fig. 29).

- Adipose fin approximately rounded (Ch. 40. 1; Fig. 29).

Hollandichthys sp. n. E

- No autapomorphy is found. See the diagnosis of this species.

Taxonomic revision of Hollandichthys

Hollandichthys Eigenmann

Hollandichthys Eigenmann, 1909:257 (diagnosed in key), 338 and 374 (lists one species for

the genus, Hollandichthys multifasciatus (Eigenmann & Norris, 1900)), 374 (lists

"Pseudochalceus perstriatus Rebeira and Pseudochalceus affinis Steindachner” as

synonyms) [type-species Hollandichthys multifasciatus by monotypy].

- Eigenmann, 1910:432 (designates Hollandichthys multifasciatus (Eigenmann &

Norris, 1900) as type-species and repeats synonym of Eigenmann, 1909).

- Eigenmann, 1921:225-226 (genus description).

- Schultz, 1966:26 (Hollandichthys as a junior synonym of Pseudochalceus Kner,

1863).

49
 - Weitzman, Menezes & Britski, 1986:344 (revalidate Hollandichthys).

Diagnosis. Hollandichthys is diagnosed from remaining characid genera by the several

synapomorphies (see synapomorphies list above). The following characters are exclusive of

Hollandichthys: median toothed portion of the maxilla, between the fifth and ninth teeth

narrower than distal and proximal tips of the maxilla (Ch. 1.1; Fig. 2a); maxillary elongated,

anterodorsal portion convex and posterodorsal portion concave, and all toothed portion

smoothly concave (Ch. 4.1; Figs. 2a and 20a); first pelvic-fin ray branched (Ch. 55.1; Fig.

13); anal fin of the males with concavity in the distal margin, formed by a reduction in length

of branched anal-fin rays 20-25 (Ch. 63.1; Figs. 23-30); presence of small bony spines in the

dorsal surface of the proximal portion of dorsal unbranched principal caudal-fin ray and/or

last dorsal procurrent caudal-fin rays and in the ventral surface of the proximal portion of

ventral unbranched principal caudal-fin ray and/or last ventral procurrent caudal-fin rays

(specimens larger than 60 mm SL) (Ch. 65.1; Fig. 15); symphyseal tooth of inner row of

premaxilla larger than subsequent teeth (Ch. 77.1; Fig. 20b); teeth of the maxillary gradually

decreasing in size from first to 5-7, and little spaced from each other (Ch. 82.1; Figs. 2a and

20a); posterior margin of scales from belly region truncate or concave (Ch. 84.1; Fig. 21).

The following characters, although not exclusive of Hollandichthys, also distinguish

the species of this genus from other characids: posterior profile of the dorsal fin rounded (Ch.

35.1); presence of longitudinal stripes in the lateral body (Ch. 85.1/2); presence of the

insemination (Ch. 92.1); presence of sperm storage area in testes (Ch. 93.1); and sperm nuclei

elongate (Ch. 94.1).

50
Distribution. All species of the genus Hollandichthys are allopatric and inhabit coastal rivers

and marine islands from Northern Rio Grande do Sul State to Southern Rio de Janeiro State,

and the upper rio Tietê in the continental upper rio Paraná drainage (Fig. 1).

Habitat description. Populations of Hollandichthys species live in small creeks or lateral

puddles with poor current, deepness, black or dim water, ooze and leaf bottom, and with very

dense riparian vegetation, in streams and rivers of the Atlantic Forest. All species were found

only in the remaining portions of this Bioma. Some populations (H. affinis, Ilha de São

Francisco do Sul, SC and Hollandichthys sp. n. D, São Sebastião, SP), were collected in

brackish water together with Awaous tajasica (Lichtenstein, 1819), Eleotris pisonis (Gmelin,

1789) and Oostethus lineatus (Kaup, 1856). According to Britski (1972) H. multifasciatus

already are caught in brackish water and can be the Characoidei more resistant the saltiness

variation.

Sexual dimorphism. Males of Hollandichthys are easily recognized by the presence of bony

hooks in the branched anal- and pelvic-fin rays (Fig. 13), completely absent in females. Hooks

are distributed between the third to 29th branched anal-fin rays (usually 5-22), and third to

fifth (usually 3-4) branched pelvic-fin rays. Hooks are more numerous on 7th to 22nd anal-fin

branched rays.

Males and females also slightly differ in pectoral and pelvic-fin lengths, and in the

anal-fin shape (Figs. 23-30). Females present anal-fin profile nearly straight. Males show

anal-fin profile nearly straight except for a concavity corresponding approximately to 20th-27th

ray; three or fourth last branched anal-fin rays are longer than the remaining, being the

penultimate ray the longest (Figs. 23-30). Caudal peduncle depth is slightly larger in males

than females, and procurrent caudal-fin rays are more prominent in males.

51
Common names. Brazil: “Lambari-listrado”(Miranda-Ribeiro, 1908); "Maria-mole" (Região

da Juréia); "Peixe-folha" (Vale do Ribeira).

Key to the species of Hollandichthys Eigenmann, 1909

1a. Presence of one black humeral spot in specimens larger than 60.0 mm SL (Table 2)........ 2

1b. Absence of humeral spots in specimens larger than 60.0 mm SL (Table 2)........................6

2a. Adipose-fin with one black pigmented along usually proximal half length (Fig. 23;

tributaries of the upper rio Tietê drainage, and coastal streams from Cubatão and Santos, SP)

..........................................................................................................................H. multifasciatus

2b. Adipose-fin not black pigmented along half proximal length, sometimes black pigmented

along its border............................................................................................................................3

3a. The body color pattern just above anal fin covered with small black dots instead of

longitudinal rows; adipose fin slightly rounded (Fig. 29; coastal streams from Bertioga to São

Sebastião, SP)....................................................................................... Hollandichthys sp. n. D

3b. The body color pattern above anal fin with faint longitudinal rows; adipose fin oval,

elongated.....................................................................................................................................4

4a. Presence of one large black humeral spot located over 7 to 13-14 lateral line scales and

extending over first longitudinal series of scales above and below lateral line (Fig. 28); orbital

diameter 27.7-34.0% of HL, mean = 30.3%) (rio Morato and rio Braço do Macaco,

Guaraqueçaba, PR)............................................................................... Hollandichthys sp. n. C

52
4b. Presence of one black humeral spot located over 7-10 lateral line scales and extending

over first longitudinal series of scales above lateral line............................................................5

5a. All fins red-orange pigmented at least in their tips; larger number scales in longitudinal

series 40-46, x = 42.5; presence of very developed spines in the dorsal surface of dorsal

unbranched caudal-fin ray (Fig. 15), and in the ventral surface of the ventral unbranched

principal caudal-fin ray in larger specimens; bony hooks present in the dorsal surface of last

dorsal procurrent ray and in the ventral surface of last ventral procurrent ray in specimens

larger than 50.0 mm SL (coastal streams from Ilhabela, SP to Mangaratiba,

RJ)......................................................................................................... Hollandichthys sp. n. E

5b. All fins yellowish pigmented; scales in longitudinal series 36-43, rarely 44, x = 41.2;

presence of small bony spines in the dorsal surface of dorsal unbranched principal caudal-fin

rays, and in the ventral surface of the ventral unbranched principal caudal-fin ray in

specimens (c&s) larger than 50.0 mm SL; bony hooks absent in the dorsal and ventral

procurrent caudal-fin ray (rio Ribeira do Iguape basin, and island and coastal drainages from

Ilha do Cardoso to Itanhaém, SP)...........................................................................H. perstriatus

6a. Presence of a small black spot covering the base of the median caudal-fin rays (Fig. 26);

dorsal procurrent caudal-fin rays 8-11 and ventral procurrent caudal-fin rays 7-8 (tributaries

of the rio Maquiné and Três Forquilhas, RS, and of the rio Mampituba,

SC)........................................................................................................ Hollandichthys sp. n. A

6b. Absence of a small black spot covering the base of the median caudal-fin rays; dorsal

procurrent caudal-fin rays 11-14 (Fig. 18) and ventral procurrent caudal-fin rays 9-12 (Fig.

19)..............................................................................................................................................7

53
7a. Yellow pigmented fins in live specimens; dorsal procurrent caudal-fin rays 11-14 ( x =

12.7) (Fig. 18), and ventral procurrent caudal-fin rays 10-12 ( x = 10.6) (Fig. 19) (tributary of

rio Araçatuba in Ressacada, Garopaba, and smaller tributary of the rio Ratones and Córrego

Grande, Ilha de Santa Catarina, Florianópolis, SC).............................. Hollandichthys sp. n. B

7b. Red pigmented ventral- and adipose-fins in live specimens (Figs. 24b-c); dorsal

procurrent caudal-fin rays 10-13 ( x = 11.4) (Fig. 18) and ventral procurrent caudal-fin rays 8-

11 ( x = 9.4) (Fig. 19) (coastal rivers and islands between rio Jaraguá, Jaraguá do Sul, SC and

rio Nhundiaquara (baia de Paranaguá), Morretes, PR).................................................H. affinis

Hollandichthys multifasciatus (Eigenmann & Norris, 1900)

(Fig. 23)

Tetragonopterus multifasciatus Eigenmann & Norris, 1900:358 (type locality: Cubatão [São

Paulo, Brazil]); CAS 61483 [ex IU 9288], 3 syntypes.

Hollandichthys multifasciatus.- Eigenmann, 1909 [in part]:338 (new combination in table and

distribution: Ribeira) [based on the description of Pseudochalceus perstriatus

Miranda-Ribeiro, 1908 specimens from rio Ribeira belong to Hollandichthys

perstriatus].

Hollandichthys multifasciatus.- Eigenmann, 1921:225 (pl. 2, fig. 4; pl. 64, figs. 1, 2, 4; pl. 95,

fig. 8; redescription and habitat).

Material examined. Syntypes: CAS 61483, 2 of 3, one juvenile 31.4 mm SL, one female

66.8 mm SL, Cubatão, São Paulo, Brazil, H. von Ihering.

Non-type specimens: Brazil: São Paulo: coastal drainages: -MCP 30559, 22, 14.3-85.8

mm SL, tributary of rio Quilombo, Quilombo, Cubatão (23o51’S 46o20’W), 21 Sep 2002. -

MCP 30560, 35 (3 c&s), 12.6-75.2 mm SL, tributary of rio Quilombo, Quilombo, Cubatão

54
(23o50’S 46o19’W). -MZUSP 16459, 2, 58.8-60.3 mm SL, Piassaguera, Cubatão (23o50’S

46o23’W), 30 Apr 1953. -MNRJ 8545, 5 of 8, 56.5-89.3 mm SL, Santos (23o57’S 46o19’W),

1943. -MZUSP 37371, 10 of 15 (2 c&s), 31.2-60.2 mm SL, rio Cabuçu, Baixada Santista,

Santos, 28 Aug 1980. -FMNH 54375, 3 of 8 (1 c&s), 55.7-71.8 mm SL, raiz da Serra, rio

Moji, [Santos], 22 Jul 1908. Upper rio Tietê drainage: Santo André: MCP 20226, 3, 26.1-

78.4 mm SL, stream about 6 km NW of Paranapiacaba (23o46’S 46o20’W), 12 Jan 1997. -

MZUSP 28848, 13, 19.8-46.2 mm SL, tributary of rio Grande, Campo Grande (23o40’S

46o32’W), 26 May 1985. -MZUSP 35232, 1, 42.4 mm SL, stream between Campo Grande

and Paranapiacaba (23o40’S 46o32’W), Nov 1985. -MZUSP 35237, 20 of 38 (3 c&s), 32.8-

90.0 mm SL, stream near of the Paranapiacaba (23o46’S 46o20’W), Nov 1985. -MZUSP

35660, 15, 24.8-43.8 mm SL, Campo Grande (23o40’S 46o32’W), 3 Aug 1985. -MZUSP

36157, 1, 62.3 mm SL, rio Grande, Paranapiacaba (23o40’S 46o32’W), 15 Sep 1985.

Diagnosis. Hollandichthys multifasciatus is distinguished from all other species of the genus

by the adipose-fin black pigmented along usually half proximal length (Fig. 23). The presence

of one black humeral spot in adults (larger than 60.0 mm SL) differs H. multifasciatus from

Hollandichthys sp. n. A, Hollandichthys sp. n. B and H. affinis. The absence of black dots

laterally in the body near the anal fin base also distinguishes this species from Hollandichthys

sp. n. D.

Description. Description based in the type material and freshly collected specimens from rio

Quilombo drainage (Cubatão, SP). Morphometric data are summarized in Table 4. Body

compressed and elongate; greatest body depth usually anterior to dorsal-fin origin. Dorsal

head profile nearly straight. Dorsal profile convex from supraoccipital bone to base of last

dorsal-fin ray, and straight from this point to adipose-fin origin. Ventral profile of head

55
convex. Ventral body profile slightly convex to nearly straight from a vertical line projected

through pectoral -fin origin to a vertical line projected through pelvic-fin origin, and straight

to slightly concave from the last point to anal-fin origin. Body profile along anal-fin base

posterodorsally slanted. Caudal peduncle elongate, nearly straight to slightly concave along

dorsal and ventral margins.

Snout gently convex or nearly straight. Head small. Mouth slightly superior. Lower

jaw protruding anteriorly to upper jaw. Maxilla long and slightly curved aligned at angle of

approximately 45 degrees to longitudinal body axis, with anterodorsal portion convex and

posterodorsal portion concave, and all toothed portion smoothly concave. Maxilla anterior

portion narrow; posterior portion enlarged.

Premaxilla with two tooth rows; outer row with 2-3, tricuspid teeth with central cusp

longer; inner row teeth 5 with 3-5 cusps (fourth or fifth cusp diminutive); central cusp three or

four times longer and broader than other cusps; symphysean tooth much larger than the

remaining. Maxilla with 14-17 (usually 14-15), tricuspid teeth, except the last two, which are

uni- or bicuspidate. Central cusp longer than lateral cusp. Four or 5 anteriormost dentary teeth

larger, with 3-5 cusps (fourth or fifth cusp diminutive), followed by medium sized tooth with

3 cusps, and 8-11 teeth with 2-3 cusps or conical, gradually decreasing in length to last teeth;

central cusp in all teeth four or five times longer and broader than other cusps. All cusp tips

slightly curved posteriorly towards mouth gape.

Dorsal-fin rays ii, 9 (ii, 9; n =20); first unbranched ray approximately half-length of

second ray. Dorsal-fin origin located posterior to middle of SL and posterior to vertical

through pelvic-fin origin. Dorsal-fin profile rounded. Adipose-fin located approximately at

vertical through insertion of base of anal-fin rays 22-24.

Anal-fin rays iv-vi, 28-32 (v, 28-29; rarely 33, x = 30.2, n = 20). First unbranched ray

normally only apparent in cleared and stained specimens. Anal-fin profile nearly straight in

56
females. Anal-fin profile nearly straight along most of its border in males, except for a

concavity in the posterior portion of the fin, corresponding approximately to 24th-28th ray;

three or four last branched anal-fin rays longer than the remaining, forming a conspicuous

anal-fin lobe; penultimate ray longest. Anal-fin origin at vertical through base of four or fifth

dorsal-fin ray. Anal-fin rays of sexually mature males bearing one pair of small, elongate,

retrorse bony hooks along posterolateral border of each segment of lepidotrichia, usually

along 5-29 branched rays; hooks more numerous along 10th through 22nd branched rays, rarely

present on 30th ray. Hooks usually located along posteriormost branched and distal 1/2 to 2/3

of each ray. Unbranched and anterior 5-7 branched anal-fin rays in males and 3-4 branched

anal-fin rays in females bearing a large posterior flap along their posterolateral borders, more

developed in larger specimens of both sexes, and extending along entire length of anterior

unbranched rays and proximal midlength of last unbranched and anterior branched rays.

Pectoral-fin rays i, 12-15 (i, 12-13; x = 13.4, n = 20). Pectoral-fin tip reaching pelvic-

fin origin in males but not in females. Pelvic-fin rays 6, i (6, i; n = 20). Pelvic-fin tip extends

beyond anal-fin origin in males but not in females. Pelvic-fin origin anterior to vertical

through dorsal-fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook per

segment of lepidotrichia along ventromedial border of third to fifth branched rays.

Caudal-fin forked, with 19 principal rays (19; one specimen 18, and other 20, n = 20).

Dorsal procurrent rays 11-14 ( x = 12.8, n = 18). Ventral procurrent rays 9-12 ( x = 9.9, n =

18). Presence of the small bony spines in the dorsal surface of dorsal unbranched principal

caudal-fin ray, and in the ventral surface of the ventral unbranched principal caudal-fin ray in

larger specimens.

Scales cycloid, moderately large. Lateral line incomplete, perforated scales 7-10 ( x =

8.6, n = 20). Longitudinal series including lateral-line scales 40-45 (40; x = 42.5, n = 20).

Scale rows between dorsal-fin origin and lateral line 6-7 (6; usually 6, x = 6.1, n = 20); scale

57
rows between lateral line and pelvic-fin origin 5-6 (5; usually 5, x = 5.1, n = 20). Predorsal

scales 13-17 (15; usually 14-15), arranged in irregular series ( x = 14.5). Scales rows around

caudal peduncle 15-16 (15; x = 15.6). Scale sheath along anal-fin base 8-13 scales in single

series, extending to base of sixth to tenth branched rays. Body scales covering the belly with

posterior margin truncate or concave (Fig. 21).

Precaudal vertebrae 16-17; caudal vertebrae 20-21; total vertebrae 37-38 (n = 3).

Supraneurals 6-7 (n = 3). Gill-rakers 7-8/10-11 (n = 2).

Color in alcohol. See figure 23 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually light brown on

sides. Black uneven lines between longitudinal rows of scales conspicuous along midlateral

and midventral portions of the body. Small black chromatophores scattered over rays of all

fins. Adults with one black humeral spot located over 7-10 lateral line scales and extending

over first longitudinal series of scales above lateral line. Adipose-fin with black spot over

almost half fin, and usually extending from the anterior and dorsal borders to the center of the

fin. Caudal-fin base without black spot. Anal fin of adult males with one longitudinal black

band along its middle portion. Specimens larger than 60.0 mm SL with one humeral spot,

located over seventh to tenth perforated lateral line scales and extending over the horizontal

series of scales just above lateral line. Color pattern of juveniles (smaller than 47.0 mm SL)

similar to that of adults, but with two diffuse black humeral spots; anterior located over

second to fourth perforated lateral line scales and extending over 1-2 horizontal series of

scales above lateral line. Second humeral spot more dark and located over seventh to ninth

perforated lateral line scales and extending over 1-2 horizontal series of scales above lateral

line. Specimens smaller than 19.0 mm SL with second humeral spot dark and very

conspicuous.

58
Color in life. Color pattern similar to described for alcohol preserved specimens, except as

follows. Dorsal, caudal, anal, pelvic and pectoral fins yellow pigmented. Dorsal and ventral

borders of caudal fin, outer rays of pectoral and pelvic fins, distal portion of the first to third

branched rays of anal fin and upper portion of dorsal fin red-orange. Anal-fin with small black

chromatophores along its border forming a narrow stripe in some specimens. Adipose fin

brilliant yellow, with black spot in the anterior portion. Body yellowish (Figs. 23b-c).

Sexual dimorphism. Males of H. multifasciatus are easily recognized by the presence of

bony hooks on branched rays of the anal and pelvic fins, by the shape of the anal fin distal

border, and by the longer pectoral and pelvic fins (see description). Caudal peduncle depth is

slightly larger in males than females, and procurrent caudal-fin rays are more prominent in

males.

Habitat description. Based in lots MCP 30559 and MCP 30560. These specimens were

caught in small stream (tributaries of rio Quilombo) with lentic (or semi-lentic), shallow,

transparent water, ooze and leaf bottom, and moderate riparian vegetation. All specimens

were hidden under pendent plants along the margins of the stream.

Ecology. Stomach content of two specimens (MCP 30560) prepared for clearing and counter

staining consisted of insect parts and insect larvae. Other species collected along with H.

multifasciatus in tributaries of the rio Quilombo were Corydoras barbatus Quoy & Gaimard

(1824), Gymnotus pantherinus Steindachner (1908), Heptapterus mustelinus (Valenciennes,

1836), Phalloceros sp., Rhamdia sp. and Rivulus sp.

59
Distribution. This species has a small geographical distribution but encompassing two major

drainage systems of South America, the upper rio Tietê, a tributary of the upper rio Paraná, La

Plata drainage, and a few coastal Atlantic drainages of southern Brazil, near Cubatão and

Santos, São Paulo (Fig. 1). All these populations can be recognized by the peculiar color

pattern of the adipose fin and are not differentiable among themselves by any other meristic or

morphometric characters. Common patterns of distribution between the upper rio Tietê and

the coastal drainages have been found for the genus of Spintherobolus Eigenmann (1911) by

Weitzman & Malabarba (1999:39-40), and for Hyphessobrycon bifasciatus Ellis (1911), H.

reticulatus Ellis (1911), Pseudocorynopoma heterandria Eigenmann (1914) and G.

pantherinus by Langeani (1989:189). Langeani (1989), based on these faunal similarities, also

suggested that both the upper rio Tietê and the coastal drainages of São Paulo state were

connected in the past. M. C. Malabarba (1998) has consistently hypothesized past connections

between the rio Tietê and coastal drainages based on the phylogeny of fossil and recent

species of the clades Lignobrycon + Triportheus and Spintherobolus + Megacheirodon

Malabarba (1998) distributed in the area.

Remarks. Description of H. multifasciatus (Eigenmann & Norris, 1900) was based on three

syntypes from "Cubatão". We have examined two of the three syntypes (CAS 61483, 31.4 and

66.8 mm SL). The syntypes have lost most of their color patterns, but the largest specimen

retains the black pigments in the proximal portion of the adipose fin that allowed its

recognition as the same species of the population from Quilombo, Cubatão, São Paulo (MCP

30559, MCP 30560). Syntypes do not differ from these lots in any of the examined meristic

and morphometric characters. The specimen with 31.4 mm SL seems to be a young specimen,

and the largest specimen (66.8 mm SL) seems to be a female, since it lacks any of the

sexually dimorphic characters of males.

60
 Hollandichthys affinis (Steindachner, 1908)

(Fig. 24)

Pseudochalceus affinis Steindachner, 1908:29 (type locality: "Rio Jaraguá bei Joinville im

Staate S. Catharina (Brasilien)", [Rio Jaraguá is a tributary of rio Itapocu at Jaraguá do

Sul, Santa Catarina, Brazil]); type material: NMW 56736, 3, NMW 56737, 2.

Material examined. Lectotype (BY PRESENT DESIGNATION): NMW 56737:1 of 3,

male, 79.1 mm SL, rio Jaraguá, Joinvile, Santa Catarina, Brazil.

Paralectotypes. NMW 56736, 2 females, 78.0-83.2 mm SL, -NMW 56737, 1 male 82.5 mm

SL, and 1 female 70.0 mm SL, same data of the lectotype.

Other specimens: Brazil: Santa Catarina: Joinvile: DZSJRP 2245, 5, 19.2-48.1 mm SL,

rio Cubatão (26o12’S 48o54’W), 31 Mar 1989. -MCP 10659, 7, 34.0-87.0 mm SL, and

MZUSP 37727, 7, 37.7-55.9 mm SL, rio Seco, tributary of rio Cubatão, km 10 of the road BR

280 (26o10’S 48o57’W), 20 Sep 1985. -MCP 30667, 4, 50.0-67.0 mm SL, rio Lindo, tributary

of rio Cubatão, in the road SC 301, 2.1 km from BR 101 (26o11’S 48o55’W), 20 Oct 2002. -

MNRJ 13527, 24, 44.5-89.1 mm SL, Joinvile (26o18’S 48o51’W). -MZUSP 18504, 2, 23.7-

26.2 mm SL, rio Cubatão (26o12’S 48o54’W), 8 Sep 1974. Santa Catarina: Araquari: MCP

30552, 4, 55.4-68.2 mm SL, tributary of rio Parati (26o22’S 48o43’W), 19 Sep 2002. Santa

Catarina: São Francisco do Sul: MCP 30553, 4, 14.9-85.2 mm SL, tributary of rio Miranda

in Ilha de São Francisco do Sul (26o19’S 48o39’W), 19 Sep 2002. -MZUSP 35426, 9, 16.4-

80.9 mm SL, stream in Itapoá, (26o14’S 48o38’W0, 22 Dec 1985. Santa Catarina: Garuva:

MCP 14060, 4, 39.6-85.1 mm SL, -MZUSP 41792, 2, 44.0-53.9 mm SL, rio Garuva, BR 101

bridge (25o59’S 48o52’W), 7 May 1989. Paraná: Guaratuba: MCP 13859, 10 (2 c&s), 53.1-

80.9 mm SL, and MZUSP 41818, 7, 45.5-72.4 mm SL, rio São João at Pedra Branca do

Araraquara, BR 376, tributary of the baia de Guaratuba (25o56’S 48o55’W), 7 May 1989. -

61
MZUSP 18618, 6, 27.4-53.3 mm SL, creek stream, 5 km S of Guaratuba (25o55’S 48o36’W),

28 Dec 1975. -MZUSP 18602, 20, 19.2-33.2 mm SL, praia de Guaratuba about 5 km N of

Matinhos (25o46’S 48o35’W), 4 Dec 1975. Paraná: Paranaguá: MNRJ 6381, 12, 12.8-80.5

mm SL, rio Vila, km 9.5 of the "Estrada do Mar" (approx. 25o32’S 48o31’W), Mar 1944. -

MNRJ 8537, 1, 77.6 mm SL, Ilha do Mel (25o31’S 48o22’W), Sep 1952. -MHNCI 6144, 10

(2 c&s), 22.2-67.5 mm SL, rio Caçual, Ilha do Mel (25o31’S 48o22’W), 18 Jun 1991. Paraná:

Morretes: MCP 30556, 1, 57.5 mm SL, tributary of rio Nhundiaquara (25o24’S 48o52’W), 23

Sep 2002. -MZUSP 18610, 1, 18.9 mm SL, rio São João (25o24’S 48o51’W), 27 Dec 1975. -

MNRJ 8533, 1, 55.9 mm SL, córrego da Divisa, Porto de Cima (25o26’S 48o52’W), Jun 1953.

-MHNCI 6802, 2, 56.9-65.0 mm SL, tributary of rio Nhundiaquara (25o28’S 48o50’W), 2 Oct

1992. -MHNCI 6813, 4, 32.8-65.2 mm SL, rio do Pinto, tributary of rio Nhundiaquara

(25o36’S 48o53’W), 2 Oct 1992.

Diagnosis. Hollandichthys affinis differs from all other species of the genus by the red

pigmented ventral- and adipose-fins in live specimens (Figs. 24b-c); other species with red-

orange pigmented fins. Hollandichthys affinis differ from Hollandichthys sp. n. A by the

absence of a black spot on the caudal-fin base; and higher number of dorsal and ventral

procurrent caudal-fin rays (10-13, x = 11.4, versus 8-11, x = 9.4; Fig. 18), and (8-11, x = 9.4,

versus 7-8, x = 7.7; Fig. 19), respectively. Hollandichthys affinis differ from Hollandichthys

sp. n. B by the smaller number of dorsal and ventral procurrent caudal-fin rays (10-13, x =

11.4, versus 11-14, x = 12.7; Fig. 18), and (8-11, x = 9.4, versus 10-12, x = 10.6, Fig. 19),

respectively. Although largely overlap the counts, these differences are statistically significant

when submitted to Kruskal-Wallis One Way Analysis of Variance on Ranks for P ≤ 0.001.

Hollandichthys affinis differ from Hollandichthys sp. n. C, H. perstriatus, Hollandichthys sp.

n. D, and H. multifasciatus by the absence of a black humeral spot in adults (larger than 60.0

62
mm SL), and additionally from H. multifasciatus by the absence of the black pigmented

adipose-fin.

Description. Description based in the type material and freshly collected specimens from rio

Cubatão Norte and rio São João drainages. Morphometric data summarized in Table 5. Body

compressed and moderately elongate; greatest body depth usually anterior to dorsal-fin origin.

Dorsal head profile nearly straight. Dorsal profile from supraoccipital bony to base of last

dorsal-fin ray strongly convex, and straight to adipose-fin origin. Ventral profile from the

anterior tip of lower jaw to pectoral-fin origin slightly convex, and from this point to anal-fin

origin convex. Body profile along anal-fin base posterodorsally slanted. Caudal peduncle

elongate, nearly straight to slightly concave along dorsal and ventral margins.

Snout gently rounded. Head small. Mouth slightly superior. Lower jaw protruding

anteriorly to upper jaw. Maxilla long and slightly curved aligned at angle of approximately 45

degrees to longitudinal body axis, with anterodorsal portion convex and posterodorsal portion

concave, and all toothed portion smoothly concave. Maxilla anterior portion narrow; posterior

portion enlarged.

Premaxilla with two tooth rows; outer row with 2-4, tricuspid teeth with central cusp

longer. Inner row teeth 5, first longer than the remaining; tricuspid, with central cusp four or

five times longer and broader than lateral cusps. Maxilla with 10-17 (usually 12-14), tricuspid

teeth, except the last two, which are uni- or bicuspidate. Central cusp longer than lateral cusp.

Four or 5 anteriormost dentary teeth larger, with 3-4 cusps (fourth cusp diminute), followed

by medium sized tooth with 3 cusps, and 8-11 teeth with 2-3 cusps or conical, gradually

decreasing in length to last teeth; central cusp in all teeth four or five times longer and broader

than other cusps. All cusp tips slightly curved posteriorly towards inside the mouth gape.

63
 Dorsal-fin rays ii, 9 (ii, 10; 3 specimens ii, 10; n = 73); first unbranched ray

approximately one-half-length of second ray. Dorsal-fin origin located posterior to middle of

SL and posterior to vertical through pelvic-fin origin. Dorsal-fin profile rounded. Adipose fin

located approximately at vertical through insertion of 20-22 anal-fin rays base.

Anal-fin rays iii-v, 27-31 (iv, 28; rarely 26 or 32-33, x = 28.6, n = 72). First

unbranched ray normally only apparent in cleared and stained specimens. Anal-fin profile

nearly straight in females, and males show anal-fin profile nearly straight except for a

concavity corresponding approximately to 20th-25th ray. Three or fourth last branched anal-fin

rays longer than the remaining, being the penultimate ray the longest. Anal-fin origin at

vertical through base of fifth dorsal-fin ray. Anal-fin rays of sexually mature males bearing

one pair of small, elongate, retrorse bony hooks along posterolateral border of each segment

of lepidotrichia, usually along 4-25 branched rays; hooks more numerous along 7th through

17th branched rays, rarely present on 26th ray. Hooks usually located along posteriormost

branched and distal 1/2 to 2/3 of each ray.

Pectoral-fin rays i, 12-16 (i, 13; x = 13.8, n = 68). Pectoral-fin tip reaching pelvic-fin

origin in males but not in females. Pelvic-fin rays 6, i (6, i; rarely 5, i; n = 72). Pelvic-fin tip

reaching beyond anal-fin origin in males but not in females. Pelvic-fin origin located anterior

to vertical through dorsal-fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook

per segment of lepidotrichia along ventromedial border of third to fifth branched rays.

Caudal-fin forked, with 19 principal rays (19; n = 68). Dorsal procurrent rays 10-13

( x = 11.4, n = 32). Ventral procurrent rays 8-11 ( x = 9.4, n = 32). No spines were observed in

the surface of unbranched caudal-fin rays of the examined c&s specimens.

Scales cycloid, moderately large. Lateral line incomplete, perforated scales 7-10 (9;

x = 8.8, n = 62). Lateral series scales including lateral-line scales 37-42 (36; rarely 35-36 or

43-44, x = 39.9, n = 63). Scale rows between dorsal-fin origin and lateral line 6-7 (7; usually

64
6, x = 6.4, n = 63); scale rows between lateral line and pelvic-fin origin 4-7 (6; usually 5, x =

5.3, n = 61). Predorsal scales 12-15 (12; usually 14), arranged in irregular series ( x = 13.7).

Scales rows around caudal peduncle 14-16 (14; x = 14.5). Scale sheath along anal-fin base

bearing 8-15 scales in single series, extending to base of eighth to fourteenth branched rays.

Body scales covering the belly with posterior margin truncate or concave.

Precaudal vertebrae 17; caudal vertebrae 20-21; total vertebrae 37-38 (n = 2).

Supraneurals 6-7 (n = 2). Gill-rakers 7-8/11 (n = 2).

Color in alcohol. See figure 24 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually light brown on

sides. Black uneven lines between longitudinal rows of scales conspicuous along midlateral

and midventral portions of the body. Small black chromatophores scattered over rays of all

fins. Adults (larger than 60.0 mm SL) without humeral spot. Adipose-fin smoothly pigmented

in the border, and with a small black spot in the anterior portion (NMW 56736:2). Caudal-fin

base without black spot. Anal fin of adult males with one longitudinal black band along its

middle portion. Color pattern of juveniles (smaller than 49.0 mm SL) similar to that of adults,

but with one or two diffuse black humeral spots; anterior located over second to fourth

perforated lateral line scales and extending over 1-2 horizontal series of scales above lateral

line. Second humeral spot more dark and located over seventh to ninth perforated lateral line

scales and extending over 1-2 horizontal series of scales above lateral line. Specimens smaller

than 25.0 mm SL with second humeral spot dark and very conspicuous.

Color in life. Color pattern similar to described for alcohol preserved specimens, except as

follows. Dorsal, caudal, anal, pelvic and pectoral fins red-orange pigmented. Adipose fin and

65
outer rays of pelvic fin reddish (Figs. 24b-c). Anal-fin with small black chromatophores along

its border forming a narrow black contour in some specimens. Body yellowish.

Sexual dimorphism. Males of H. affinis are easily recognized by the presence of bony hooks

on the anal- and pelvic-fin branched rays, by the shape of the anal fin distal border, and by the

longer pectoral and pelvic fins (see description).

Habitat description. Based in lots MCP 30552, MCP 30553, MCP 30556 and MCP 30667.

These specimens were caught in small stream with lentic or with very low current, shallow,

transparent water or black water (MCP 30552), sand, ooze and leaf bottom, and moderate

riparian vegetation. In most of these localities the specimens were hidden under pendent

plants or near the margin of stream.

Ecology. Species collected with H. affinis in localities of the lots MCP 30552, MCP 30553,

MCP 30556 and MCP 30667 were: A. tajasica, Characidium sp., C. barbatus, Deuterodon

langei Travassos (1957), Geophagus brasiliensis (Quoy & Gaimard, 1824), H. bifasciatus, H.

reticulatus, Mimagoniates microlepis, Phalloceros sp. 1, Phalloceros sp. 2 and Schizolecis

guentheri (Miranda-Ribeiro, 1918).

Distribution. Hollandichthys affinis is known from coastal rivers and islands between rio

Jaraguá, Jaraguá do Sul, Santa Catarina and rio Nhundiaquara (baia de Paranaguá), Morretes,

Paraná, Brazil (Fig. 1).

Remarks. Original description of H. affinis was based on five syntypes from rio Jaraguá, near

Joinvile, SC. We have examined all these syntypes (NMW 56736:2 females 78.1-83.2 mm

66
SL, NMW 56737:1 female, 70.0 mm SL and 2 males, 79.1-82.9 mm SL). The syntypes are

relatively well preserved and its color pattern conserved. The specimens of lot NMW 56736

have black pigments in the proximal portion and in borders of the adipose fin, absent in the

specimens of the lot NMW 567737. They also differ in body depth, depth and length caudal

peduncle (see Table 5). Due to these differences, we prefer to designate one specimen of

NMW 56737 as lectotype for H. affinis (Fig. 24a).

Hollandichthys perstriatus (Miranda-Ribeiro, 1908)

(Fig. 25)

Pseudochalceus perstriatus Miranda-Ribeiro, 1908: (pages not numbered) (type locality:

"Corregos de Iporanga", [Iporanga, São Paulo, Brazil]), holotype probably lost.

Material examined. Neotype (BY PRESENT DESIGNATION): Brazil: São Paulo:

Iporanga: FMNH 54313, 1 of 3, male, 79.4 mm SL (approx. 24o35’S 48o36’W), 1 Dec 1908,

J. D. Haseman.

Other specimens: Brazil: São Paulo (rio Ribeira do Iguape basin): DZSJRP 2237, 1, 49.1

mm SL, rio Ribeira do Iguape, Registro (24o30’S 47o51’W), 5 Oct 1987. -DZSJRP 2293, 5,

21.6-81.3 mm SL, stream at Seara farm, Jacupiranga (24o42’S 48o1’W), 23 Aug 1985. -

DZSJRP 2309, 1, 76.6 mm SL, rio Jacupiranga, km 461 of road BR 116, Pariquera-Açu

(24o42’S 48o0’W), 6 May 1986. -DZSJRP 2318, 1, 60.6 mm SL, rio Sabadela at Intervales

farm, Sete Barras (24o23’S 47o56’W), 4-6 Mar 1994. -DZSJRP 3151, 3, 32.0-56.9 mm SL,

same date as DZSJRP 2293, 16 Aug 1985. -DZSJRP 3184, 3, 19.6-33.9 mm SL, Sítio

Guamiranga, Jacupiranga (24o42’S 48o1’W), 15 Aug 1985. -FMNH 54313, 2, 29.5-74.8 mm

SL, collected with neotype. -MNRJ 23489, 10 of 18, 43.0-85.6 mm SL, small stream at NW

of Juquiá, Santiago farm, Juquiá (24o18’S 47o40’W), 16 Feb 2002. -MZUSP 16381, 8 of 12,

67
39.2-65.9 mm SL, ribeirão Poço Grande tributary of rio Juquiá, Poço Grande farm, Juquiá

(24o20’S 47o38’W), Jan 1950. -MZUSP 17232, 10, 64.2-83.7 mm SL, Barra do Icapava

(24o41’S 47o26’W), 28 Jul 1966. -MZUSP 17802, 8, 13.0-85.1 mm SL, ribeirão da fazenda

Canta Galo, Juquiá (24o20’S 47o38’W), 5 Sep 1969. -MZUSP 36582, 24 (3 c&s), 25.4-53.8

mm SL, small margin lake, km 10 of road between Jacupiranga to Eldorado, Jacupiranga

(approx. 24o42’S 48o1’W), 14 Oct 1985. -MZUSP 40211, 2, 59.3-68.5 mm SL, rio Pariquera-

Mirim, Sítio Margom, Registro (24o30’S 47o50’W), 16 Aug 1985. -MZUSP 45197, 6, 14.2-

36.0 mm SL, tributary of rio Ribeira do Iguape, Eldorado (24o32’S 48o7’W), 5 Mar 1993. -

MZUSP 62884, 18 (1 c&s), 16.8-79.6 mm SL, stream in track of Cachoeira, Estação

Ecológica Juréia-Itatins (E.E.J.I.), Iguape (24o32’S 47o10’W), 24 Feb 1985. -UFRJ 4318, 1,

58.7 mm SL, stream from 20 km S at Jacupiranga, Jacupiranga (24o47’S 48o4’W), 30 Jul

1993. São Paulo (coastal drainages): MCP 30554, 8, 19.9-57.2 mm SL, stream Cachoeira

das Antas at Guaraú, Peruíbe (24o21’S 47o2’W), 20 Sep 2002. -MCP 30555, 9, 23.7-60.1 mm

SL, stream Cachoeira da Lagoa at Guaraú, Peruíbe (24o22’S 47o3’W), 20 Sep 2002. -MCP

30561, 2, 40.6-51.5 mm SL, same data as MCP 30554. -MZUSP 16455, 2, 36.5-58.2 mm SL,

Itanhaém (24o11’S 46o48’W), Mar 1953. -MZUSP 16546, 1, 73.6 mm SL, Pedrinhas, Ilha

Comprida (24o54’S 47o47’W), 26 Apr 1959. -MZUSP 51925, 4, 44.7-65.9 mm SL, river

crossing the road to Cachoeira, Ilha do Cardoso, Cananéia (approx. 25o8’S 47o56’W), 28 Oct

1989. -MZUSP 58728, 3, 45.6-75.0 mm SL, rio Itinguçu, tributary of rio Una do Prelado, E.

E. J. I., Peruíbe (24o23’S 47o7’W), Jun 1997. -MZUSP 58732, 3, 29.7-44.9 mm SL, rio

Tetequera, tributary of rio Guaraú, E.E.J.I., Peruíbe (24o24’S 47o4’W), Mar 1997. -UFRJ

4451, 1, 68.4 mm SL, stream near at Mandira, Cananéia (24o59’S 48o4’W), 30 Dec 1993. -

UFRJ 5440, 6, 22.8-61.0 mm SL, stream in track to Lage of beach, Ilha do Cardoso, Cananéia

(25o8’S 47o56’W), 26 Jul 2001. -USNM 297984, 12 (2 c&s), 25.4-76.6 mm SL, innominate

68
blackwater stream crossing road SP 193 at km 56, NW of Cananéia (24o56’S 47o58’W), 21

Feb 1988.

Diagnosis. The presence of one black humeral spot in specimens larger than 60.0 mm SL

(Fig. 25) distinguishes H. perstriatus from Hollandichthys sp. n. A, Hollandichthys sp. n. B

and H. affinis, without humeral spots in adults. The size of the humeral spot, smaller, covering

7-10 lateral line scales and extending over first longitudinal series of scales above lateral line,

versus humeral spot located over 7 to 13-14 lateral line scales and extending over first

longitudinal series of scales above and below lateral line, differ H. perstriatus from

Hollandichthys sp. n. C, respectively. Hollandichthys perstriatus differs from H.

multifasciatus by the absence of the adipose-fin black pigmented, and from Hollandichthys

sp. n. D by the absence of the black dots in the body portion near of the anal fin base.

Hollandichthys perstriatus differ from Hollandichthys sp. n. E by the smaller number scales

in longitudinal series (39-43, rarely 36-38 or 44, x = 41.2, n = 62, versus 40-46, x = 42.5, n =

42, respectively).

Description. Description based in the specimens collected in the rio Ribeira do Iguape basin.

Morphometric data are summarized in Table 6. Body compressed and elongate; greatest body

depth usually anterior to dorsal-fin origin. Dorsal head profile nearly straight or slightly

concave. Dorsal profile from supraoccipital bone to base of last dorsal-fin ray convex, and

straight from this point to adipose-fin origin. Ventral profile of head convex. Ventral body

profile slightly convex to nearly straight from pectoral-fin origin to pelvic-fin origin, and

straight to slightly concave from this point to anal-fin origin. Body profile along anal-fin base

posterodorsally slanted. Caudal peduncle elongate, nearly straight to slightly concave along

dorsal and ventral margins.

69
 Snout gently convex or nearly straight. Head small. Mouth slightly superior. Lower

jaw protruding anteriorly to upper jaw. Maxilla long and slightly curved aligned at angle of

approximately 45 degrees to longitudinal body axis, with anterodorsal portion convex and

posterodorsal portion concave, and all toothed portion smoothly concave. Maxilla anterior

portion narrow; posterior portion enlarged.

Premaxilla with two tooth rows; outer row with 2-3 tricuspid teeth, with central cusp

longer; inner row teeth 5 with 3-5 cusps (fourth or fifth cusp diminutive); central cusp three or

four times longer and broader than other cusps; medial tooth longer than the remaining.

Maxilla with 11-19 (usually 14-16), tricuspid teeth, except the last two, which are uni- or

bicuspidate. Central cusp longer than lateral cusp. Four or 5 anteriormost dentary teeth larger,

with 3-5 cusps (fourth or fifth cusp diminutive), followed by medium sized tooth with 3

cusps, and 8-14 teeth with 2-3 cusps or conical, gradually decreasing in length to last teeth;

central cusp in all teeth four or five times longer and broader than other cusps. All cusp tips

slightly curved posteriorly towards mouth gape.

Dorsal-fin rays ii, 9 (ii, 9; one specimen ii, 8, n = 69); first unbranched ray

approximately half-length of second ray. Dorsal-fin origin located posterior to middle of SL

and posterior to vertical through pelvic-fin origin. Dorsal-fin profile rounded. Adipose-fin

located approximately at vertical through insertion of 20th to 24th anal-fin rays bases.

Anal-fin rays iv-vi, 27-31 (iv, 30; rarely 26 or 32, x = 28.7, n = 69). First unbranched

ray normally only apparent in cleared and stained specimens. Anal-fin profile nearly straight

in females. Anal-fin profile nearly straight along most of its border in males, except for a

concavity in the posterior portion of the fin, corresponding approximately to 21th-26th ray;

three or four last branched anal-fin rays longer than the remaining, forming a conspicuous

anal-fin lobe; penultimate ray the longest. Anal-fin origin at vertical through base of four or

fifth dorsal-fin ray. Anal-fin rays of sexually mature males bearing one pair of small,

70
elongate, retrorse bony hooks along posterolateral border of each segment of lepidotrichia,

usually along 4-27 branched rays; hooks more numerous along 8th through 18th branched rays,

rarely present on 28th ray. Hooks usually located along posteriormost branched and distal 1/2

to 2/3 of each ray. Unbranched and anterior 5-6 branched anal-fin rays in males and 3-4

branched anal-fin rays in females bearing a large posterior flap along their posterolateral

borders, more developed in larger specimens of both sexes, and extending along entire length

of anterior unbranched rays and proximal midlength of last unbranched and anterior branched

rays.

Pectoral-fin rays i, 12-16 (i, 13; x = 13.6, n = 68). Pectoral-fin tip reaching pelvic-fin

origin in males but not in females. Pelvic-fin rays 6, i (6, i; one specimen 5, i, n = 69). Pelvic-

fin tip extends beyond anal-fin origin in males but not in females. Pelvic-fin origin anterior to

vertical through dorsal-fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook per

segment of lepidotrichia along ventromedial border of third to fifth branched rays.

Caudal-fin forked, with 19 principal rays (19; one specimen 18, and others 20-21, n =

67). Dorsal procurrent rays 11-15 ( x = 12.9, n = 46). Ventral procurrent rays 9-12 ( x = 10.3, n

= 46). Presence of the small bony spines in the dorsal surface of dorsal unbranched principal

caudal-fin rays, and in the ventral surface of the ventral unbranched principal caudal-fin ray in

larger specimens.

Scales cycloid, moderately large. Lateral line incomplete, perforated scales 7-11 (9;

x = 8.5, n = 64). Longitudinal series including lateral-line scales 39-43 (43; rarely 36-38 or

44, x = 41.2, n = 62). Scale rows between dorsal-fin origin and lateral line 6-7 (7; usually 6,

x = 6.4, n = 65); scale rows between lateral line and pelvic-fin origin 5-6 (6; x = 5.5, n = 65).

Predorsal scales 13-16 (16; usually 14-15), arranged in irregular series ( x = 14.6, n = 55).

Scales rows around caudal peduncle 14-16 (14; x = 15.5, n = 61). Scale sheath along anal-fin

71
base 8-17 scales in single series, extending to base of sixth to fifteen branched rays. Body

scales covering the belly with posterior margin truncate or concave.

Precaudal vertebrae 15-17; caudal vertebrae 20-21; total vertebrae 36-38 (n = 4).

Supraneurals 6-7 (n = 4). Gill-rakers 7-8/10-11 (n = 4).

Color in alcohol. See figures 25 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually light brown on

sides. Black uneven lines between longitudinal rows of scales conspicuous along midlateral

and midventral portions of the body. Small black chromatophores scattered over rays of all

fins. Adults with one black humeral spot located over 7-10 lateral line scales and extending

over first longitudinal series of scales above lateral line. Adipose-fin with small black spot

proximal portion, and usually extending in the border. Caudal-fin base without black spot.

Anal fin of adult males with one longitudinal black band along its middle portion. Specimens

larger than 60.0 mm SL with one humeral spot, located over seventh to tenth perforated lateral

line scales and extending over the horizontal series of scales just above lateral line. Color

pattern of juveniles (smaller than 50.0 mm SL) similar to that of adults, but with two diffuse

black humeral spots; anterior located over second to fourth perforated lateral line scales and

extending over 1-2 horizontal series of scales above lateral line. Second humeral spot more

dark and located over seventh to ninth perforated lateral line scales and extending over 1-2

horizontal series of scales above lateral line.

Color in life. Description of color based in MCP 30554, MCP 30555, MCP 30561, and one

specimen collected in the Estação Ecológica Juréia-Itatins by O. T. Oyakawa. Color pattern

similar to described for alcohol preserved specimens, except as follows. Dorsal, adipose,

72
caudal, anal, pelvic and pectoral fins yellowish pigmented. Anal-fin with small black

chromatophores along its border forming a narrow stripe in some specimens. Body yellowish.

Sexual dimorphism. Males of H. perstriatus are easily recognized by the presence of bony

hooks on the anal- and pelvic-fin branched rays, by the shape of the anal fin distal border, and

by the longer pectoral and pelvic fins (see description). Caudal peduncle depth is slightly

larger in males than females, and procurrent caudal-fin rays are more prominent in males.

Distribution. Hollandichthys perstriatus is widely distributed in rio Ribeira do Iguape basin,

and island and coastal drainages from Ilha do Cardoso to Itanhaém, São Paulo, Brazil (Fig. 1).

Common names. Brazil: São Paulo: “Lambari-listrado”(Miranda-Ribeiro, 1908); "Maria-

mole" (Região da Juréia); "Peixe-folha" (Vale do rio Ribeira do Iguape).

Remarks. Hollandichthys perstriatus (Miranda-Ribeiro, 1908) was described from "Corregos

de Iporanga" [Iporanga, SP]. Judging from the description it was probably based in only one

specimen measuring "9 cm", but no type specimens were listed or designated. Miranda-

Ribeiro (1953:390) lists Pseudochalceus perstriatus in the MNRJ catalog of the types of the

species and subspecies described by Alipio Miranda-Ribeiro, but gives no catalog number or

any additional information about the holotype or any additional type material for this species.

According to P. A. Buckup (pers. comm.) the location of the type-specimen(s) is unknown,

being probably lost or destroyed. Our effort to locate the type-material was unfruitful.

Although, Miranda-Ribeiro did not expressly designated the type specimen(s), we

found the Hollandichthys population from the rio Ribeira do Iguape basin, including

Iporanga, constitutes a valid species, and not a junior synonym of H. multifasciatus, as

73
anteriorly proposed. We consider a Neotype designation is needed to define the nominal taxon

objectively (ICZN, 2000: Article 75.1), and choose a specimen from the same locality of the

original description (Iporanga, SP).

Hollandichthys sp. n. A

(Fig. 26)

Holotype. MCP 30000, 85.8 mm SL, male, tributary of rio do Ouro, Barra do Ouro, Maquiné,

Rio Grande do Sul, Brazil (29o35’S 50o17’W), 25 Jan 2001, L. R. Malabarba, V. A. Bertaco,

J. Anza & T. Hasper.

Paratypes. Brazil: Rio Grande do Sul: rio Maquiné drainage: MCP 26969, 15, 32.6-85.8

mm SL, collected with the holotype. -MCP 24621, 13 (3 c&s), 38.6-87.8 mm SL, same

locality of the holotype, 31 Aug 1999, F. G. Becker, T. Finkler & M. Vassiliou. -MCP 25273,

18 (3 c&s), 40.4-85.8 mm SL, same locality of the holotype, 9 Dec 1999, F. G. Becker, V. A.

Bertaco & P. Colombo. -MCP 25416, 8, 36.8-69.5 mm SL, arroio Encantado, Maquiné

(29o36’S 50o12’W), 14 Nov 1999, F. G. Becker, P. Colombo & A. Brandt. -MCP 25417, 1,

63.4 mm SL, arroio Pinheiro, Maquiné (29o38’S 50o13’W), 15 Nov 1999, F. G. Becker, P.

Colombo & A. Brandt.

Additional material (non-types). Brazil: Rio Grande do Sul: rio Três Forquilhas

drainage: MCP 29244, 15 (2 c&s), 12.5-63.9 mm SL, creek tributary of arroio da Barra, Itati

(29o25’S 50o10’W). Santa Catarina: rio Mampituba drainage: MCP 23625, 3, 35.7-78.5

mm SL, -MCP 29241, 9, 29.4-45.6 mm SL, -MCP 29242, 12 (2 c&s), 22.8-58.1 mm SL,

arroio Molha Coco, Praia Grande (29o10’S 49o59’W). -MCP 29243, 4 (1 c&s), 37.1-71.9 mm

SL, rio Mampituba, Praia Grande (29o15’S 50o7’W).

74
Diagnosis. Hollandichthys sp. n. A differs from all other species of the genus by the presence

of a small black spot covering the base of the median caudal-fin rays (Fig. 26); and by the

longitudinal stripes faint pigmented and not clearly defined in the midventral portion of the

body. The body scales are usually more evenly pigmented than in the other Hollandichthys

species turning the longitudinal stripes more diffuse. All other species have the longitudinal

stripes strongly dark pigmented and clearly defined. Hollandichthys sp. n. A is also

recognized by the smallest number of dorsal procurrent caudal-fin rays (8-10, x = 9.4, versus

10-15, x ≥ 11.4 remaining species; Fig. 18) and by the smallest number ventral procurrent

caudal-fin rays (7-8, x = 7.7, versus 8-12 x ≥ 9.4 remaining species; Fig. 19) among

Hollandichthys species.

Description. Description based on specimens from rio Maquiné drainage. Morphometric data

are summarized in Table 7. Body compressed and elongate; greatest body depth usually

anterior to dorsal-fin origin. Dorsal head profile gently concave. Dorsal profile from

supraoccipital bone to base of last dorsal-fin ray slightly convex, and from this point to

adipose-fin origin straight. Ventral profile from the anterior tip of lower jaw to pelvic-fin

origin convex, and straight to anal-fin origin. Body profile along anal-fin base posterodorsally

slanted. Caudal peduncle elongate, nearly straight to slightly concave along dorsal and ventral

margins.

Snout gently convex or nearly straight. Head small. Mouth slightly superior. Lower

jaw protruding strongly anterior to upper jaw. Maxilla long and slightly curved aligned at

angle of approximately 45 degrees to longitudinal body axis, with anterodorsal portion

convex and posterodorsal portion concave, and all toothed portion smoothly concave. Maxilla

anterior portion narrow; posterior portion enlarged.

75
 Premaxilla with two tooth rows; outer row with 2-3, rarely 4, tricuspid teeth with

central cusp longer; inner row teeth 5 with 3-5 cusps (fourth or fifth cusp diminute); central

cusp three or four times longer and broader than other cusps; medial tooth longer than the

remaining. Maxilla with 13-17 (usually 13-14) tricuspid teeth, with central cusp longer. Four

or 5 anteriormost dentary teeth larger, with 3-4 cusps (fourth cusp diminutive), followed by

medium sized tooth with 3 cusps, and 7-12 teeth with 2-3 cusps or conical; central cusp in all

teeth three or four times longer and broader than other cusps. All cusp tips slightly curved

posteriorly towards mouth gape.

Dorsal-fin rays ii, 9 (ii, 9; n = 32); first unbranched ray approximately one-half length

of second ray. Dorsal-fin origin located posterior to middle of SL and posterior to vertical

through pelvic-fin origin. Dorsal-fin profile rounded. Adipose-fin located approximately at

vertical through insertion of 20th to 22th anal-fin rays. Unbranched and anterior 3 to 4 dorsal-

fin rays bearing a large posterior flap along their posterolateral borders, more developed in

larger specimens of both sexes, and extending along entire length of first unbranched ray and

proximal midlength of second unbranched and anterior branched rays.

Anal-fin rays iii-v, 27-30 (iii, 28; rarely 26, x = 28.6, n = 32). First unbranched ray

normally only apparent in cleared and stained specimens. Anal-fin profile nearly straight in

females. Anal-fin profile nearly straight along most of its border in males, except for a

concavity in the posterior portion of the fin, corresponding approximately to 20th to 25th ray;

three or four last branched anal-fin rays longer than the remaining, forming a conspicuous

anal-fin lobe; penultimate ray the longest. Anal-fin origin at vertical through the base of third

to fourth dorsal-fin branched rays. Anal-fin rays of sexually mature males bearing one pair of

small, elongate, retrorse bony hooks along posterolateral border of each segment of

lepidotrichia, along 3 to 27 (usually 4-23) branched rays; hooks more numerous along 7th

through 20th branched rays, rarely present on 27th ray. Hooks usually located along

76
posteriormost branch and distal 1/2 to 2/3 length of each ray. Unbranched and anterior 2 to 3

anal-fin rays bearing a large posterior flap along their posterolateral borders, more developed

in larger specimens of both sexes, and extending along entire length of anterior unbranched

rays and proximal midlength of last unbranched and anterior branched rays.

Pectoral-fin rays i, 11-14 (i, 14; x = 13.2, n = 32). Pectoral-fin tip reaching pelvic-fin

origin in males but not in females. Pelvic-fin rays 6, i (6, i; rarely 5, i, n = 32). Pelvic-fin tip

extends beyond anal-fin origin in males but not in females. Pelvic-fin origin anterior to

vertical through dorsal-fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook per

segment of lepidotrichia along ventromedial border of third to fifth branched rays.

Caudal-fin forked, with 19 principal rays (19; one specimen 18, n = 31). Caudal-fin

lobes profile rounded. Dorsal procurrent caudal-fin rays 8-11 ( x = 9.4, n = 20). Ventral

procurrent caudal-fin rays 7-8 ( x = 7.7, n = 20). Presence of the small bony spines in the

dorsal surface of dorsal unbranched caudal-fin ray, and in the ventral surface of the ventral

unbranched principal caudal-fin ray in larger specimens.

Scales cycloid, moderately large. Lateral line incomplete, perforated scales 6-8 (6; x =

7, n = 31). Longitudinal series of scales including lateral-line scales 37-40 (39; rarely 35 or

41-42, x = 38.5, n = 32). Scale rows between dorsal-fin origin and lateral line 6-7 (7; usually

7, x = 7, n = 32); scale rows between lateral line and pelvic-fin origin 5-6 (6; usually 6, x = 6,

n = 32). Predorsal scales 12-16 (14; usually 14), arranged in irregular series ( x = 13.7). Scale

rows around caudal peduncle 14-16 (14; x = 14.6). Scale sheath along anal-fin base 8-15

scales in single series, extending to base of sixth to twenty branched rays. Body scales

covering the belly with posterior margin truncate or concave.

Precaudal vertebrae 16-18; caudal vertebrae 20-21; total vertebrae 36-39, usually 37 (n

= 6). Supraneurals 6-7, usually 6 (n = 6). Gill-rakers 7/10-11 (n = 6).

77
Color in alcohol. See figures 26 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually light brown on

sides. Black longitudinal lines usually forming a zigzag pattern between longitudinal rows of

scales; more conspicuous in middle portion of body, and more visible in specimens larger

50.0 mm SL. Small black chromatophores scattered over rays of all fins. Adipose-fin

smoothly pigmented in the border. Small black spot in the base of the median caudal-fin rays.

Specimens larger than 60.0 mm SL without humeral spot. Specimens measuring 42.0-50.0

mm SL with one or two faint humeral spots. Color pattern of juveniles (smaller than 41.0 mm

SL) similar to that of adults, but with two diffuse black humeral spots; anterior located over

first to fourth perforated lateral line scales and sometimes extending over the horizontal series

of scales just above lateral line. Second humeral spot more dark located over seventh to eighth

perforated lateral line scales and sometimes extending over the horizontal series of scales just

above lateral line.

Color in life. Color pattern similar to described for alcohol preserved specimens, except as

follows. Dorsal, anal and caudal fins yellow pigmented. Pelvic fin yellowish with tip of outer

rays red-orange. Anal-fin with small black chromatophores along its border forming a narrow

stripe in some specimens. Distal portion of the branched first to third rays of anal fin red-

orange. Adipose fin brilliant yellow, with small chromatophores along its border. Body

yellowish (Fig. 26c).

Sexual dimorphism. Males of Hollandichthys sp. n. A are easily recognized by the presence

of bony hooks on the anal- and pelvic-fin branched rays, by the shape of the anal fin distal

border, and by the longer pectoral and pelvic fins (see description). Caudal peduncle depth is

78
slightly larger in males than females, and procurrent caudal-fin rays are more prominent in

males.

Habitat description. All specimens were caught in lateral puddles of rivers and very small

tributaries with lentic waters, shallow, black water, ooze and leaf bottom, and very dense

riparian vegetation, similar to observed by Sabino & Castro (1990) for H. multifasciatus

(=Hollandichthys sp. n. E, Ubatuba, SP) in an Atlantic Forest stream in Southeastern Brazil.

The larger specimens (70.0-80.0 mm SL) were found isolated and small specimens (30.0-40.0

mm SL) were captured in groups of 3 to 6 fishes, also observed by Sabino & Castro (1990).

Although extensive collection efforts have been applied in sampling fishes the last 20 years in

the rio Maquiné, rio Três Forquilhas and rio Mampituba drainages, Hollandichthys sp. n. A

was never caught in the main river itself, with lotic water, stones, clean water, and few

riparian vegetation.

Ecology. Stomach contents of three specimens (MCP 24621) prepared for clearing and

staining consisted of spider, ant, beetle, and insect parts. Other species collected along with

Hollandichthys sp. n. A were Astyanax sp., Cyphocharax voga (Hensel, 1870), Gymnotus sp.,

H. luetkenii (Boulenger, 1887), Characidae gen. n., M. rheocharis Menezes & Weitzman

(1990), Phalloceros sp. and Steindachnerina biornata (Braga & Azpelicueta, 1987).

Conservation note. Hollandichthys sp. n. A was listed as a threatened species in the state of

Rio Grande do Sul, Brazil (Marques et al., 2002), but named as H. multifasciatus.

Distribution and geographical variation. This species in know from smaller tributaries of

the rio Maquiné and rio Três Forquilhas, Rio Grande do Sul and of the rio Mampituba

79
drainage, Santa Catarina, Brazil, and represents the meridional limit of distribution of the

genus (Fig. 1). Comparing the type material (rio Maquiné) and non type specimens (rio Três

Forquilhas and rio Mampituba) we found some small differences in morphometric characters:

caudal peduncle length, snout length, orbital diameter, and interorbital width. These

differences may be correlated with the low number of individuals of the samples, and smaller

size of the specimens from rio Três Forquilhas and rio Mampituba (see Table 7). These small

differences do permit the recognition of more than one species within what is herein identified

as Hollandichthys sp. n. A.

Hollandichthys sp. n. B

(Fig. 27)

Holotype. Brazil: Santa Catarina: Ilha de Santa Catarina: Florianópolis: MCP 30001,

88.1 mm SL, male, stream Córrego Grande near the Área de Preservação Ambiental (APA) da

CASAN, Lagoa da Conceição (27o35’S 48o28’W), 25 Oct 2001, L. R. Malabarba, V. A.

Bertaco & M. A. Azevedo.

Paratypes. Brazil: Santa Catarina: Ilha de Santa Catarina: Florianópolis: MCP 28737,

35 (3 c&s), 34.4-88.1 mm SL, collected with the holotype. -MCP 28732, 9, 15.2-69.8 mm SL,

-MCP 28747, 6, 22.7-60.7 mm SL, tributary of rio Ratones (27o30’S 48o29’W), same date and

collectors of holotype. -UFRJ 1957, 7, 20.2-53.3 mm SL, same locality of the holotype, 22

Mar 1993, R. Sachsse.

Additional material (non-types). Brazil: Santa Catarina: Garopaba: MCP 28734, 4, 54.3-

74.0 mm SL, -MCP 29080, 20 (2 c&s), 17.2-77.8 mm SL, tributary of rio Araçatuba at

Ressacada (28o4’S 48o42’W).

80
Diagnosis. Hollandichthys sp. n. B is distinguished from Hollandichthys sp. n. A by the

absence of a black spot on the caudal-fin base, and by the number of dorsal procurrent caudal-

fin rays (11-14, x = 12.7, versus 8-11, x = 9.4; Fig. 18) and ventral procurrent caudal-fin rays

(10-12, x = 10.6, versus 7-8, x = 7.7; Fig. 19, respectively). Hollandichthys sp. n. B differ

from H. affinis by color in life (fins yellow pigmented (Fig. 27) versus fins red-orange

pigmented (Fig. 24b-c), respectively), and larger number of dorsal and ventral procurrent

caudal-fin rays (11-14, x = 12.7, versus 10-13, x = 11.4; Fig. 18), and (10-12, x = 10.6,

versus 8-11, x = 9.4; Fig. 19), respectively. Smaller orbital diameter differ Hollandichthys sp.

n. B from Hollandichthys sp. n. C (23.9-31.1% HL, x = 28.3% versus 27.7-34% HL, x = 30.3,

respectively; Table 8). The absence of one black humeral spot in specimens larger than 60.0

mm SL differs Hollandichthys sp. n. B from Hollandichthys sp. n. C, H. perstriatus,

Hollandichthys sp. n. D, and H. multifasciatus. It is also distinguished from H. multifasciatus

by the lack of a black spot on adipose-fin.

Description. Description based with the specimens from streams of Ilha Santa Catarina,

Florianópolis, SC. Morphometric data for Hollandichthys sp. n. B are summarized in Table 8.

Body compressed and moderately elongate; greatest body depth usually anterior to dorsal-fin

origin. Dorsal head profile nearly straight. Dorsal profile from supraoccipital bone to dorsal-

fin origin slightly convex, and from this point to base of last dorsal-fin ray strongly convex,

and straight to adipose-fin origin. Ventral profile from the anterior tip of lower jaw to

pectoral-fin origin convex, and from this point to anal-fin origin slightly convex. Body profile

along anal-fin base posterodorsally slanted. Caudal peduncle elongate, nearly straight to

slightly concave along dorsal and ventral margins.

Snout gently rounded. Head small. Mouth slightly superior. Lower jaw protruding

strongly anterior to upper jaw. Maxilla long and slightly curved aligned at angle of

81
approximately 45 degrees to longitudinal body axis, with anterodorsal portion convex and

posterodorsal portion concave, and all toothed portion smoothly concave. Anterior portion of

maxilla narrow, and posterior portion enlarged.

Premaxilla with two tooth rows; outer row with 2-3, tricuspid teeth with central cusp

longer; inner row teeth 5, first longer than the remaining; tricuspidate, with central cusp four

or five times longer and broader than lateral cusps. Maxilla with 11-19 (usually 15-17),

tricuspidate teeth, except the last two, which are uni- or bicuspidate. Central cusp longer than

lateral cusp. Four or 5 anteriormost dentary teeth larger, with 3-4 cusps (cusp fourth

diminute), followed by medium sized tooth with 3 cusps, and 10-14 teeth with 2-3 cusps or

conical, gradually decreasing in length to teeth last; central cusp in all teeth four or five times

longer and broader than other cusps. All cusp tips slightly curved posteriorly towards inside

mouth gape.

Dorsal-fin rays ii, 9 (ii, 9; n = 38); first unbranched ray approximately one-half length

of second ray. Dorsal-fin origin located posterior to middle of SL and posterior to vertical

through pelvic-fin origin. Dorsal-fin profile rounded. Adipose-fin located approximately at

vertical through insertion of 22-23 anal-fin ray bases.

Anal-fin rays iv-vi, 27-30 (iv, 28; rarely 26, x = 28.2, n = 39). First unbranched ray

normally only apparent in cleared and stained specimens. Anal-fin profile nearly straight in

females; males show anal-fin profile nearly straight except for a concavity corresponding

approximately to 20th-25th ray. Three or fourth last branched anal-fin rays of males longer

than the remaining, being the penultimate ray the longest one. Anal-fin origin at vertical

through base of fifth dorsal-fin ray. Anal-fin rays of sexually mature males bearing one pair of

small, elongate, retrorse bony hooks along posterolateral border of each segment of

lepidotrichia, usually along 4-23 branched rays; hooks more numerous along 7th through 20th

82
branched rays, rarely present on 26th ray. Hooks usually located along posteriormost branched

and distal 1/2 to 2/3 length of each ray.

Pectoral-fin rays i, 14-17 (i, 14; x = 14.4, n = 38). Pectoral-fin tip reaching pelvic-fin

origin in males but not in females. Pelvic-fin rays 6, i (n = 38). Pelvic-fin tip outstrips anal-fin

origin in males but not in females. Pelvic-fin origin located anterior to vertical through dorsal-

fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook per segment of

lepidotrichia along ventromedial border of third to fifth branched rays.

Caudal-fin forked, with 19 principal rays (19; one specimen 18). Dorsal procurrent

rays 11-14 ( x = 12.7, n = 20). Ventral procurrent rays 10-12 ( x = 10.6, n = 20). Presence of

small bony spines in the dorsal surface of dorsal unbranched principal caudal-fin ray, and in

the ventral surface of the ventral unbranched principal caudal-fin rays in adults (larger than

50.0 mm SL).

Scales cycloid, moderately large. Lateral line incomplete, perforated scales 6-11 (8;

x = 8.5, n = 38). Longitudinal series of scales including lateral-line scales 37-40 (38; rarely 36

or 41, x = 38.3, n = 38). Scale rows between dorsal-fin origin and lateral line 6-7 (6; usually

6, x = 6.1, n = 38); scale rows between lateral line and pelvic-fin origin 5-6 (5; usually 5, x =

5.1, n = 38). Predorsal scales 13-15 (13; usually 14), arranged in irregular series ( x = 14).

Scales rows around caudal peduncle 14-16 (14; x = 14.2). Scale sheath along anal-fin base 8-

14 scales in single series, extending to base of eighth to fourteenth branched rays. Body scales

covering the belly with posterior margin truncate or concave.

Precaudal vertebrae 16-17; caudal vertebrae 20-21; total vertebrae 37-38, usually 37 (n

= 3). Supraneurals 5-7 (n = 6). Gill-rakers 7/10-11 (n = 3).

Color in alcohol. See figure 27 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually yellow on sides.

83
Black zigzag longitudinal lines between longitudinal body rows of scales, more distinct in

median region of body. Adults without humeral spot. Small black chromatophores scattered

over rays of all fins. Adipose-fin smoothly pigmented in the border. Caudal-fin base without

black spot. Color pattern of juveniles (smaller than 41.0 mm SL) similar to that of adults

(larger than 51.0 mm SL), but with two black diffuses humeral spots; anterior located over

first to fourth perforated lateral line scales and extending over 1-2 horizontal series of scales,

including lateral line. Second humeral spot located over seventh to ninth perforated lateral

line scales and extending over 1-2 horizontal series of scales, including lateral line. Specimens

sub-adults (42.0-50.0 mm SL) with one or two faint humeral spots.

Color in life. Color pattern similar to described for alcohol preserved specimens, except as

follows. Dorsal, anal and caudal fins yellow pigmented. Pelvic fin yellowish with tip of outer

rays red-orange. Anal-fin with small black chromatophores along its border forming a narrow

stripe in some specimens. Distal portion of the branched first to third rays of anal fin red-

orange. Adipose fin brilliant yellow, with small chromatophores along its border. Body

yellowish (Fig. 27).

Sexual dimorphism. Males of Hollandichthys sp. n. B are easily recognized by the presence

of bony hooks on the anal- and pelvic-fin branched rays, by the shape of the anal fin distal

border, and by the longer pectoral and pelvic fins (see description). Caudal peduncle depth is

slightly larger in males than females, and procurrent caudal-fin rays are more prominent in

males.

Habitat description. The population sample from Córrego Grande, near the Área de Proteção

Ambiental, CASAN, Lagoa da Conceição, was collected in a small stream running in gallery

84
forest. The specimens were collected in a stream about 1.5 m, wide water depth 60 cm,

transparent and lentic water, little emergent vegetation, and bottom with sand and ooze. The

sampled place was deforested and the marginal vegetation replaced by grass, but a few meters

above the stream was covered with dense vegetation. The specimens from a tributary of rio

Araçatuba at Garopaba, SC, were caught in small stream with clear water, faint current water,

about 1.5 m wide and 30-90 cm deep, sand, ooze and leaf bottom, and very riparian

vegetation. All specimens were hidden under pendent plants in margin of stream.

Ecology. Stomach contents of two specimens (MCP 28737) prepared for clearing and staining

consisted of a spider, ants, insect and plants parts. Species collected with Hollandichthys sp.

n. B in streams of Ilha de Santa Catarina, SC: Characidium sp., C. barbatus, G. brasiliensis,

Phalloceros sp. and Rhamdia sp.

Distribution. This species in know from a tributary of rio Araçatuba at Ressacada, Garopaba,

and smaller tributaries of the rio Ratones and Córrego Grande, Ilha de Santa Catarina,

Florianópolis, Santa Catarina, Brazil (Fig. 1).

Hollandichthys sp. n. C

(Fig. 28)

Holotype. MCP 32038, 79.3 mm SL, tributary of rio Tagaçaba in the road PR 404 (25o12’S

48o26’W), 23 Sep 2002, L. R. Malabarba, V. A. Bertaco & M. A. Azevedo.

Paratypes. Brazil: Paraná: Guaraqueçaba: MCP 30557, 9, 44.6-62.4 mm SL, tributary of

rio Serra Negra in the road PR 404 (25o10’S 48o25’W), 23 Sep 2002, L. R. Malabarba, V. A.

Bertaco & M. A. Azevedo. -MCP 30558, 5, 24.3-71.5 mm SL, tributary of rio Tagaçaba in the

road PR 404 (25o12’S 48o26’W), same date and collectors as MCP 30557. -MHNCI 6396, 4,

85
28.9-72.0 mm SL, rio Braço do Macaco at Laranja Azeda (25o6’S 48o19’W), 6 Feb 1992, A.

M. Cordeiro, A. P. Barreto & E. Grando. -MHNCI 7823, 13 (2 c&s), 36.2-80.8 mm SL, rio

Engenho tributary of the rio Morato, Figueira farm, (25o13’S 48o19’W), 18 Jun 1991, W. B.

Wosiacki & J. C. Mateus.

Diagnosis. Hollandichthys sp. n. C is distinguished from all other species of the genus by

presence of one larger black humeral spot in specimens larger than 60.0 mm SL, located over

7 to 13-14 lateral line scales and extending over first longitudinal series of scales above and

below lateral line (Fig. 28), versus humeral spot located over 7-10 lateral line scales and

extending over first longitudinal series of scales above lateral line in H. perstriatus, H.

multifasciatus, Hollandichthys sp. n. D, and Hollandichthys sp. n. E, and humeral spot absent

in Hollandichthys sp. n. A, Hollandichthys sp. n. B, and H. affinis. It is further distinguished

from Hollandichthys sp. n. B by the larger orbital diameter (23.9-31.1% HL, x = 28.3%,

versus 27.7-34.0% HL, x = 30.3%, respectively; Table 8). It is also distinguished from H.

multifasciatus by the lack of a black spot on adipose-fin.

Description. Morphometric data for Hollandichthys sp. n. C are summarized in Table 8.

Body compressed and elongate; greatest body depth usually anterior to dorsal-fin origin.

Dorsal head profile nearly straight. Dorsal profile from supraoccipital bone to base of last

dorsal-fin ray slightly convex, and from this point to adipose-fin origin straight. Ventral

profile from the anterior tip of lower jaw to pelvic-fin origin convex, and straight to anal-fin

origin. Body profile along anal-fin base of sexually mature males slight convex, and nearly

straight in females. Caudal peduncle elongate, nearly straight to slightly concave along dorsal

and ventral margins.

86
 Snout gently rounded. Head small. Mouth slightly upper terminal. Lower jaw

protruding strongly anterior to upper jaw. Maxilla long and slightly curved aligned at angle of

approximately 45 degrees to longitudinal body axis, with anterodorsal portion convex and

posterodorsal portion concave, and all toothed portion smoothly concave. Maxilla anterior

portion narrow; posterior portion enlarged.

Premaxilla with two tooth rows; outer row with 2-3, rarely 4, tricuspid teeth with

central cusp longer; inner row teeth 5, first longer than the remaining; with 3-5 cusps (fourth

or fifth cusp diminute), with central cusp three or four times longer and broader than other

cusps. Maxilla with 13-17 (usually 13-14), tricuspid teeth, with central cusp longer. Four or 5

anteriormost dentary teeth larger, with 3-4 cusps (fourth cusp diminutive), followed by

medium sized tooth with 3 cusps, and 7-12 teeth with 2-3 cusps or conical; central cusp in all

teeth three or four times longer and broader than other cusps. All cusp tips slightly curved

posteriorly towards inside of mouth.

Snout gently rounded. Head small. Mouth slightly upper terminal. Lower jaw

protruding strongly anterior to upper jaw. Maxilla long and slightly curved aligned at angle of

approximately 45 degrees to longitudinal body axis, with anteroventral border slightly

concave and posterodorsal border concave. Maxilla anterior portion narrow and wide in the

posterior portion.

Premaxilla with two tooth rows; outer row with 3, tricuspid teeth with central cusp

longer; inner row teeth 5, first longer than the remaining; with 3 cusps, with central cusp three

or four times longer and broader than other cusps. Maxilla with 11-16 (usually 14-15),

tricuspid teeth, with central cusp longer. Four or 5 anteriormost dentary teeth larger, with 3-4

cusps (fourth cusp diminutive), followed by medium sized tooth with 3 cusps, and 8-12 teeth

with 2-3 cusps or conical; central cusp in all teeth three or four times longer and broader than

other cusps. All cusp tips slightly curved posteriorly towards inside of mouth.

87
 Dorsal-fin rays ii, 9 (ii, 9; 2 specimens ii, 8; n = 31); first unbranched ray

approximately one-half length of second ray. Dorsal-fin origin located posterior to middle of

SL and posterior to vertical through pelvic-fin origin. Dorsal-fin profile rounded. Adipose-fin

located approximately at vertical through insertion of 22-23 anal-fin rays base.

Anal-fin rays iv-vi, 28-30 (iv, 29; one specimen 25, rarely 31, x = 28.7, n = 31). First

unbranched ray normally only apparent in cleared and stained specimens. Anal-fin profile

nearly straight in females, and males show anal-fin profile nearly straight except for a

concavity corresponding approximately to 22th-25th ray. Three or four last branched anal-fin

rays longer than the remaining, being the penultimate ray the longest one. Anal-fin origin at

vertical through bases of third to fourth dorsal-fin branched ray. Anal-fin rays of sexually

mature males bearing one pair of small, elongate, retrorse bony hooks along posterolateral

border of each segment of lepidotrichia, usually along 5-23 branched rays; hooks more

numerous along 8th through 17th branched rays, rarely present on 27th ray. Hooks usually

located along posteriormost branched and distal 1/2 to 2/3 length of each ray.

Pectoral-fin rays i, 12-15 (i, 13; x = 14.2, n = 31). Pectoral-fin tip reaching pelvic-fin

origin in males but not in females. Pelvic-fin rays 6, i (6, i; one specimen 5, i, n = 31). Pelvic-

fin tip outstrips anal-fin origin in males but not in females. Pelvic-fin origin located anterior to

vertical through dorsal-fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook per

segment of lepidotrichia along ventromedial border of third to fifth branched rays.

Caudal-fin forked, with 19 principal rays (19; n = 31). Caudal-fin lobes profile

rounded. Dorsal procurrent rays 11-14 ( x = 12.4, n = 23). Ventral procurrent rays 9-12 ( x =

10.0, n = 23). Presence of small bony spines in the dorsal surface of unbranched principal

caudal-fin ray, and in the ventral surface of the ventral unbranched principal caudal-fin ray in

adults (larger than 50.0 mm SL).

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 Scales cycloid, moderately large. Lateral line incomplete, perforated scales 8-10 (9;

one specimen 6 and 12, x = 8.9, n = 30). Lateral series scales including lateral-line scales 39-

43 (41; rarely 38 or 44-46, x = 40.4, n = 31). Scale rows between dorsal-fin origin and lateral

line 6-7 (6; usually 6, x = 6.2, n = 31); scale rows between lateral line and pelvic-fin origin 5-

6 (5; usually 5, x = 5.2, n = 31). Predorsal scales 13-17 (14; usually 14-15), arranged in

irregular series ( x = 14.4). Scale rows around caudal peduncle 14-16 (14; x = 14.7). Scale

sheath along anal-fin base with 8-14 scales in single series, extending to base of seventh to

twenty branched rays. Body scales covering the belly with posterior margin truncate or

concave.

Precaudal vertebrae 17; caudal vertebrae 20; total vertebrae 37 (n = 2). Supraneurals 7,

(n = 2). Gill-rakers 7-8/11 (n = 2).

Color in alcohol. See figures 28 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually yellow on sides.

Black zigzag longitudinal lines between longitudinal body rows of scales, more distinct in

median region of body. Adult males and females with one large humeral spot located over 7

to 13-14 lateral line scales and extending over first longitudinal series of scales above and

below lateral line. Small black chromatophores scattered over rays of all fins. Adipose-fin

smoothly pigmented in the border. Caudal-fin base without black spot. Color pattern of

juveniles (36.2 mm SL) similar to that of adults (larger than 51.0 mm SL), but with two black

diffuses humeral spots; anterior located over first to fourth perforated lateral line scales and

extending over 1-2 horizontal series of scales, including lateral line. Second humeral spot

located over seventh to ninth perforated lateral line scales and extending over 1-2 horizontal

series of scales, including lateral line. Specimens sub-adults (47.7-52.5 mm SL) with one

small humeral spot (equivalent the second humeral spot in juveniles).

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Color in life. Color pattern similar to described for alcohol preserved specimens, except as

follows. Dorsal, pectoral, pelvic, adipose, anal and caudal fins yellow pigmented. Anal-fin

with small black chromatophores along its border forming a narrow stripe in some specimens.

Adipose fin brilliant yellow, with small chromatophores along its border. Body yellowish.

Sexual dimorphism. Males of Hollandichthys sp. n. C are easily recognized by the presence

of bony hooks on the anal- and pelvic-fin branched rays, by the shape of the anal fin distal

border, and by the longer pectoral and pelvic fins (see description). Caudal peduncle depth is

slightly larger in males than females, and procurrent caudal-fin rays are more prominent in

males.

Habitat description. Based in lots MCP 30557 and MCP 30558. These specimens were

caught in small stream with lentic or with very low current, 30-50 cm depth, transparent

water, sand and some stones bottom, and moderate riparian vegetation. In most localities the

specimens were hidden under pendent plants or tree-trunks in water near margin of stream.

Ecology. The male specimen (62.0 mm SL; MHNCI 7823), possessed in the stomach two

specimens of the M. lateralis (Nichols, 1913) (23.6-27.0 mm SL), and one shrimp. The only

species collected with Hollandichthys sp. n. C was Phalloceros sp.

Distribution. This species in know from smaller tributary of the rio Morato and rio Braço do

Macaco, Guaraqueçaba, Paraná, Brazil (Fig. 1).

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 Hollandichthys sp. n. D

(Fig. 29)

Holotype. MZUSP uncat (removed from MZUSP 63130), 96.4 mm SL, stream in the km 208

of road Rio-Santos, 5 km of Riviera São Lourenço (23o46’S 45o59’W), 5 Jan 1998,

Expedition MZUSP.

Paratypes. Brazil: São Paulo: Bertioga: MZUSP 63130, 11 of 16 (2 c&s), 30.6-75.9 mm

SL, 24 Out 2000, -MZUSP 53683, 16, 25.2-85.5 mm SL, same data of the holotype. -UFRJ

5442, 9, 26.2-63.75 mm SL, stream in praia de Guaratuba (approx. 23o45’S 45o50’W), 16

Nov 2001, W. Costa, C. Bove & B. Costa. São Paulo: São Sebastião: MCP 30658, 2, 76.5-

83.8 mm SL, stream in the Juqueí beach about 1 km of road SP 55 (23o45’S 45o42’W), 15 Oct

2002, V. A. Bertaco, J. F. Pezzi da Silva & P. Lehmann. -MCP 30659, 1, 94.7 mm SL,

córrego da Barra about 400 m of road BR 101, -MCP 30660, 2, 59.4-88.3 mm SL, córrego da

Barra (23o47’S 45o33’W), same date and collectors as MCP 30658. -MZUSP 21871, 4, 65.9-

88.6 mm SL, rio do Alto da Serra (23o48’S 45o25’W), Mar 1982, CESP. -MZUSP 45249, 22

(2), 29.3-58.4 mm SL, rio Juqueí in the Juqueí beach (23o45’S 45o42’W), 19 Jan 1993, F.

Lima. -MZUSP 49933, 3, 13.4-49 mm SL, tributary of rio Juqueí in the Juqueí beach

(23o45’S 45o42’W), 30 Jan 1994, F. Lima. -MZUSP 53687, 4, 34.8-40.3 mm SL, tributary of

rio Una, Barra do Una beach, km 182 Rio-Santos road, 29 Jan 1998, F. Lima & E. Baena. -

MZUSP 54167, 4, 24.1-75.6 mm SL, rio Boiçucanga at Boiçucanga, 14 Aug 1995, A. Akama,

M. C. de Pinna & E. Trajano. -MZUSP 54475, 7, 21.6-57.3 mm SL, rio das Pedras, Camburi

beach, 13 Aug 1995, A. Akama, M. C. de Pinna & E. Trajano.

Diagnosis. The body color pattern just above anal fin, covered with small black dots (Fig. 29)

(more clearly discernible in larger specimens) instead of longitudinal rows, distinguishes

Hollandichthys sp. n. D from all other species of the genus. The shape of the adipose fin,

91
rounded and elevated (Fig. 29), also distinguish this species from all other species with more

elongate adipose fins.

Description. Morphometric data for Hollandichthys sp. n. D are summarized in Table 9.

Body compressed and elongate; greatest body depth usually anterior to dorsal-fin origin.

Dorsal head profile nearly straight to gently concave. Dorsal profile from supraoccipital bone

to base of last dorsal-fin ray slightly convex, and from this point to adipose-fin origin straight.

Ventral profile from the anterior tip of lower jaw to pelvic-fin origin convex, and straight to

anal-fin origin. Body profile along anal-fin base of sexually mature males slight convex, and

straight in females. Caudal peduncle elongate, nearly straight to slightly concave along dorsal

and ventral margins.

Snout gently convex or nearly straight. Head small. Mouth slightly superior. Lower

jaw protruding strongly anterior to upper jaw. Maxilla long and slightly curved aligned at

angle of approximately 45 degrees to longitudinal body axis, with anterodorsal portion

convex and posterodorsal portion concave, and all toothed portion smoothly concave. Maxilla

anterior portion narrow; posterior portion enlarged.

Premaxilla with two tooth rows; outer row with 2-3, tricuspid teeth with central cusp

longer; inner row teeth 5, tricuspid, first longer than the remaining, with central cusp three or

four times longer and broader than other cusps. Maxilla with 12-19 (usually 15-17), tricuspid

teeth (teeth 1 or 3 last with conical), with central cusp longer. Four or 5 anteriormost dentary

teeth larger, with 3-4 cusps (fourth cusp diminute), followed by medium sized tooth with 3

cusps, and 11-16 teeth with 2-3 cusps or conical; central cusp in all teeth three or four times

longer and broader than other cusps. All cusp tips slightly curved posteriorly towards mouth

gape (Fig. 20).

92
 Dorsal-fin rays ii, 9 (ii, 9; one specimen ii, 8; n = 58); first unbranched ray

approximately one-half length of second ray. Dorsal-fin origin located posterior to middle of

SL and posterior to vertical through pelvic-fin origin. Dorsal-fin profile rounded. Adipose-fin

rounded in larger specimens, and located approximately at vertical through insertion of 20th to

22th anal-fin ray. Unbranched and anterior 3 to 4 dorsal-fin rays bearing a large posterior flap

along their posterolateral borders, more developed in larger specimens of both sexes, and

extending along entire length of first unbranched ray and proximal midlength of second

unbranched and anterior branched rays.

Anal-fin rays iv-v, 27-30 (iv, 29; rarely 26 or 31, x = 28.5, n = 58). First unbranched

ray normally only apparent in cleared and stained specimens. Anal-fin profile nearly straight

in females, and males show anal-fin profile nearly straight except for a concavity

corresponding approximately to 21th-26th ray. Three or fourth last branched anal-fin rays

longer than the remaining, being the penultimate ray the longest. Anal-fin origin at vertical

through bases of third to fourth dorsal-fin branched ray. Anal-fin rays of sexually mature

males bearing one pair of small, elongate, retrorse bony hooks along posterolateral border of

each segment of lepidotrichia, along 4-29 (usually 5-26) branched rays; hooks more numerous

along 9th through 20th branched rays, rarely present on 29th ray. Hooks usually located along

posteriormost branched and distal 1/2 to 2/3 of each ray. Unbranched and anterior 5-7

branched anal-fin rays in males and 3-4 branched anal-fin rays in females bearing a large

posterior flap along their posterolateral borders, more developed in larger specimens of both

sexes, and extending along entire length of anterior unbranched rays and proximal midlength

of last unbranched and anterior branched rays.

Pectoral-fin rays i, 12-15 (i, 14; x = 13.8, n = 53). Pectoral-fin tip reaching pelvic-fin

origin in males but not in females. Pelvic-fin rays 6, i (6, i; rarely 5, i, n = 58). Pelvic-fin tip

reaching beyond anal-fin origin in males but not in females. Pelvic-fin origin located anterior

93
to vertical through dorsal-fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook

per segment of lepidotrichia along ventromedial border of third to fifth branched rays.

Caudal-fin forked, with 19 principal rays (19; n = 51). Caudal-fin lobes profile

rounded. Dorsal procurrent rays 12-15 ( x = 13.2, n = 34). Ventral procurrent rays 10-12 ( x =

10.6, n = 34). Presence of small bony spines in the dorsal surface of dorsal unbranched

principal caudal-fin ray, and in the ventral surface of the ventral unbranched principal caudal-

fin ray in adults (larger than 50.0 mm SL).

Scales cycloid, moderately large. Lateral line incomplete, perforated scales 7-11 (6;

one specimen 6 and 12, x = 8.8, n = 56). Longitudinal series of scales including lateral-line

scales 38-44 (43; rarely 36 or 46, x = 41.5, n = 52). Scale rows between dorsal-fin origin and

lateral line 5-7 (6; x = 6.2, n = 53); scale rows between lateral line and pelvic-fin origin 4-6

(5; x = 5.3, n = 53). Predorsal scales 13-17 (15; usually 14-15), arranged in irregular series

( x = 15). Scales rows around caudal peduncle 14-16 [14] ( x = 15.1). Scale sheath along anal-

fin base 8-15 scales in single series, extending to base of sixth to thirteen branched rays. Body

scales covering the belly with posterior margin truncate or concave.

Precaudal vertebrae 16-17; caudal vertebrae 20-21; total vertebrae 36-37 (n = 4).

Supraneurals 6-7, (n = 4). Gill-rakers 7-8/11 (n = 3).

Color in alcohol. See figure 29 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually yellow on sides.

Black dots in the body portion near anal fin base (more visible in adults); body with multiple

series of black longitudinal stripes positioned along areas of overlap of contiguous horizontal

rows of body scales. Adult males and females with one humeral spot located over 7 to 9

lateral line scales and extending over first longitudinal series of scales above lateral line.

Small black chromatophores scattered over rays of all fins. Adipose-fin smoothly pigmented

94
in its border, and more pigmented in the anterior portion. Specimens larger than 44.0 mm SL

with one small humeral spot located over seventh to ninth perforated lateral line scales and

extending over the horizontal series of scales just above lateral line. Color pattern of juveniles

(smaller than 30.5 mm SL) similar to that of adults (larger than 66.0 mm SL), but with two

black diffuses humeral spots; anterior located over first to fourth perforated lateral line scales

and extending over 1-2 horizontal series of scales, including lateral line. Second humeral spot

located over seventh to eighth perforated lateral line scales and extending over 1-2 horizontal

series of scales, including lateral line. The specimens collected in the coastal streams at

Bertioga, SP, have body color and longitudinal lines more dark.

Color in life. Description based in lots MCP 30659 and MCP 30660 (São Sebastião). Color

pattern similar to described for alcohol preserved specimens, except as follows. Dorsal,

caudal, anal, pelvic and pectoral fins red-orange pigmented. Adipose fin yellowish. Anal-fin

with small black chromatophores along its border forming a narrow stripe in some specimens.

Body yellowish and orange.

Sexual dimorphism. Males of Hollandichthys sp. n. D are easily recognized by the presence

of bony hooks on the anal- and pelvic-fin branched rays, by the shape of the anal fin distal

border, and by the longer pectoral and pelvic fins (see description). Caudal peduncle depth is

slightly larger in males than females, and procurrent caudal-fin rays are more prominent in

males.

Habitat description. The specimens of the lots MCP 30658, MCP 30659 and MCP 30660

were caught in lentic waters of shallow small streams, with transparent water, sand and stones

bottom, and lots of riparian vegetation. All specimens were hidden under pendent plants in

95
margin of stream. Other species collected along with Hollandichthys sp. n. D were A.

tajasica, Eleotris pisonis, G. brasiliensis, G. pantherinus, Oostethus lineatus, Phalloceros sp.

and S. guentheri.

Distribution. This species in know from coastal streams in the road from Rio de Janeiro to

Santos, between Bertioga and São Sebastião, São Paulo, Brazil (Fig. 1).

Hollandichthys sp. n. E

(Fig. 30)

Holotype. Brazil: Rio de Janeiro: Parati: MCP 32039, 83.8 mm SL, male, córrego da Toca

do Boi, about 50 m of lodge of the Condomínio Larangeiras (23o19’S 44o40’W), 17 Oct 2002,

V. A. Bertaco, J. F. Pezzi da Silva & P. Lehmann.

Paratypes. Brazil: Rio de Janeiro: Parati: MCP 30664, 59, 35.5-79.5 mm SL, collected

with the holotype. -MNRJ 14846, 14 (2 c&s), 23.3-84.3 mm SL, same locality of the

holotype, 3 Mar 1989, Ictiologia. -MNRJ 20247, 20 of 41 (3 c&s), 30.6-76.6 mm SL, same

locality of the holotype, 23 Apr 2000, M. Melo & P. A. Buckup.

Additional material (non-types). Brazil: São Paulo: Ilhabela: Ilha de São Sebastião:

MCP 30661, 18, 9.2-53.2 mm SL, stream in the road for praia de Castelhanos (23o49’S

45o21’W), 16 Oct 2002. -MZUSP 49941, 9 (2 c&s), 20.0-60.8 mm SL, same locality of MCP

30661, 30 Oct 1993. -MZUSP 49943, 6, 31.5-58.1 mm SL, same locality of MCP 30661, 4

Apr 1994. -MCP 30662, 7, 55.5-70.4 mm SL, stream in praia de Perequê (23o49’S 45o21’W),

16 Oct 2002. São Paulo: Ubatuba: MCP 30663, 4, 60.8-67.2 mm SL, Cachoeira

Ipiranguinha, affluent of rio Grande, about 2.4 km of SP 125 road, (23o25’S 45o7’W), 17 Oct

2002. -MZUSP 18450, 6, creek streams affluent of the right margin near mouth of rio da

Lagoa Itaguá (23o26’S 45o4’W), Jan 1974. -MZUSP 18542, 2, 50.2-58.7 mm SL, ribeirão

96
Ipiranguinha (23o25’S 45o6’W), 6 Mai 1977. -MZUSP 27114, 19 (2 c&s), 32.4-65.1 mm SL,

creek between the Jardim Acaraú and Rio de Janeiro-Santos road (23o26’S 45o4’W), 24 Jul

1975. -MZUSP 27505, 2, 52.4-68.7 mm SL, rio Silva in the SP 125 road, between Ubatuba to

Taubaté (approx. 23o12’S 45o20’W), 27 Oct 1982. -MZUSP 39988, 1, rio Indaiá (23o24’S

45o3’W), Apr 1984 to Mai 1985. -UFRJ 3382, 5, 48.3-87.7 mm SL, rio do Surdo, Ubatumirim

(approx. 23o21’S 44o52’W), 13 Oct 1985. Rio de Janeiro: Parati: (locality of Tarituba):

MCP 30665, 1, 62.3 mm SL, rio São Roque in BR 101 road (23o4’S 44o41’W), 18 Oct 2002. -

MCP 30666, 11, 36.0-75.5 mm SL, stream affluent of rio Taquari (23o2’S 44o41’W), 18 Oct

2002. -MNRJ 21038, 10, 33.3-52.5 mm SL, stream in Tarituba (23o3’S 44o35’W), 20 Sep

1972. -UFRJ 0587, 4, 64.8-69.4 mm SL, same locality of the MCP 30666, 23 Jul 1991. -UFRJ

0589, 12 of 13 (3 c&s), 36.7-77.8 mm SL, rio Iriri (23o3’S 44o35’W), 25-28 Jul 1991. Rio de

Janeiro: Angra dos Reis: MNRJ 5561, 10, 37.4-75.8 mm SL, rio Ariró Pequeno (23o1’S

44o19’W), 1948. -MNRJ 5608, 2, 54.8-60.5 mm SL, riacho Vermelho, affluent of rio

Mambucaba (23o1’S 44o32’W), 1948. -MNRJ 5991, 12 (2 c&s), 38.4-73.0 mm SL, fazenda

de Japuhyba (23o1’S 44o19’W), Aug 1942. -UFRJ 2148, 1, 34.0 mm SL, stream affluent of

rio da Guarda, in road Angra dos Reis to Lídice (22o55’S 44o19’W), 30 Dec 1992. -UFRJ

3983, 1, 47.9 mm SL, same locality of UFRJ 2148, Apr 1996. Rio de Janeiro: Mangaratiba:

MNRJ 10952, 9 (1 c&s), 49.7-94.5 mm SL, rio Itinguçu in Itacuruçá, Mangaratiba (22o54’S

43o53’W), 28 Jan 1954.

Diagnosis. Hollandichthys sp. n. E is distinguished from all species of the genus by the red-

orange pigmented fins. Hollandichthys sp. n. E is distinguished from Hollandichthys sp. n. A,

Hollandichthys sp. n. B and H. affinis by the presence of one black humeral spot in specimens

larger than 60 mm SL. Also differs from Hollandichthys sp. n. A and H. affinis by the larger

number of dorsal procurrent caudal-fin rays (12-14, x = 13, versus 8-11, x = 9.4, Fig. 18, and

97
10-13, x = 11.4, Fig. 19, respectively) and from Hollandichthys sp. n. A by the larger number

of ventral procurrent caudal-fin rays (9-11, x = 10.3, versus 7-8, x = 7.7, respectively, Fig.

19). Hollandichthys sp. n. E differ from Hollandichthys sp. n. C by absence of the large black

humeral spot in adults (larger than 50 mm SL), from H. multifasciatus by absence the

adipose-fin black pigmented along half proximal length, and from Hollandichthys sp. n. D by

the absence of black dots laterally in the body near the anal fin base. Hollandichthys sp. n. E

differ from H. perstriatus by the larger number scales in longitudinal series (40-46, x = 42.5,

n = 42, versus 39-43, rarely 36-38 or 44, x = 41.2, n = 62, respectively).

Description. Morphometric data for Hollandichthys sp. n. E are summarized in Table 10.

Body compressed and elongate; greatest body depth usually anterior to dorsal-fin origin.

Dorsal head profile nearly straight or slightly concave. Dorsal profile from supraoccipital

bone to base of last dorsal-fin ray convex, and from this point to adipose-fin origin straight.

Ventral profile of head convex. Ventral body profile slightly convex from pectoral-fin origin

to pelvic-fin origin, and straight to anal-fin origin. Body profile along anal-fin base

posterodorsally slanted. Caudal peduncle elongate, nearly straight to slightly concave along

dorsal and ventral margins.

Snout gently convex or nearly straight. Head small. Mouth slightly superior. Lower

jaw protruding anteriorly to upper jaw. Maxilla long and slightly curved aligned at angle of

approximately 45 degrees to longitudinal body axis, with anterodorsal portion convex and

posterodorsal portion concave, and all toothed portion smoothly concave. Maxilla anterior

portion narrow; posterior portion enlarged.

Premaxilla with two tooth rows; outer row with 2-3, tricuspid teeth with central cusp

longer; inner row teeth 5 with 3-5 cusps (fourth or fifth cusp diminutive); central cusp three or

four times longer and broader than other cusps; medial tooth longer than the remaining.

98
Maxilla with 11-20 (usually 14-16) tricuspid teeth, with central cusp longer. Four or 5

anteriormost dentary teeth larger, with 3-4 cusps (fourth cusp diminute), followed by medium

sized tooth with 3 cusps, and 12-14 teeth with 2-3 cusps or conical; central cusp in all teeth

three or four times longer and broader than other cusps. All cusp tips slightly curved

posteriorly towards mouth gape.

Dorsal-fin rays ii, 9 (ii, 9; one specimen ii, 10, n = 41); first unbranched ray

approximately one-half length of second ray. Dorsal-fin origin located posterior to middle of

SL and posterior to vertical through pelvic-fin origin. Dorsal-fin profile rounded. Adipose-fin

located approximately at vertical through insertion of 22th to 24th anal-fin rays. Unbranched

and anterior 3 to 4 dorsal-fin rays bearing a large posterior flap along their posterolateral

borders, more developed in larger specimens of both sexes, and extending along entire length

of first unbranched ray and proximal midlength of second unbranched and anterior branched

rays.

Anal-fin rays iv-vi, 27-30 (iv, 28; x = 28.4, n = 42). First unbranched ray normally

only apparent in cleared and stained specimens. Anal-fin profile nearly straight in females.

Anal-fin profile nearly straight along most of its border in males, except for a concavity in the

posterior portion of the fin, corresponding approximately to 22th to 25th ray; three or four last

branched anal-fin rays longer than the remaining, forming a conspicuous anal-fin lobe;

penultimate ray the longest. Anal-fin origin at vertical through the base of third to fourth

dorsal-fin branched rays. Anal-fin rays of sexually mature males bearing one pair of small,

elongate, retrorse bony hooks along posterolateral border of each segment of lepidotrichia,

along 4 to 28 (usually 5-25) branched rays; hooks more numerous along 8th through 21th

branched rays, rarely present on 28th ray. Hooks usually located along posteriormost branched

and distal 1/2 to 2/3 of each ray. Unbranched and anterior 2 to 3 anal-fin rays bearing a large

posterior flap along their posterolateral borders, more developed in larger specimens of both

99
sexes, and extending along entire length of anterior unbranched rays and proximal midlength

of last unbranched and anterior branched rays.

Pectoral-fin rays i, 13-16 (i, 14; x = 14.5, n = 42). Pectoral-fin tip reaching pelvic-fin

origin in males but not in females. Pelvic-fin rays 6, i (6, i; rarely 5, i, n = 42). Pelvic-fin tip

extends beyond anal-fin origin in males but not in females. Pelvic-fin origin anterior to

vertical through dorsal-fin origin. Pelvic fin of males usually bearing 1 retrorse bony hook per

segment of lepidotrichia along ventromedial border of third to fifth branched rays (Fig. 13).

Caudal-fin forked, with 19 principal rays (19; one specimen 18, n = 42). Caudal-fin

lobes profile rounded. Dorsal procurrent caudal-fin rays 12-14 ( x = 13, n = 17). Ventral

procurrent caudal-fin rays 9-11 ( x = 10.3, n = 17). Presence of the bony spines very

developed in the dorsal surface of dorsal unbranched caudal-fin ray, and in the ventral surface

of the ventral unbranched principal caudal-fin ray in larger specimens. Also present bony

spines in the dorsal surface of last dorsal procurrent ray and in the ventral surface of last

ventral procurrent ray in specimens larger than 50.0 mm SL.

Scales cycloid, moderately large. Lateral line incomplete, perforated scales 6-9 (9; x =

7.8, n = 42). Longitudinal series of scales including lateral-line scales 40-46 (42; x = 42.5, n =

42). Scale rows between dorsal-fin origin and lateral line 6-7 (6; usually 6, x = 6.2, n = 42);

scale rows between lateral line and pelvic-fin origin 5-6 (6; usually 6, x = 5.9, n = 42).

Predorsal scales 14-16 (15; x = 14.8), arranged in irregular series. Scales rows around caudal

peduncle 16. Scale sheath along anal-fin base 8-15 scales in single series, extending to base of

sixth to twenty branched rays. Body scales covering the belly with posterior margin truncate

or concave.

Precaudal vertebrae 16-17; caudal vertebrae 20; total vertebrae 36-37, usually 37 (n =

5). Supraneurals 6 (n = 5). Gill-rakers 7/10 (n = 2).

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Color in alcohol. See figure 30 for the preserved color patterns of adult males and females.

Ground color brown in dorsal region of body and head, becoming gradually light brown on

sides. Black longitudinal lines usually forming a zigzag pattern between longitudinal rows of

scales; more conspicuous in middle portion of body. Small black chromatophores scattered

over rays of all fins. Adipose-fin smoothly pigmented in the border, and more pigmented in

the anterior portion. Specimens larger than 60.0 mm SL with one humeral spot located over

seventh to ninth perforated lateral line scales and extending over the horizontal series of

scales just above lateral line. Specimens measuring 50.0-60.0 mm SL with one or two faint

humeral spots. Color pattern of juveniles (smaller than 48.0 mm SL) similar to that of adults,

but with two diffuse black humeral spots; anterior located over first to fourth perforated

lateral line scales and sometimes extending over the horizontal series of scales just above

lateral line. Second humeral spot darker located over seventh to eighth perforated lateral line

scales and sometimes extending over the horizontal series of scales just above lateral line.

Color in life. Color pattern similar to described for alcohol preserved specimens, except as

follows. All fins orange pigmented. Anal-fin with small black chromatophores along its

border forming a narrow stripe in some specimens. Adipose fin brilliant orange pigmented,

with small chromatophores along its border. Body black yellowish or orange.

Sexual dimorphism. Males of Hollandichthys sp. n. E are easily recognized by the presence

of bony hooks on the anal- and pelvic-fin branched rays, by the shape of the anal fin distal

border, and by the longer pectoral and pelvic fins (see description). Caudal peduncle depth is

slightly larger in males than females, and procurrent caudal-fin rays are more prominent in

males.

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Habitat description. The specimens of the lots MCP 30664 were caught in lentic or faint

current water of shallow streams, with transparent water, sand and some stones bottom, and

very riparian vegetation. All specimens were hidden under pendent plants or tree-trunks in

water near margin of stream.

Ecology. Stomach contents of one specimen (68.2 mm SL, MNRJ 14846) prepared for

clearing and staining consisted of one spider, one grasshopper, and insect parts. The only fish

species collected with Hollandichthys sp. n. E was Phalloceros sp.

Distribution. This species is known from coastal streams between Ilhabela, Ilha de São

Sebastião, São Paulo and Mangaratiba, Rio de Janeiro, Brazil (Fig. 1).

Remarks. The population of Hollandichthys sp. n. E collected in the rio Itinguçu, Itacuruçá,

Mangaratiba, Rio de Janeiro (MNRJ 10952, 9 specimens) has the largest number of branched

anal-fin rays (30-33, x = 30.9, n = 9, versus 26-31, rarely 32-33, x ≤ 29.4, in other

Hollandichthys species); and largest number of scales in longitudinal series (43-45, x = 44.3,

n = 8, versus 35-43, rarely 44-46, x ≤ 42.5, in other Hollandichthys species). These specimens

were collected nearly the middle of the last century, and are relatively well preserved, but its

color pattern is faint (pale). The small number of examined specimens avoid the recognition

of this population as a distinct new species. This population is the septentrional limit of the

genus. Larger specimens of Hollandichthys sp. n. E present bony spines very developed in the

dorsal surface of dorsal unbranched principal caudal-fin ray, and in the ventral surface of

ventral unbranched principal caudal-fin ray, also present in the dorsal surface of last dorsal

procurrent caudal-fin ray, and in the ventral surface of last ventral procurrent caudal-fin ray.

In a recent collecting expedition (October/2002) to rio Itinguçu, Mangaratiba, RJ, we could

102
observe habitat degradation. The river presents shallow and lentic water, sand and ooze

bottom, with some rock and stones. The water was polluted with garbage, and lacking most of

the riparian vegetation. In the upper portion of the river, a small dam was constructed for

water supply. None specimen of the Hollandichthys was caught, and some specimens of the

Phalloceros sp., and several specimens of the black tadpole were observed in a stretch of this

river.

Biogeography of Hollandichthys

All species of the genus Hollandichthys inhabit coastal rivers and marine islands from

Northern Rio Grande do Sul State to Southern Rio de Janeiro State, and the upper rio Tietê in

the continental upper rio Paraná drainage (Fig. 1). Such a distribution pattern strongly

corroborates the recognition of a large area of endemism comprising the coastal drainages of

South and Southeastern Brazil, including rio Tietê headwaters. This is similar or largely

congruent with the distribution given to the species of the genera Deuterodon Eigenmann

(1907) by Lucena & Lucena (1992; 2002), Mimagoniates Regan (1907) by Menezes &

Weitzman (1990), Spintherobolus Eigenmann (1911) by Weitzman & Malabarba (1999), and

Hemipsilichthys Eigenmann & Eigenmann (1889) by Pereira & Reis (2002).

Along the distribution of Hollandichthys, some sub-regions can be recognized. The

most evident, and inhabited by the most basal species of the genus, Hollandichthys sp. n. A, is

the area comprised by the rio Maquiné and rio Três Forquilhas (rio Tramandaí drainage), and

rio Mampituba. Malabarba & Isaia (1992) identified the area comprising these rivers as an

area of fish species endemism, later extended to the next northern drainage of the rio

Araranguá by evidence presented by Reis & Schaefer (1999). We have no records of any

Hollandichthys species from the rio Araranguá drainage, but this is possibly due to the lack of

collection efforts in the specific habitats were these fishes are found. The taxa with

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geographic distribution similar to Hollandichthys sp. n. A and that support the recognition of

an area of endemism comprising the rio Maquiné, rio Três Forquilhas, rio Mampituba and rio

Araranguá are the characid Mimagoniates rheocharis described by Menezes & Weitzman

(1990), Deuterodon stigmaturus (Gomes, 1947) by Lucena & Lucena (1992; 2002), one

species of an undescribed Characidae genus (Malabarba & Weitzman, in press); the loricariid

genus Epactionotus described by Reis & Schaefer (1999), and two monophyletic groups of

loricariids of the genus Hemipsilichthys (Pereira & Reis, 2002); and the characid genus

Odontostoechus Gomes (1947), with one species from the rio Tramandaí and rio Mampituba

drainages, and a closely related species from the rio Araranguá drainage (Malabarba, 1998).

A distribution pattern similar to H. affinis, in coastal streams between Joinvile, SC and

Morretes, PR, is found in Deuterodon langei (Lucena & Lucena, 2002), and Spintherobolus

ankoseion described by Weitzman & Malabarba (1999).

The rio Ribeira do Iguape and adjacent coastal drainages also seems to form an area of

endemism, with the occurrence of H. perstriatus along with Spintherobolus leptoura

described by Weitzman & Malabarba (1999), Deuterodon iguape Eigenmann (1907) (Lucena

& Lucena, 1992; 2002), two species of the endemic loricariid genus Isbrueckerichthys Derijst

(1996) (Pereira & Reis, 2002), and of two species of Pimelodella Eigenmann & Eigenmann

(1888), P. kronei (Miranda-Ribeiro, 1907) and P. transitoria Miranda-Ribeiro (1907)

(Guazelli, 1997).

Hollandichthys multifasciatus has a small geographical distribution but encompassing

two major drainage systems of South America, the upper rio Tietê, a tributary of the upper rio

Paraná, La Plata drainage, and a few coastal Atlantic drainages of southeastern Brazil, near

Cubatão and Santos, SP (Fig. 1). Common patterns of distribution between the upper rio Tietê

and the coastal drainages have been found for the genus of Spintherobolus by Weitzman &

Malabarba (1999:39-40), and for Hyphessobrycon bifasciatus, H. reticulatus,

104
Pseudocorynopoma heterandria and G. pantherinus by Langeani (1989:189). Langeani

(1989), based on these faunal similarities, also suggested that both the upper rio Tietê and the

coastal drainages of São Paulo state were connected in the past. M. C. Malabarba (1998) has

consistently hypothesized past connections between the rio Tietê and coastal drainages based

on the phylogeny of fossil and recent species of the clades Lignobrycon + Triportheus and

Spintherobolus + Megacheirodon distributed in the area.

No taxa were found with geographic distribution similar to Hollandichthys sp. B,

Hollandichthys sp. C, Hollandichthys sp. D, and Hollandichthys sp. E.

The speciation through isolation of formerly widespread species in south and

southeastern Brazilian coastal drainages is possibly fostered by changes in sea level as

discussed by Weitzman et al. (1988). Most populations of Hollandichthys species live in

small creeks or lateral puddles isolated in streams and rivers of the Atlantic Forest found

nearly sea level. Some populations (H. affinis, Ilha de São Francisco do Sul, SC and

Hollandichthys sp. n. D, São Sebastião, SP), were captured in brackish water together with

Awaous tajasica, Eleotris pisonis and Oostethus lineatus. According to Britski (1972) H.

multifasciatus has been caught in brackish water and may be the Characoidei more resistant

the saltiness variation. We suggest that this evidence together ecological changes and lowland

interchange of freshwaters during flood periods can have favored, in the recent past, the

dispersion and speciation of some populations of Hollandichthys along of the all distribution

area of genus. This may also explain its occurrence in sea islands along the coast.

The distribution pattern of each Hollandichthys species, along with the hypothesis of

relationships presented herein, allows the analysis of historical connections between these

coastal drainages. The basal position of Hollandichthys sp. n. A as a sister species to all other

Hollandichthys species supports the hypothesis of an ancient isolation of the southernmost

drainages, namely rio Maquiné, rio Três Forquilhas and rio Mampituba (possibly also rio

105
Araranguá - see discussion of areas of endemism above) from the remaining drainages. This

vicariant event would be followed by the isolation of a group of populations from Garopaba

and Ilha de Santa Catarina, corresponding to Hollandichthys sp. n. B, and of a group of

populations from the coastal streams between Joinvile, SC and Morretes, PR, corresponding

to H. affinis, from the remaining populations distributed to the North, up to Southern Rio de

Janeiro. Vicariant events isolating each of the five species recognized in the Northern

distribution of the genus (Fig. 1), namely Hollandichthys sp. n. C, H. perstriatus, H.

multifasciatus, Hollandichthys sp. n. D and Hollandichthys sp. n. E, would be the most recent

of those.

The phylogeneticall analysis also hypothesizes close relationships between the

Hollandichthys Clade, distributed along the Atlantic Brazilian coastal drainages between the

states of Rio Grande do Sul and Rio de Janeiro, and the Pseudochalceus Clade, distributed in

the Pacific drainages of Ecuador and Colombia, in the Western side of Andes. No related

fishes are found in the area between. Such a distribution pattern is not exclusive to this group

of fishes and is congruent with that described for the Cyprinodontiforms fishes of the family

Anablepidae and the Characid genera Cheirodon Girard (1854) and Nanocheirodon of the

Cheirodontinae. The genus Jenynsia Günther (1866) of the Anablepidae is distributed

exclusively along the Atlantic drainages of Argentina, South Bolivia, Uruguay, South and

Southeastern Brazil, while its sister group genus Anableps Scopoli (1777) occurs in Central

America and Northern South America, in both sides of Andes (Ghedotti, 1998). The genus

Cheirodon has a distribution pattern similar to Jenynsia, occurring along the Atlantic

drainages of Argentina, Uruguay, South and Southeastern Brazil, but also occurring in the

Pacific drainages of Chile in Southern South America, while its sister group genus

Nanocheirodon is found exclusively in the western side of the Andes, in Colombia

106
(Malabarba, 1998). Again, no related fishes are found in the area between the distribution of

these sister clades, found in Southeastern and Northeastern South America, in these examples.

DISCUSSION

The comparative analysis of almost all available material of Hollandichthys, along

with new collection efforts along the coastal drainages of South and Southern Brazil, allows

the recognition of eight species in the genus, while only one has been recognized since the

proposition of the genus by Eigenmann (1909). This high diversity was found closely

associated to the Atlantic Forest, in such a way remaining Hollandichthys populations were

found only in well preserved areas or little refuges of still forested places, during all field

expeditions performed in the last two years. Also notable is the discovery and description of

the most basal species of the genus (Hollandichthys sp. A), whose first specimens were

collected only four years ago (1999). Although coastal drainages of South and Southeastern

Brazil are relatively well explored, as indicated by various taxa review (e. g. Menezes &

Weitzman, 1990; Malabarba & Isaia, 1992; Lucena & Lucena, 1992, 2002; Pereira & Reis,

2002; Reis & Schaefer, 1999; Weitzman & Malabarba, 1999), the recognition of undescribed

species in those drainages indicated that a large effort to study of the fish fauna is still

necessary in areas or environments moderately explored.

List of the comparative material.

Astyanax lineatus: MCP 15617, 5 (1 c&s), 42.8-88.8 mm SL, stream at road between

Cuiabá/Cáceres, Cáceres, Mato Grosso, Brazil; MCP 15569, 2, 30.9-55.7 mm SL, tributary of

rio do Bugre, km 165 of BR 174, Porto Esperidião, Mato Grosso, Brazil. Brycon pesu: MCP

17092, 1 c&s, 54.9 mm SL, rio Tapajós, São Luiz, Pará, Brazil. Bryconops sp.: MCP 17077, 1

c&s, 57.7 mm SL, rio Tapajós at Itaituba, Pará, Brazil. Bryconops melanurus: MCP 15807, 46

107
(3 c&s), 40.2-82.2 mm SL), stream at 13 km S of Nova Olímpia, Barra dos Bugres, Mato

Grosso, Brazil. Diapoma terofali: MCP 9071, 6 c&s, 38.1-43.3 mm SL, rio Santa Maria on

km 246 of road BR 293, Dom Pedrito, Rio Grande do Sul, Brazil; MCP 16391, 26, 23.5-42.5

mm SL, rio Negro, Bagé, Rio Grande do Sul, Brazil. Aphyocharax anisitsi: MCP 9261, 9 (1

c&s), 24.5-29.5 mm SL, rio Jaguari, São Francisco do Sul, Rio Grande do Sul, Brazil; MCP

9272, 2 c&s, 22.3-27.3 mm SL, stream between Santa Maria/Mata, Rio Grande do Sul,

Brazil; MCP 16212, 60, 26.5-28.5 mm SL, praia da Formosa, Uruguaiana, Rio Grande do Sul,

Brazil. Hemibrycon dariensis: MCP 27072, 12 (1 c&s), 19.8-44.7 mm SL, rio Pirre, El Real,

Darién, Panamá; MCP 27074, 19 (1 c&s), 23.6-41.0 mm SL, rio Pucuro, above of rio Tuira,

Pucuro, Darién, Panamá. Nematobrycon palmeri: CAS 70883, 43 (2 c&s), 10.2-28.2 mm SL,

rio Condoto, tributary of rio San Juan, Departamento Choco, Colombia; MHNG 2182.86, 7 (2

c&s), 25.0-30.2 mm SL, Colombia (aquarium center). Nematocharax venustus: MCP 17773,

8 (1 c&s), 25.6-51.7 mm SL, rio Branco, Arataca, Bahia, Brazil; MCP 17977, 49 (2 c&s),

15.1-31.6 mm SL, rio Jequitinhonha, Itaobim, Minas Gerais, Brazil; MCP 17978, 7, 40.4-47.3

mm SL, rio California near mouth of rio Água Fria, Itambé, Bahia, Brazil; MCP 17987, 29 (3

c&s), 15.9-44.7 mm SL, tributary of rio Pratas at São José, Buerarema, Bahia, Brazil; MCP

17991, 30 (2 c&s), 16.1-51.8 mm SL, rio Branco, tributary of rio Una, Camaçã, Bahia, Brazil.

Odontostilbe fugitiva: INPA 18465, 73 (4 c&s), 30.8-36.1 mm SL, rio Solimões, Ilha de

Marchantaria, Manaus, Amazonas, Brazil; INPA 18510, 100, 16.5-28.9 mm SL, rio Solimões,

Manaus, Amazonas, Brazil. Poptella paraguayensis: MCP 10960, 2 c&s, 49.0-51.2 mm SL,

stream at Transpantaneira road, Poconé, Mato Grosso, Brazil; MCP 15717, 18, 36.5-45.3 mm

SL, stream on MT 124 road, Tangará da Serra/Barra dos Bugres, Nova Olímpia, Mato

Grosso, Brazil. Pseudochalceus kyburzi: USNM 324462, 9 of 11 (4 c&s), 37.8-64.8 mm SL

(aquarium center). Pseudochalceus lineatus: ANSP 75905, 1, 42.4 mm SL, rio Santiago,

Esmeraldas, Ecuador; ANSP 75906, 1, 80.2 mm SL, rio Toachi near Santo Domingo of Los

108
Colorados, Pichincha, Ecuador; ANSP 75907, 1, 48.4 mm SL, rio Cupa, Esmeraldas,

Ecuador; MCZ 48730, 2, 37.3-61.2 mm SL, tributary of rio Branco, Pichincha, Ecuador;

USNM 287747, 5 (1 c&s), 24.6-43.7 mm SL, rio Chirape tributary of rio Guayllabama, R.

Esmeraldos de Paraiso, Ecuador. Pseudochalceus longianalis: ANSP 140067, 1 paratype,

34.3 mm SL, rio Guiza tributary of rio Mira, Colombia; MHNUC 113, 2, 63.2-73.1 mm SL;

MHNUC 137, 1, 42,5 mm SL; MHNUC 139, 6, 18.5-22.5 mm SL, stream Cantadelicia

tributary of rio Guapí, Guapí, Cauca, Colombia; ICNMHN 4965, 4, 34.0-40.9 mm SL;

ICNMHN 2338, 5, 39.4-59.6 mm SL, stream Angostura tributary of rio Mira, Narino,

Colombia. Pseudochalceus sp.: MHNG 2536.96, 22, 20.6-48.7 mm SL; MCP 31914, 2 c&s,

36.7-39.0 mm SL; tributary of rio Mira, border with Colombia; USNM 287746, 4 (1 c&s),

36.8-58.4 mm SL, Esmeraldas, Ecuador. Rachoviscus crassiceps: MZUSP 18564, 8 (1 c&s),

18.6-23.4 mm SL, praia de Brejatuba, Guaratuba, Paraná, Brazil; MZUSP 35732, 3 (1 c&s),

13.6-14.5 mm SL; MHNG 2188.72, 1, 14.1 mm SL, praia de Guaratuba, Guaratuba, Paraná,

Brazil. Rachoviscus graciliceps: MHNG 2514.34, 4 (1 c&s), 13.8-16.6 mm SL, stream at 13

km S of Prado, Bahia, Brazil; MZUSP 38366, 6 (1 c&s), 11.5-19.3 mm SL, tributary of rio

São José, Reserva Florestal de Soretama, Espírito Santo, Brazil.

ACKNOWLEDGMENTS

We are grateful to Mark Sabaj and John Lundberg (ANSP), David Catania (CAS),

Francisco Langeani (DZSJRP), Barry Chernoff (FMNH), Ivan Mojica (ICNMHN), Karsten

Hartel (MCZ), Vinicius Abilhoa (MHNCI), Claude Weber (MHNG), Pablo Lehmann

(MHNUC), Paulo Buckup (MNRJ), Osvaldo Oyakawa (MZUSP), Helmut Wellendorf

(NMW), Wilson Costa (UFRJ), Susan Jewett (USNM) for selection and loan of specimens; to

the Centro de Microscopia e Microanálises - CEMM, PUCRS for the SEM preparations.

Fernando Becker, José Pezzi da Silva, Juan Anza, Marco Azevedo, Pablo Lehmann, Verônica

109
Baumbach helped in field collecting. The senior author thanks the Conselho Nacional de

Desenvolvimento Científico e Tecnológico - CNPq for the student fellowship and the Museu

de Ciências e Tecnologia da PUCRS for the support. Thanks to PRONEX (Project

"Conhecimento, conservação e utilização racional da diversidade da fauna de peixes do

Brasil") and CNPq (Proc. 464545/00-5) by financial support for collecting expedition.

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