% * l *

UNIVERSIDADE FEDERAL DE LAVRAS

SPECIES FORMERLY IN THE GENERA

Trichillum HAROLD, 1868 AND
Pedaridium HAROLD, 1868

(COLEOPTERA: SCARABAEIDAE)

FERNANDO ZAGURY VAZ DE MELLO

2003





55560

FERNi» 4DO ZAGURY VAZ DE MELLO

SPECIES FORMERLY IN THE GENERA
Trichillum HAROLD, 1868 AND Pedaridium HAROLD,

1868 (COLEOPTERA: SCARABAEIDAE)

Dissertação apresentada à Universidade Federal
de Lavras, como parte das exigências do Curso
de Mestrado em Entomologia, para obtenção do
título de "Mestre".

Orientador

Prof. Júlio Neil Cassa Louzada

LAVRAS

MINAS GERAIS - BRASIL

2003

DESCARTADO

ELIfô

Data

L
;SíNATURA

U /JUJ1
BIBLIOTECA UNIVERSITÁRIA

ÜF»-A

vitor
New Stamp

vitor
New Stamp



Ficha Catalográfica Preparada pela Divisão de Processos Técnicos da
Biblioteca Central da UFLA

Vaz de Mello. Fernando Zagury
Species formerly in the genera Trichillum Harold, 1868 and Pedaridium

Harold, 1868 (Coleoptera: Scarabaeidae) / Fernando Zagury Vaz de Mello. --
Lavras : UFLA, 2003.

266 p.: il.

Orientador: Júlio Neil Cassa Louzada.

Dissertação (Mestrado) - UFLA.
Bibliografia.

1. Systematics. 2. Taxonomy. 3.Neotropical dung beeties. 4. Phylogeny. I.
Universidade Federal de Lavras. II. Título.

CDD-595.7649



FERNANDO ZAGURY VAZ DE MELLO

SPECIES FORMERLY IN THE GENERA

Trichillum HAROLD, 1868 AND Pedaridium HAROLD,
1868 (COLEOPTERA: SCARABAEIDAE)

Dissertação apresentada à Universidade Federal
de Lavras, como parte das exigências do Curso
de Mestrado em Entomologia, para obtenção do
título de "Mestre".

APROVADA em 26 de fevereiro de 2003

Dr. Sérgio Ide Instituto Biológico

Profa. Brígida Souza UFLA

(Orientador)

LAVRAS

MINAS GERAIS - BRASIL



I dedicate this dissertation to my wife, Silvia...

... and to my son Léo and his forthcoming brothers and sisters, hoping they will

live in a world where titles arenot so important, andthen dissertations and thesis

not so necessary.



Por onde, pelo comum, poder-se corrigir o ridículo ou o grotesco, até
levá-lo ao sublime; seja daí que seu entrelimite é tão tênue. E não será esse um
caminho por onde o perfeitíssimo se alcança? Sempre que algo de importante e
grande se faz, houve um silogismo inconcluso, ou, digamos, umpulo do cômico
ao excelso.

Conflui, portanto, que:

Os dedos são anéis ausentes?

Há palavras assim: desintegração...
Oaréo que não se vê, fora e dentro das pessoas.
O mundo é Deus estando em toda parte.
O mundo, para umateu, é Deus não estando nuncaem nenhuma parte.
Copo não basta: é preciso um cálice ou dedal com água, para as

grandes tempestades.
O O é um buraco não esburacado.

O que é - automaticamente?
O avestruz é umagirafa; só o que tem é que é umpassarinho.
Haja a barriga semo rei. (Isto é, o homem semalgum rei na barriga.)
Entre Abel e Caim, pulou-se um irmão começado por B.
Se o tolo admite, seja nem que um instante, que é nele mesmo que está o

que não o deixa entender, já começou a melhorarem argúcia.
A peninha no rabo do gato não é apenas "para atrapalhar".
Há uma rubra ou azul ausência no roxo (e no não roxo).
O copo com água até a metade: está meio cheio, ou meio vazio?
Saudade é o predomínio do que não está presente, diga-se, ausente.
Diz-se de um infinito - rendez-vous das paralelas todas.
O silêncio proposital dá a maior possibilidade de música.
Se viemos do nada, é claro que vamos para o tudo.
Veja-se, vezes, prefácio como todos gratuito.

Ergo:

O livro pode valer pelo muito que nele não deveu caber.

Quod erat demonstrandum."

(João Guimarães Rosa, Aletria e Hermenêutica, in Tutaméia - Terceiras
Estórias)



ACKNOWLEDGEMENTS

To Silvia, who accompanied this work since its very beginning, and so

had to support hearing about ali those horrible names, without knowing exactly

what was I talking about, but patiently trying to understand and suggesting

improvements, especially making new names less disgusting. I though it was not

necessary to tell that family, both old and new members, need to be

acknowledged by supporting me in similar ways but not so densely as Silvia

herself.

Silvia and Léo are responsible by mostevents that permitted me to forget

this work in criticai occasions, and that was essential to its completing.

A number of people helped me in the antecedents and during the course

of that work; I will try to name them ali, but surely, it will not be possible, so

please feel acknowledged if your name is nothere.

That work would notbe possible without thehelp and encouragement of

many people that was present since very early in my entomological life. Those

people were responsible for most of good things I learaed. Those are, in nearly

appearance order: Ângelo Machado, Celso Godinho, Ayr de Moura Bello,

Moacyr Alvarenga, Everardo Grossi, Antônio Martínez (in memoriam), Patrick

Arnaud, Maria Aparecida Vulcano, W. David Edmonds, Henry Howden, Brett

Ratcliffe, just forcitingthose prior to my undergrad.

The nucleus of this dissertation begun during my undergrad in Viçosa,

and many people there helped me a lot in many ways. Some of those are Zhé,

Duka, Carlos, Cristiano, Gabriela, Jovane and other people at Laboratório de

Ecologia de Comunidades, Unidade de Estudos em Ecologia de Comunidades

and Entomologia.



Mario Zunino taught me several new ways of examining and thinking
about scarabs, and, although it was not possible for me to apply that as I wanted
here, I hope to do so for life.

Alberto Ballerio is especially acknowledged by, apart of loaning
specimens, examining types at Paris Museum.

Without the possibility to going to Canada in 2000 it would not be

possible for me to see important types and to have avery important and special
advisor in the beginiiing ofthis project, that was François Génier. That trip and
logistical support there were possible by his efforts and by those by Bob
Anderson (and by extension to the Canadian Museum of Nature), Henry
Howden, Bruce Gill and my mother.

Moving to Lavras wasn't easy but that moving and finalization of the

work were possible thanks to the efforts of Júlio, Luísa, Marina, Juliana,

Berenice, Igor, Tatá, Teresa, Andreíza, Raquel, Jaqueline, Paulo, Mario, Malú,

Fernando, Maria Cristina, Brígida and people at both Laboratório de Ecologia
and Departamento de Entomologia.

Sérgio Ide, Cleide Costa and Fermín Martín-Piera (in memoriam) are
acknowledged byencouraging me inavery necessary way and moment.

A number of curators helped with specimens from collections under their

care, sometimes loaned to other people (who permitted me to view that

specimens), but most times to myself. Those curators and evolved intermediaries

are: José Verdú and Eduardo Galante (Universitat d'Alicant), François Génier
and Henry Howden (Canadian Museum of Nature, Aylmer), Eva Sprecher
(Naturhistorisches Museum Basel), Hella Wendt (Museum fíir Naturkunde der

Humboldt-Universitãt zu Berlin), Mareei Cludts (Institut Royal des Sciences
Naturelles de Belgique, BruxeUes), Otto Merkl (Magyar Természettudomány
Muzéum, Budapest), Axel Bachmann (Museo Argentino de Historia Natural

"Bernardino Rivadavia", Buenos Aires), Enio Cano and Jack Schuster



(Universidad dei Valle de Guatemala), Carlos A. H. Flechtmann (Faculdade de

Engenharia de Ilha Solteira da Universidade Estadual Paulista), Brett Ratcliffe

and Mary Liz Jameson (University of Nebraska State Museum, Lincoln),

Malcolm Kerley and François Génier (The Natural History Museum, London),

F. Dingemans-Backels and Alexey Tishechkin (Natuurhistorisch Museum

Maastricht), Antony Davies (Canadian National Collection of Insects, Ottawa),

Yves Cambefort, Olivier Montreuil, Alberto Ballerio and François Génier

(Museum National d'Histoire Naturelle, Paris), Roberto A. Zucchi and Luiz C.

Marchini (Escola Superior deAgricultura "Luiz deQueirós" da Universidade de

São Paulo, Piracicaba), Fernando R. Meyer and Paschoal Grossi (Museu

Anchieta, Porto Alegre), Maria Helena M. Galileo and José Henrique

Schoereder (Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre), Josef

Jelinek and François Génier (Národní Muzeum, Praha), Giovanni Onore and

Carlos Carpio (Pontifícia Universidad Católica dei Ecuador, Quito), Miguel A.

Monné and Paulo R. Magno (Museu Nacional da Universidade Federal do Rio

de Janeiro), Sérgio Ide and Júlio Louzada (Coleção Entomológica "Adolph

Hempel", Instituto Biológico, São Paulo), Ubirajara R. Martins, Carlos

Campaner, Sérgio Ide, José Henrique Schoereder and Júlio Louzada (Museu de

Zoologia da Universidade de São Paulo), Francisco Racca Filho and Paschoal

Grossi (Universidade Federal Rural do Rio de Janeiro, Seropédica), Arturo

Terán (Fundación Miguel Lillo, San Miguel deTucumán), Sacha Konstantinov,

Steve Lingafelter, Nancy Adams and Bruce Gill (United States National

Museum, Washington).

The following people following contributed loaning ordonating material

from their personal collections, or simply collecting specimens, and are much

acknowledged: Albert Allen, Moacyr Alvarenga, Gerardo Arriágada, Alberto

Ballerio, Ayr de Moura Bello, Gervásio Carvalho, Nicolas Degallier, Leonardo

Delgado, Ivone Diniz, Ana Aline Endres, François Feer, Antoine Foucart,



Eurides Furtado, Bruce Gill, Celso Godinho, Everardo and Paschoal Grossi,
Gonzalo Halffter, Malva I. M. Hernández, Henry Howden, Bert Kohlmann,
Wilson Werner Koller, Cristiano Lopes-Andrade, José Lopes, Sebastião
Lourenço de Assis Júnior, Júlio Louzada, Ângelo Machado, Carlos Augusto
Matrangolo, Marcelo Rocha Mattos, Claudia Medina, ísis M. Medri, MicheUe

Milhomem, Esteia Monteresino, José Luis Moreno, Miguel A. Morón, Sérgio
Roberto Rodrigues, Miguel Ángel Ruíz Díaz Villalba, Pamela Scheffler,
Gustavo Schiffler, Rodrigo Diniz Silveira, Angel Soh's, Carlos Sperber, Eduardo
Stehling, Daniela Maeda Takiya, Alexey Tishechkin, José R.Verdú, José Cola
Zanúncio.

Elliot Kitajima and Eduardo Alves taught me to use the Scanning
Electron Microscope that made possible illustration of most taxa. Cristiano

Lopes-Andrade did fírst photos and helped with most others. François Génier
did drawings in chapter 2and some useful photos of types; Violeta and Gonzalo
Halffter kindly gave me some ofthe photos inchapter 4.

Peter Hargreaves carefully read and improved English language in afírst
draft of chapter 1. Danielle Bastos Miranda and Enrico Bernard were also

victims in smaller similar situatíons. Teresa Telles Gonçalves, Jaqueline
Sicupira Rodrigues and Jorge Marcelo Padovani Porto were my victims in a
rather boringway of helping.

Many more people contributed in several other ways but there is not
enough space to list their names.

CAPES is acknowledged by the MS grant, and CNPq by expanding my
possibilities of working densely thanks to the new stereoscope. Universidade
Federal de Viçosa and Universidade Federal de Lavras are acknowledged by the
possibility of my formation as an entomologist.

And, last but not least, Pd like to thank to ali those (fortunately few)
people and institutions that discouraged me and/or refused helping or even



muddled me: without you, that dissertation would be much less careful and I

would have learned much less about human relations.



SUMMARY

Page

DISCLAIM i

RESUMO ü

ABSTRACT iü

CHAPTER 1 1

1 General Introduction 2

2 Theoretical Reference 4

2.1Taxonomic andSystematic history 4

2.2Present systematic status 13

2.3 Present taxonomic status 14

2.4 Present state of names 20

3 Bibliographical References 24

CHAPTER 2: A Review of Arrow*s Types of Trichillum and Pedaridium with

Description ofTwo New Species ofPedaridium 33

1 Resumo 34

2Abstract 35

3 Introduction 36

4 Material and Methods 37

5 Results and Discussion 38

5.1 New Species 50

6 Bibliographical References 57

CHAPTER 3: Lectotype Designations, New Synonymies and New Species in

the Genera Trichillum and Pedaridium 62

1 Resumo 63



2 Abstract ^4

3 Introduction 65

4 Material and Methods 66

5 Results and Discussion 67

5.1 New species 70

6 Bibliographical References 73

CHAPTER 4: A New Species of Pedaridium from México and Guatemala 77
1Resumo 70

2 Abstract 70

3 Introduction oq

4 Material and Methods gj

5 Results g2

6 Discussion g6

7 Bibliographical References g7

CHAPTER 5: A New Species ofPedaridium from Colômbia 90
1Resumo gi

2 Abstract Q2

3 Introduction 93

4 Material and Methods 94
5 Results 05

6 Discussion ng

7 Bibliographical References 99

CHAPTER 6: Phylogeny and Generic Rearrangement for the Group Trichillum-
Pedaridium 201

1Resumo 202

2 Abstract 203

3 Introduction 204

4 Materials and Methods 206



5 Results and Discussion 110

5.1 Characters used in cladistic analyses 110

5.2Results of cladisticanalyses 116*

5.3 New generical rearrangement fortheTrichillum-Pedaridium group 117

6 Bibliographical References 130

CHAPTER 7: Taxonomic Synopsys of theGroup Trichillum-Pedaridium 160

1 Resumo 161

2 Abstract 162

3 Introduction 163

4 Material and Methods 164

5 Results and Discussion 165

5.1 Key to the American Scarabaeidae genera and subgenera with fusionated

abdominal sternites and setae at elytral apex 165

5.2Besourenga Vaz-de-Mello, n. gen 168

5.2.1 Species included: 168
5.3Boreopedaridium Vaz-de-Mello, n. gen 174

5.3.1 Species included: 174
5.4 Boreotrichillum Vaz-de-Mello, n. gen 174

5.4.1 Species included: 174

5.5 Bradypodidium Vaz-de-Mello, n. gen 176

5.5.1 Species included: 176
5.6 Degallieridium Vaz-de-Mello, n. gen 179

5.6.1 Species included: 180

5.7 Eutrichillum Martínez, 1968, n. status 180

5.7.1 Species included: 180

5.8 Feeridium Vaz-de-Mello, n. gen 187

5.8.1 Species included: 187

5.9 Genieridium Vaz-de-Mello, n. gen 188



5.9.1 Key tothe species ofGenieridium Igg

5.9.2 Species included: Igg

5.10 Gillidium Vaz-de-Mello, n. gen 19g

5.10.1 Species included: 298

5.11 Horridotrichillum Vaz-de-Mello, n. gen 19g
5.11.1 Species included: 298

5.12 Howdenidium Vaz-de-Mello, n. gen 199
5.12.1 Species included: 199

5.13 Leotrichillum Vaz-de-Mello, n. gen 199

5.13.1 Key tothe species ofLeotrichillum 199

5.13.2 Species included: 200

5.14 Martinezidium Vaz-de-Mello, n. gen 201

5.14.1 Key tothe species ofMartinezidium 201

5.14.2 Species included: 202

5.15 Onoreidium Vaz-de-Mello, n. gen 205

5.15.1 Key tothe species ofOnoreidium 205
5.15.2 Species included: 206

5.16 Pedaridium Harold, 1868 208

5.16.1 Key tospecies ofPedaridium 208

5.16.2 Species included: 209

5.17 Pereiraidium Vaz-de-Mello, n. gen 211
5.17.1 Species included: 211

5.18 Silvia Vaz-de-Mello, n. gen 212
5.18.1 Species included: 212

5.19 Trichillidium Vaz-de-Mello, n. gen 212

5.19.1 Key tothe species of Trichillidium 212

5.19.2 Species included: 212

5.20 Trichillum (Paratrichillum) Vaz-de-Mello, n.subgen 215



5.20.1 Key tothespecies of Trichillum (Paratrichillum) 215

5.20.2 Species included: 215

5.21 Trichillum (Trichillum) Harold, 1868 217

5.21.1 Species included: 217

5.22 Youngidium Vaz-de-Mello, n. gen 240

5.22.1 Species included: 240

6 Bibliographical References 241

APPENDDC 264

omfad£Sorb'emS'""«2*25-**«*Pnnted in «he body
for r, pleaseread <?
forE,pÍeaseread 9



DISCLABVI

This dissertation does not intent to be a valid publication under articles

7-9 of the International Code of Zoological Nomenclature, so new zoological

names, synonyms and combinations published here are not valid or available

until their publication in other sources.



RESUMO

VAZDEMELLO, Fernando Zagury. Espécies originalmente nos gêneros
Trichillum Harold, 1868 e Pedaridium Harold, 1868(Coleoptera:
Scarabaeidae). 2003.266p. Dissertação (Mestrado emEntomologia)-
Universidade Federal de Lavras, Lavras.*

Este trabalho apresenta uma revisão dos Scarabaeidae (Coleoptera),
anteriormente agrupados nos gêneros Trichillum e Pedaridium. Esse grupo é
caracterizado pela distribuição neotropical, aliada à presença de pilosidade
dorsal, pelo menos no ápice elitral e fusão dos esternitos abdominais visíveis no
meio. Separa-se, à primeira vista, de Aphengium, que compartilha esses
caracteres, por carecer de carena lateral nos élitros. O grupo contava,
anteriormente ao presente estudo, com 26 espécies no gênero Pedaridium e 15
no gênero Trichillum, com dois subgêneros, além de duas espécies de gênero
incerto. Os capítulos 2 e 3 resultam de exame de tipos de espécies
duvidosamente identificadas e apresentam como resultado a descrição de cinco
novas espécies anteriormente identificadas erroneamente, e a sinonimização de
cinco espécies anteriormente descritas. Os capítulos 4 e 5 apresentam a
descrição de duas novas espécies morfológica e biogeograficamente muito
significativas. O capítulo 6 apresenta uma análise filogenética do grupo,
agrupando suas espécies em 20 gêneros (um com dois subgêneros), sendo cinco
baseados emespécies também novas (uma descrita nocapítulo 4). Finalmente, o
capítulo 7 apresenta uma sinopse taxonômica do grupo em nível específico,
resultando num total de 97 espécies, sendo 60 novas. Pela nova estruturação
genérica do grupo, as espécies estão agrupadas como segue (número total de
espécies, seguido pelo número de espécies descritas no presente trabalho, entre
parênteses após o nome do gênero ou subgênero): Besourenga (12, 9),
Boreopedaridium (1, 1), Boreotrichillum (1, 0); Bradypodidium (5, 2),
Degallieridium (1, 1), Eutrichillum (6, 3), Feeridium (1, 1), Genieridium (8, 2),
Gillidium (1, 1), Horridotrichillum (1, 0), Howdenidium (1, 0),Leotrichillum (2,
1), Martinezidium (7, 5), Onoreidium (4, 1), Pedaridium (2, 1), Pereiraidium (1,
0), Silvia (1, 1), Trichillidium (2, 0), Trichillum (Paratrichillum) (2, 0),
Trichillum (Trichillum) (37, 31), Youngidium (1, 0).

* Comitê Orientador: Dr. Júlio Neil Cassa Louzada - UFLA (Orientador), Dr.
Gonzalo Halffter Salas - BEcol, Dr. Sérgio Ide - IB-SP e Dr. Mario Zunino -
UniUrb.

li



ABSTRACT

VAZ DE MELLO, FernandoZagury. Species formerly in the genera
Trichillum Harold, 1868 and Pedaridium Harold, 1868(Coleoptera:
Scarabaeidae). 2003.266p.Dissertation (Master Program in Entomology)-
Universidade Federal de Lavras, Lavras.*

This work presents a revision of the Scarabaeidae (Coleoptera) formerly
grouped inthe genera Trichillum and Pedaridium. This group ischaracterized by
its Neotropical distribution, presence of dorsal setae at least in the elytral apex,
and visible abdominal stemites fusionated in the middle. It differs from
Aphengium, that have these same characters, by lacking a lateral elytral carina.
This group included, prior to the present study, 26 species in the genus
Pedaridium and 15 in the genus Trichillum, divided in two subgenera, and two
species of dubious generic placement. Chapters 2 and 3 present the results of
examining types of species with doubtfull identities, and result in thedescription
of five new species, and synonymization of five previously described species.
Chapters 4 and 5 present descriptions of two new species, of great
morphological or biogeographical significance. Chapter 6 presents phylogenetic
analyses of the group, grouping its included species in 20 genera (one with two
subgenera), five based on new species (one described in chapter 4). Finally,
chapter 7 presents a taxonomic synopsis of the group at specific levei, resulting
in 97 valid species, 60 being new. Following the new generical structuration of
thegroup, numbers of species are as follows (total number of species, followed
bynumber of species described in this work, are in brackets after each genus or
subgenus name): Besourenga (12, 9), Boreopedaridium (1, 1), Boreotrichillum
(1, 0); Bradypodidium (5, 2), Degallieridium (1, 1), Eutrichillum (6, 3),
Feeridium (1, 1), Genieridium (8, 2), Gillidium (1, 1), Horridotrichillum (1, 0),
Howdenidium (1,0), Leotrichillum (2, 1), Martinezidium (7, 5), Onoreidium (4,
1), Pedaridium (2, 1), Pereiraidium (1, 0), Silvia (1, 1), Trichillidium (2, 0),
Trichillum (Paratrichillum) (2, 0), Trichillum (Trichillum) (37, 31), Youngidium
(1,0).

* Guidance Committee: Júlio Neil Cassa Louzada - UFLA (Main Advisor),
Gonzalo Halffter Salas- IEcol, Sérgio Ide - IB-SPand Mario Zunino-UniUrb.

in



CHAPTER!



1 General Introduction

In the past few years agrowing interest in the systematics ofNeotropical
Scarabaeinae has prompted few workers to revise certain Neotropical genera as
this group of beeties is used in rapid assessment of biodiversity. More
specifically an effort has been done to revise poorly known groups such as
Trichillum and Pedaridium. For various reasons it was not possible for these
workers to study type material deposited inEuropean museums at the time. As

usual these revisions have instigated agreater interest in these groups and rurther
investigation has revealed that additional work is required as more systematic
problems have been found at the generic as well as specific levei in the genera.

The aim of this work is to redefine generic and specific entities in the
group Trichillum-Pedaridium, verifying its monophyly, solving systematic and
taxonomic problems, based primarily on phylogenetic inferences, using mainly
morphological characters and relating itto biogeographic patterns.

This work is divided in five main sections: Section 1 (Chapter 1)
summarizes current knowledge ofthe group. Section 2(Chapters 2and 3) results
from observation of primary types not examined by previous reviewers of the
group and designates necessary lectotypes, presents some new synonymies and
describes some new species that have been misidentified in previous works.
Section 3 (Chapters 4 and 5) describes new species that were under study by
other specialists when this project begun. Section 4 (Chapter 6) presents a
phylogenetic analysis of the group studied and its generic reestructuration,
describing as new type-species of those genera not represented by already
described species and a very unusual species belonging to one ofthe previously
known genera, whoose description is necessary to define generic lirnits. Finally,
Section 4 (Chapter 7) presents keys for generic and specific identifications and
reviews genera asdefined inChapter 6 inspecific levei.



The fírst section is co-authored by François Génier (Canadian Museum

of Nature, Ottawa, Canada), the second is separately co-authored by respectively

Gonzalo and Violeta Halffter (Instituto de Ecologia, Xalapa, México, Chapter 4)

and Bruce Gill (Agriculture Canada, Ottawa, Canada, Chapter 5) and the

remaining sections belong exclusively to the present author.



2 Theoretícal Reference

2.1Taxonomic and Systematic history

The genus Pedaria was erected by Laporte (1832), by a single species,
P. nigra, from Senegal. He stated that the genus could be distinguished from ali
other scarabaeids by its abdômen with only one segment (that is, fused

abdominal stemites). Later, the same author (as Castelnau, 1840) described two

other species of Pedaria also from África, he commented that the genus was
very close to Choeridium Serville, 1825 (now Ateuchus Weber, 1801), differing
by its elongated and convex body form, emarginated clypeus, elevated
pronotum, elytra covering completely the abdômen, "solded abdominal

segments" and absence of hind wings (!). He also commented they resemble
Aphodius üliger 1798 and both were endemic to África.

In 1859, Harold described the fírst non-African species of this genus,
Pedaria hirsuta, based ona number ofspecimens from Brazil, from his own and

StumVs collections. Harold (1875) tranferred the Australian species Aphodius
geminatus MacLeay 1871 to Pedaria.

In 1868, Harold erected one more scarabaeid genus, Trichillum,
including in it, T. heydeni Harold, 1868, based on specimens from Brazil, from

various collections. Li the same paper, Harold established the genus Pedaridium
for the American species of Pedaria, and differentiated itfrom Trichillum by the
shorter hind basitarsomere and unexpanded epipleuron. He did not designate a
type species for Pedaridium, however, as only Pedaria hirsuta was described

from the Américas, itcan be considered an implicit designation.

Gemminger &Harold (1869) cited Trichillum heydeni from Brazil. They
made, for the fírst time, the combination Pedaridium hirsutum, cited it from
Brazil to.



Borre (1880) described another species of Trichillum: T.

externepunctatum, based on a specimen from the Thomson collection labeled

"Colômbia", and "Uroxys hirtusn, the latter by Guérin-Ménéville, and cited

three more specimens of T. heydeni from Brazil, inCandèze collection.

Ohaus (1909) briefly commented on a new species of Trichillum, in

Ecuador, breeding on cow dung without building a nest. That was the first

citation of species inthe studied group for Ecuador, probably it refers to both T.

ohausi and T. cristatum, described by Arrow in 1931. That was also the first

comment onbreeding without nesting in true dung beeties (Scarabaeidae s. str.).

This observation wascitedalsoby Halffter & Matthews (1966).

Gillet (1911) cited Pedaridium hirsutum, Trichillum heydeni and T.

externepunctatum in his catalogue. He also cited for the first time T. heydeni for

Argentina. He followed Harold (1875) and considered A. geminatus in Pedaria.

Arrow(1913) described three more species of Pedaridium: P. cryptops,

from Brazil, based on three specimens, one from Jatai, Goiás, and the others

from Natal, Rio Grande do Norte; P.fulgens, basedon two specimens from Rio

Salado, Gran Chaco, Argentina; and P. argentinum, based on an unspecified

number of specimens from Icano and Rio Salado, Gran Chaco, Argentina. This

last species has been erroneously called P. rugiceps in the key given just after

the description of P. cryptops. In the comments under P. argentinum, the author

compared it with Pedaridium setosum. This last named has never been formally

described and shouldbe considered in litteris or lapsus for P. hirsutum (Harold,

1859).

Lea (1923) described another Australian species of Pedaria, P.

alternata.

Boucomont (1928) described Trichillum hirsutum based on a single

specimen from São Paulo, Brazil, and Trichillum bradyporum from a specimen



taken in 1925 by Nevermann in the ânus of a sloth in Santa Clara Province,
Costa Rica.

Arrow, in 1931, revised the genus Trichillum and described three new

species: T. hystrix, based on specimens from Argentina (La Noria, Rio San

Javier, Santa Fé); T. ohausi and T. cristatum, both from Ecuador, the former

from Loja, Calvário and Piscobamba, and the later from Loja and Piscobamba.
In this paper, the author did not mention the species described in 1928 by
Boucomont.

The same author, one year later (Arrow, 1932) described two more

species of Pedaridium: P. paranense, based on four specimens from Castro,
Paraná, Brazil, and P. quadridens, based on asingle specimen from Santa Elena,
Entre Rios, Argentina. He also gave an updated key for the genus Pedaridium
and comments about Boucomonfs (1928) descriptions ofTrichillum, adding that
T. hirsutum wouldbe closeto T. hystrix Arrow, 1931.

Saylor (1935) described three more species of Trichillum, ali based on

single specimens and ali from Horquetá, Paraguay. The described species were
T. minutum, T. arrowi and T. boucomonti, the second was related to T. hystrix
and the last to T. ohausi.

Carter (1936) described athird Australian species of Pedaria.

Paulian (1936) described the homonymous T. arrowi based on asingle
specimen from Pará, Brazil, giving a new key for the species known in this

genus, except for that for Saylor (1935), whose work was surely unknown to

him. The author cited once more T. heydeni Harold, 1868 as occuning in
Argentina.

Balthasar (1938) described Pedaridium bidens based on nine specimens
from Jatai, Goiás, Brazil, and Horquetá, Paraguay. In the same paper, the author
presented a key for ali species of Pedaridium known at that time.



The same author (Balthasar, 1939) reviewed the genus Trichillum and

described two new species, T. elongatum and T. vejdovskyi, the former from

Argentina, Cordoba, Rosário, and the later from Bolivia, Santa Cruz,

Buenavista, based on a single specimen. In the same paper the author cited T.

heydeni from Porto Alegre, Rio Grande do Sul, Brazil, T. externepunctatum

from Paraguay, T. boucomonti from Buenos Aires, Argentina, T. hirsutum from

Tucumán, Argentina, T. hystrix from Paraguay, and gave a new name for T.

arrowi Paulian: T. pauliani.

Pessoa & Lane (1941) cited P. hirsutum from Franca, São Paulo state,

Brazil, and T heydeni fromVila Nova, Bahia, Brazil. They also stated that there

was a common species in São Paulo dobtfully identifiedas T. externepunctatum.

Balthasar, in 1942, described Trichillum depilatum based on a single

specimen from São Paulo, and related it to T. externepunctatum.

Blackwelder (1944) cited in his catalog ali species known at that time of

Pedaridium, and ali Trichillum except those described by Balthasar (1939,

1942). Healso gave a new name, T. homonymum, for T. arrowiPaulian, 1936.

Pereira (1946) described a horned Pedaridium, P. almeidai, based on

specimens from Piraquara, Guaraúna and Itupava, Paraná, and Glória, Rio

Grande do Sul, Brazil.

Martínez (1947), in his addenda y corrigenda to Blackwelder (1944)

listing, added species described by Balthasar (1939) and Pereira (1946), and

synonymized Trichillum homonymum Blackwelder, 1944 with T. pauliani

Balthasar, 1939.

Robinson (1948) described Trichillum pilosus based on two specimens

from Barro Colorado Island, Panamá, and compared it with T. bradyporum

Boucomont, 1928.

Martínez (1951) described Pedaridium mansosotoi from Paraguay and

Argentina, andcompared it with P. bidens and P. cryptops.



In addition, Martínez (1959) cited in his catalogue of Argentinean

scarabaeids: Pedaridium argentinum, P. fulgens, P. mansosotoi, P. quadridens,
Trichillum elongatum, T. externepunctatum, T. heydeni and T. hystrix. He still

cited T. externepunctatum from Brazil, Bolivia, Uruguay, Paraguay and
Colômbia, and hypothesized that T. hystrix and T. boucomonti could be

synonyms.

In another paper in the same year, Pereira & Martínez (1959)
redescribed Trichillum arrowi Saylor, 1935, comparing it to T.
externepunctatum Borre, 1880 and T. depilatum Balthasar, 1942, based on

specimens from Boquerón, Paraguay.

Vulcano & Pereira (1967) included both Pedaridium and Trichillum

among the Scarabaeidae occuring in Amazonian Region. They cited the only
described Pedaridium species that occurs there, P. cryptops Arrow, 1913, and

gave a key to Trichillum species in this region, including T. externepunctatum,
T. cristatum, T. ohausi, T. vejdovskyi and T. pauliani.

Martínez (1968) erected the subgenus Eutrichillum for Trichillum,
including in it T. boucomonti (type species), T. minutum, T. vejdovskyi, and,

based on descriptions, T. hystrix and T. pauliani. He also synonymized (again,
done before in 1947 byhimself) T. homonymum with T. pauliani, hypothesized
that T. pauliani also could be (with T. hystrix) a synonym of T. boucomonti. He

also transferred T. bradyporum, T. ohausi and T. elongatum to Pedaridium,
hypothesized T. hirsutum could also be a Pedaridium and in this case would

need a new name, and considered the last and T. cristatum as incertae sedis. In

the same paper the author reviewed the subgenus Trichillum, describing the
following new species: T. (T.) adjunctum, from São Paulo and Paraná, Brazil, T.

(T.) pereirai from São Paulo and Minas Gerais, Brazil, T. (T.) halffteri from

Misiones, Argentina, and Santa Catarina and Rio Grande do Sul, Brazil, and T.

(subgenus?) horacioi, from Nueva Moka, Sara, Santa Cruz, Bolivia. The



following species were considered in the subgenus Trichillum: T. heydeni, T.

externepunctatum, T. depilatum and 7". arrowi. The new citations included T.

heydeni for Córdoba, Argentina, T. externepunctatum from Brazil, Bolivia,

Paraguay and Argentina, and T. depilatum from Córdoba, Argentina. In this

paper, the author omitted anycomments about T. pilosus Robinson, 1948.

Martínez (1974) described "Pedaridium (?) caingua" based on three

females from Argentina, and stated that this new species, together with P.

quadridens, should forma new taxon under studyby him.

In 1976, Matthews erected the new genus Notopedaria for Australian

species of Pedaria.

Bacchus (1978) listed Arrow's syntypes and holotypes of his described

Trichillum and Pedaridium, but did not designate lectotypes among this

material, as he considered necessary to dissect them to differentiate males and

females. In his list, he called sometimes Pedaridium either as Pedarium or

Pedarius. He considered seven syntypes for P. argentinum, three for T.

cristatum, three for P. cryptops (pointing out that there was a fourth specimen

missing - andthis was not corroborated by the original description), eight for T.

hystrix, five for T. ohausi, four for P. paranense, and the holotype for P.

quadridens.

Ratcliffe (1980) described adults, larvae and pupae of a species of

Trichillum, T. adisi, taken from the pellage óf three-toed sloths in Brazilian

Amazônia. Healso gave both keys for subgenera of Trichillum and for species in

the subgenus Eutrichillum, including T. hirsutum Boucomont, 1928, and stating

that it has elytral intervals uniseriately punctate, instead of biseriately punctate

as stated by the author of this last species. The same author (Ratcliffe, 1981)

redescribed the type of Trichillum hirsutum. In this paper, he also cited both T.

hirsutum and T. boucomonti as occurringin Manaus, Amazonas, Brazil.



Howden & Young (1981) described two species of Pedaridium from

Panamá, P. bottimeri and P. brevisetosum, both based on unique specimens from

Barro Colorado Island. They also assigned Trichillum pilosus Robinson, 1948 to

Pedaridium, based on characters used by Paulian (1936) and Martínez (1968) to

distinguish these genera, cited P. pilosum from Costa Rica and Ecuador, and

commented on the confusion regarding distinguishing characters between

Trichillum and Pedaridium, and the need of comprehensive revisions of both.

They also stated the possibility of occurrence of Pedaridium bradyporum
(Boucomont, 1928) in Panamá, comparing it to P. pilosum.

In 1986, Matthews & Stebnicka synonymized Notopedaria with

Demarziella Balthasar, 1961, considered since its description as belonging tothe

Aphodiidae, and forming the monobasic tribe Demarziellini (Matthews &

Stebnicka, 1986).

Martínez (1987), in his catalog of scarabaeid species occurring in Salta,

cited from there Pedaridium argentinum, P. quadridens, Trichillum

externepunctatum, T. boucomonti; once more stated that he found no differences

between this later species and Trichillum hystrix, and synonymized Trichillum
elongatum Balthasar, 1939 withPedaridium argentinum.

D'Hotman & Scholtz (1990) included Pedaridium among examined

genera in a paper dealing with male genitalia in the Scarabaeoidea, but did not

illustrate or comment on the genus specifically. The same occurred with other

papers dealing with general Scarabaeoidea systematics (e.g. Browne & Scholtz

1995, 1998, 1999). Nel &Scholtz (1990) in apaper on adult scarab mouthparts
citedPedaria pilosaRobinson (sic) among examined material.

Martínez (1992) described a Venezuelan species, P. bordoni, based on

specimens from Guárico, Barinas, Monagas and Anzoátegui states. In the

abstract of this paper, in both Spanish and English, the author cited Pedaridium

maya, an undescribed species, from México.

10



Ferreira & Galileo (1993) revised the genus Pedaridium, excepting for

P. bordoni Martínez, 1992, that was probably not known to them. In this

revision, they transferred T. adisi to Pedaridium, misspelling it P. adissi, and

synonymized Pedaridium rugiceps Arrow, 1913 objectively under P.

argentinum Arrow, 1913. Six species were described in this revision: P.

equatoriensis and P. howdeni, from Ecuador, P. brasiliensis, from Brazil, P.

venezuelensis, from Venezuela, and P. martinsi and P. martinezi, both from

Argentina. They apparentiy erroneously considered Pedaridium male in gender,

and not neuter as stated by the suffix "idium", as they modified some species

names and used the malegender for the new ones. P. bottimeri wascited for the

first time from Venezuela, P. mansosotoi from Brazil, P. quadridens from

Boliviaand Paraguay, and P. pilosum from Bolivia. It is important to point out

that the authors did not exam most types, which arein European museums.

Monteresino et ai. (1996) citedP. argentinum, P.fulgens, P. quadridens,

Trichillum depilatum, T. externepunctatum, T. heydeni and T. boucomonti from

Córdoba, Argentina. They apparentiy did not know the Ferreira & Galileo

(1993) revision, asother species were cited from Córdoba in this last.

Verdú & Galante (1997) described Trichillum morellii, from Uruguay,

comparing it to T. externepunctatum, and gave a new key for species in the

subgenus Trichillum.

Montreuil (1998), in a paper on phylogenetic relations between

Ateuchini and Coprini, examined Pedaridium (the type species and, possibly,

some additional species not stated), and placed it in an unsolved clade including

Demarziella, Pedaria, Sarophorus Erichson, 1847, and the clade Bdelyrus

Harold, 1869 + Bdelyropsis Pereira, Vulcano & Martínez, 1960. This paper did

not included Trichillum.

11



Vkwfe<'1rtQ^

Vaz-de-Mello & Canhedo (1998) described Pedaridium louzadaorum

and P. zanunciorum, both from Minas Gerais state, Brazil, and included those

and P. bordoni Martínez, 1992 in the Ferreira &Galileo (1993) key for species.
Escobar (2000) cited a Pedaridium species close to P. pilosum from the

Cinturón Árido Pericaribeno, Trichillum hystrix from Amazônia, and T.
externepunctatum with no locality, from Colômbia.

Vaz-de-Mello (2000) listed for Brazil, ali Pedaridium species cited from

that country by Ferreira &Galileo (1993), adding the species described by Vaz-
de-Mello &Canhedo (1998), and ali Trichillum previously cited from Brazil.

In 2001, Aguilar described Pedaridium amarillai, based on a single
female specimen from Paraguay. The author did not cite the revision by Ferreira
& Galileo (1993), and stated that the new species had sinuate epipleura, a
character that would place the species in Trichillum according to Ferreira &
Galileo (1993).

Verdú & Galante (2001) described larvae and breeding behaviour of
Pedaridium brasiliense andP. almeidai. They also reexamined larvae ofP. adisi

and concluded the genus seems to be polyphyletic based on larval morphological
characters. A short comment was made on Trichillum externepunctatum
corroborating Ohaus' (1909) and their own data on Pedaridium, that those

genera do not present nesting behaviour.

Barbero (2001) cited for the first time Pedaridium pilosum from

Nicarágua, establishing the depaitment of Rio San Juan as the northern limit for
that genus.

Medina et ai. (2001) cited Pedaridium pilosum, Trichillum
externepunctatum, T. hystrix and T. hirsutum from Colômbia.

Génier &Vaz-de-Mello (2002) diagnosed and designated lectotypes for
those species described by Arrow in bot Trichillum and Pedaridium,
synonymized P. equatoriensis under T. cristatum, transferring that last to

12



Pedaridium, P. martinezi under P. fulgens, P. mansosotoi under P. cryptops,

observed gender consideration mistakes made by Ferreira & Galileo (1993) for

Pedaridium, and described P. margareteae for the species erroneusly identified

as P. cryptops, and P. galileoae for the species erroneusly identified as P.

fulgens in that last work.

Solís & Kohlmann (2003) described Trichillum (Eutrichillum) arcus

from Costa Rica, relating it to T. boucomonti, and illustratedand commented the

distribution of Pedaridium bradyporum, cited that speciesfor the first time from

the Pacific slope in Costa Rica.

Vaz-de-Mello, Halffter & Halffter (in press) described Pedaridium maya

from Southern México and Guatemala.

Gill & Vaz-de-Mello (in press) described Pedaridium medinae, from

Cundinamarca, Risaralda and Quindio in Colômbia.

22 Present systematic status

Since the beginning, both genera have been since the beginning placed

near Ateuchus Weber, 1801, and tiras are considered actually belonging to the

tribe Ateuchini (subtribe Ateuchina of some authors). Morphological differences

between Pedaridium and Pedaria have never been pointed out in literature, and

need to be exposed, since the only commented difference between them is

geographical.

Morphological differences between Pedaridium and Trichillum havenot

been clearly stated. Harold (1868) distinguished them in his key by shape of

epipleuron and hind tarsi. In later keys (e.g. Luederwaldt, 1931, Paulian, 1936,

Pessoa & Lane, 1941, Pereira, 1954, Vulcano & Pereira, 1967) the only noted

difference is in hind tarsi. However, P. almeidai Pereira, 1946 shows Trichillum-

like hind tarsi (i.e., basitarsomere longer than second tarsomere). The genus

13



Aphengium Harold, 1868 is commonly placed near Trichillum and Pedaridium

in keyscited above, sharing with these genera the fused abdominal stemites and

pilose elytra, being separated from Pedaridium generally by longer hind
basitarsus and from Trichillum by shape of pygidium. However, other characters

such as meso- and metasternal shape, and genitalia, appear to put it closer to

Ateuchus than to the Trichillum-Pedaridium complex (personal observation).

Martínez (1968) wrote short comments on this, transferring some
species from Trichillum to Pedaridium, but did not clearly point out differences

between them. In addition, Martínez (1974) put doubts on the generic
classification of the group Trichillum-Pedaridium in his description of P. (?)
caingua.

Ferreira & Galileo (1993) used only the epipleuron to stablish generic

differences, transferring some more species from Trichillum to Pedaridium

based on this, but did not comment on characters other than tarsi, that was

considered useless. The monophyly of both genera have never been commented

on, and most characters vary considerably among species in both genera, giving
rise to the consideration that this classification, based solely on one character
(shape of epipleuron), is notnow a good one.

Larval characters did not appear to support Pedaridium as a

monophyletic group (Verdú & Galante, 2001).

23 Present taxonomic status

Up to now, the genus Pedaridium contains 29 valid species, four names

that are synonyms of valid species, one nomen nudum (that can be simply a
lapsus), and one misspelling of species name. The genus name has been

misspelled twice, either as Pedarium or Pedarius. Some species names have

14



been described as male nouns, however Pedaridium is clearly a neutral one and

those names are in need of correction.

The genus Trichillum hasno synonyms and includes two subgenera. The

nominotypical one contains eight valid species names, and no synonyms at

species levei. The other, Trichillum (Eutrichillum), contains seven valid species

names, and two synonyms at species levei, one of which is a sênior synonym,

homonym of aname previously in Trichillum (Trichillum). One species names is

considered as incertae sedis, without subgeneric placement.

The following lines summarize the current understanding of the group,

listing modifications that occurred in status, combination and spelling of names.

Genus 1. Pedaridium Harold, 1868

Pedarius- misspeUingin Bacchus, 1978

Pedarium - misspelling in Bacchus, 1978

Species 1.1. Pedaridium hirsutum (Harold, 1859)

Pedaria hirsuta Harold, 1859

Pedaridium hirsutum - comb. in Gemminger & Harold (1869)

Wedaridium setosum - Arrow, 1913 (lapsusT)

Species 1.2. Pedaridium cryptops Arrow, 1913

Pedarium cryptopsArrow, 1913 - misspel. in Bacchus(1978)

Pedaridium mansosotoi Martínez, 1951 - syn. in Génier & Vaz-

de-Mello (2002)

Species 1.3. Pedaridium fulgens Arrow, 1913

Pedarius fulgens Arrow 1932 - misspel. in Bacchus (1978)

Species 1.4. Pedaridium argentinum Arrow, 1913

Pedaridium rugiceps Arrow, 1913 (lapsus) - syn. in Ferreira &

Galileo (1993)

Trichillum elongatum Balthasar, 1939 - syn. in Martínez (1987)

15



Pedaridium elongatum (Balthasar, 1939) - comb. in Martmez
(1968)

Pedarium argentinum Arrow, 1913 - misspel. in Bacchus
(1978)

Species 1.5. Pedaridium bradyporum (Boucomont, 1928)

Trichillum bradyporum Boucomont, 1928

Pedaridium bradyporum (Boucomont, 1928) - comb. in
Martínez (1968)

Species 1.6. Pedaridium ohausi (Arrow, 1931)

Trichillum ohausi Arrow, 1931

Pedaridium ohausi (Arrow, 1931) - comb. in Martínez (1968)
Species 1.7. Pedaridium cristatum (Arrow, 1931)

Trichillum cristatum Arrow, 1931

Pedaridium cristatum (Arrow, 1931) - comb. in Génier & Vaz-

de-Mello (2002)

Pedaridium equatoriensis Ferreira & Galileo, 1993 - syn. in
Génier& Vaz-de-Mello (2002)

Species 1.8. Pedaridium paranense Arrow, 1932

Pedaridium paranensis Arrow, 1932 - misspel. in Ferreira &
Galileo (1993)

Species 1.9. Pedaridium quadridens Arrow, 1932

Species 1.10. Pedaridium bidens Balthasar, 1938

Species 1.11. Pedaridium almeidai Pereira, 1946

Species 1.12. Pedaridium pilosum (Robinson, 1948)

Trichillum pilosus Robinson, 1948

Pedaridium pilosum (Robinson, 1948) - comb. in Howden &
Young(1981)

16



Pedaria pilosa (Robinson, 1948) - miscombination in Nel &

Scholtz (1990)

Species 1.13. Pedaridium caingua Martínez, 1974

Pedaridium (?) cainguaMartínez, 1974

Pedaridium caingua Martínez, 1974 - Ferreira & Galileo (1993)

Species 1.14.Pedaridium adisi (Ratcliffe, 1980)

Trichillum (Eutrichillum) adisi Ratcliffe, 1980

Pedaridium adissi (Ratcliffe, 1980) - comb7misspel. in Ferreira

& Galileo (1993)

Species 1.15. Pedaridium brevisetosum Howden & Young, 1981

Species 1.16. Pedaridium bottimeri Howden & Young, 1981

Species 1.17. Pedaridium bordoni Martínez, 1992

Species 1.18. Pedaridium howdeni Ferreira & Galileo, 1993

Species 1.19. Pedaridium brasiliensis Ferreira & Galileo, 1993

Species 1.20. Pedaridium venezuelensis Ferreira & Galileo, 1993

Species 1.21. Pedaridium martinsi Ferreira & Galileo, 1993

Species 1.22. Pedaridium martinezi Ferreira & Galileo, 1993

Species 1.23. Pedaridium louzadaorum Vaz-de-Mello & Canhedo, 1998

Species 1.24. Pedaridium zanunciorum Vaz-de-Mello & Canhedo, 1998

Species 1.25. Pedaridium amarillai Aguilar, 2001

Species 1.26. Pedaridium margareteae Génier &Vaz-de-Mello, 2002

Species 1.27. Pedaridium galileoae Génier &Vaz-de-Mello, 2002

Species 1.28. Pedaridium maya Vaz-de-Mello, Halffter & Halffter (in

press)

Pedaridium maya Martínez, 1992 - nomen nudum

Species 1.29. Pedaridium medinae Gill&Vaz-de-Mello (in press)

17



Nomen nudum 1.1. Pedaridium setosum Arrow, 1913, seeremarks under

P. hirsutum (Harold, 1859)

Genus 2. Trichillum Harold, 1868

Subgenus 1. Trichillum (s. str.) - defined byMartínez (1967)

Species 2.1. Trichillum (Trichillum) heydeni Harold, 1868

Trichillum heydeni Harold, 1868

Trichillum (Trichillum) heydeni Harold, 1868 - comb. in

Martínez (1968)

Species 2.2. Trichillum (Trichillum) externepunctatum Borre, 1880

Uroxys hirta - nomen in litteris attributed to Guérin-Ménéville

by Borre (1880)

Trichillum externepunctatum Borre, 1880

Trichillum (Trichillum) externepunctatum Borre, 1880 - comb.

in Martínez (1968)

Species 2.3. Trichillum (Trichillum) arrowi Saylor, 1935

Trichillum arrowi Saylor, 1935 (not Paulian, 1936)

Trichillum (Trichillum) arrowi Saylor, 1935 - comb. in Martínez

(1968)

Species 2.4. Trichillum (Trichillum) depilatum Balthasar, 1942

Trichillum depilatum Balthasar, 1942

Trichillum (Trichillum) depilatum Balthasar, 1942 - comb. in

Martínez (1968)

Species 2.5. Trichillum (Trichillum) adjunctum Martínez, 1968

Species 2.6. Trichillum (Trichillum) pereirai Martínez, 1968

Species 2.7. Trichillum (Trichillum) halffteri Martínez, 1968

Species 2.8. Trichillum (Trichillum) moiellii Verdú &Galante, 1997
Subgenus 2. Trichillum (Eutrichillum) Martínez, 1968

Species 2.9. Trichillum (Eutrichillum) hirsutum Boucomont, 1928

18



Trichillum hirsutum Boucomont, 1928 - as incerta sedis in

Martínez (1968)

Trichillum (Eutrichillum) hirsutum Boucomont, 1928 - comb. in

Ratcliffe (1980)

Species 2.10. Trichillum (Eutrichillum) hystrix Arrow, 1931

Trichillum hystrix Arrow, 1931

Trichillum (Eutrichillum) hystrix Arrow, 1931 - comb. in

Martínez (1968)

Species 2.11. Trichillum (Eutrichillum) boucomonti Saylor, 1935

Trichillum boucomonti Saylor, 1935

Trichillum (Eutrichillum) boucomonti Saylor, 1935 - comb. in

Martínez (1968) (type species)

Species 2.12. Trichillum (Eutrichillum) minutum Saylor, 1935

Trichillum minutum Saylor, 1935

Trichillum (Eutrichillum) minutum Saylor, 1935 - comb. in

Martínez (1968)

Species 2.13. Trichillum (Eutrichillum) pauliani Balthasar, 1939

Trichillum arrowi Paulian, 1936 (not Saylor, 1935)

Trichillum pauliani Balthasar, 1939- nomennovum

Trichillum homonymum Blackwelder, 1944 - nomen novum -

syn. in Martmez (1947)

Trichillum (Eutrichillum) pauliani Balthasar, 1939 - comb. in

Martmez (1968)

Species 2.14. Trichillum (Eutrichillum) vejdovskyi Balthasar, 1939

Trichillum vejdovskyi Balthasar, 1939

Trichillum (Eutrichillum) vejdovskyi Balthasar, 1939 - comb. in

Martínez (1968)

Species 2.15. Trichillum (Eutrichillum) arcus Solís & Kohlmann, 2003

19



Incertae sedis (stated byMartínez, 1968):

Species 2.16. Trichillum (?) horacioi Martmez, 1968

2.4 Present state of names

(species epithets with original genus in parenthesis):

1. adisi Ratcliffe, 1980 (Trichillum) - valid, now in Pedaridium (Ferreira &
Galileo, 1993)

2. adissi (Ratcliffe, 1980) (Pedaridium) - misspelling of P. adisi (Ratcliffe,
1980) in Ferreira & Galileo (1993)

3. adjunctum Martínez, 1968 (Trichillum (Trichillum)) - valid
4. almeidai Pereira, 1946 (Pedaridium) - valid

5. amarillai Aguilar, 2001 (Pedaridium) - valid

6. arcus Solís &Kohlmann, 2003 (TrichillumfEutrichillum)) - valid
7. argentinum Arrow, 1913 (Pedaridium) - valid

8. arrowi Paulian, 1936 (not, Saylor 1935) (Trichillum) - sênior synonym of
Trichillum (Eutrichillum) pauliani Balthasar, 1939 (Balthasar, 1939)

9. arrowi Saylor, 1935 (Trichillum) - valid, now in Trichillum (Trichillum)
(Martínez, 1968)

10. bidens Balthasar, 1938 (Pedaridium) - valid

11. bordoni Martínez, 1992 (Pedaridium) - valid

12. bottimeri Howden &Young, 1981 (Pedaridium) - valid

13. boucomonti Saylor, 1935 (Trichillum) - valid, now in Trichillum
(Eutrichillum) (type species) (Martínez, 1968)

14. bradyporum Boucomont, 1928 (Trichillum) - valid, now in Pedaridium
(Martínez, 1968)

20



15. brasiliensis Ferreira & Galileo, 1993 (Pedaridium) - valid, but misspelled -

would be P. brasiliense

16. brevisetosum Howden & Young, 1981 (Pedaridium) - valid

17. caingua Martínez, 1974 (Pedaridium*!) - valid, as Pedaridium (Ferreira &

Galileo, 1993)

18. cristatum Arrow, 1931 (Trichillum) - valid, now in Pedaridium (Génier &

Vaz-de-Mello, 2002)

19. cryptops Arrow, 1913 (Pedaridium) - valid

20. depilatum Balthasar, 1942 (Trichillum) - valid, now in Trichillum

(Trichillum) (Martínez, 1968)

21. elongatum Balthasar, 1939 (Trichillum) - synonym of Pedaridium

argentinum Arrow, 1913 (Martínez, 1987)

22. equatoriensis Ferreira & Galileo, 1993 (Pedaridium) - misspelled - would

be P. equatoriense, synonym of P. cristatum (Arrow, 1931) (Génier &Vaz-

de-Mello)

23. Eutrichillum Martínez, 1968(subgenus of Trichillum) - valid

24. externepunctatum Borre, 1880 (Trichillum) - valid, now in Trichillum

(Trichillum) (Martínez, 1968)

25. fulgens Arrow, 1913 (Pedaridium) - valid

26. galileoae Génier &Vaz-de-Mello, 2002 (Pedaridium) - valid

27. halffteri Martínez, 1968 (Trichillum (Trichillum)) - valid

28. heydeni Harold, 1868 (Trichillum) - valid, now in Trichillum (Trichillum)

(type species) (Martínez, 1968)

29. hirsuta Harold, 1859 (Pedaria) - valid, now Pedaridium hirsutum

(Gemminger & Harold, 1869)

30. hirsutum Boucomont, 1928 (Trichillum) - valid, now in Trichillum

(Eutrichillum) (Ratcliffe, 1980)

21



31. hirta Guérin-MénéviUe, in litteris (Uroxys) - synonym of Trichillum
(Trichillum) externepunctatum Borre, 1880 (cited by Borre, 1880)

32. homonymum Blackwelder, 1944 (Trichillum) - júnior synonym of
Trichillum (Eutrichillum) pauliani Balthasar, 1939 (Martínez, 1947,1968)

33. horacioi Martínez, 1968 (Trichillum) - valid, incerta sedis (Martmez, 1968)
34. howdeni Ferreira &Galileo, 1993 (Pedaridium) - valid

35. hystrix Arrow, 1931 (Trichillum) - valid, now in Trichillum (Eutrichillum)
(Martínez, 1968)

36. louzadaorum Vaz-de-Mello &Canhedo, 1998 (Pedaridium) - valid
37. mansosotoi Martínez, 1951 (Pedaridium) - synonym of P. cryptops Arrow,

1913 (Génier &Vaz-de-Mello, 2002)

38. margareteae Génier &Vaz-de-Mello, 2002 (Pedaridium) - valid
39. martinezi Ferreira &Galileo, 1993 (Pedaridium) - synonym of P. fulgens

Arrow, 1913 (Génier &Vaz-de-Mello, 2002)

40. martinsi Ferreira &Galileo, 1993 (Pedaridium) - valid

41. maya Martínez, 1992 (Pedaridium) - nomen nudum, synonym of P. maya
Vaz-de-Mello, Halffter &Halffter (in press)

42. maya Vaz-de-Mello, Halffter &Halffter (in press) (Pedaridium) - valid
43. medinae Gill &Vaz-de-Mello (in press) (Pedaridium) - valid

44. minutum Saylor, 1935 (Trichillum) - valid, now in Trichillum (Eutrichillum)
(Martínez, 1968)

45. morellii Verdú &Galante, 1997 (Trichillum (Trichillum)) - valid
46. ohausi Arrow, 1931 (Trichillum) - valid, now in Pedaridium (Martínez,

1968)

47. paranense Arrow, 1932 (Pedaridium) - valid

48. paranensis Arrow, 1932 (Pedaridium) - misspeUing ofP. paranense Arrow,
1932 in Ferreira & Galileo (1993)

22



49. pauliani Balthasar, 1939 (Trichillum) - valid, now in Trichillum

(Eutrichillum) (Martínez, 1968)

50. Pedaridium Harold, 1868 - valid

51. Pedarium - misspelling of Pedaridium in Bacchus (1978)

52. Pedarius - misspelling of Pedaridium in Bacchus (1978)

53. pereirai Martínez 1968 (Trichillum (Trichillum)) - valid

54. pilosum (Robinson 1948) (Pedaridium) - correct spelling and new

combination for Trichillum pilosus Robinson, 1948 (Howden & Young,

1981)

55. pilosus Robinson, 1948 (Trichillum) - valid, now inPedaridium (Howden &

Young, 1981), misspelled, shouldbe T. pilosum.

56. quadridens Arrow, 1932 (Pedaridium) - valid

57. rugiceps Arrow, 1913 (Pedaridium) - sênior objective synonym of

Pedaridium argentinum Arrow, 1913 (Ferreira &Galileo, 1993)

58. setosum Arrow, 1913 (Pedaridium) - nomen nudum or lapsus for P.

hirsutum (Harold 1859)

59. Trichillum Harold, 1869 - valid

60. vejdovskyi Balthasar, 1939 (Trichillum) - valid, now in Trichillum

(Eutrichillum) (Martínez, 1968)

61. venezuelensisFerreira & Galileo, 1993 (Pedaridium) - valid, but mispelled -

would be P. venezuelense

62. zanunciorumVaz-de-Mello & Canhedo, 1998 (Pedaridium) - valid

23



3 Bibliographical References

AGUILAR JÚLIO, C. A. Una nueva espécie de Pedaridium (Coleoptera:
Scarabaeidae: Scarabaeinae) proveniente dei Parque Nacional Cerro Cora.
Boletín dei Museo Nacional de Historia Natural dei Paraguay, San Lorenzo,
v. 13, p. 1-4,2001.

ARROW, G. J. Some new species ofLamellicom beeties from Brazil. The

Annals and Magazine ofNatural Hístory, London, v. 11, S. 8, p. 456-466,
1913.

ARROW, G. J. The coleopterous genus Trichillum (Copridae), with akey to the
species. The Annals and Magazine of Natural Hístory, London, v. 8, S. 10, p.
609-611,1931.

ARROW, G. J. New species of Lamellicom beeties (subfamily Coprinae) from
South America. Stylops, London, v. 1, n. 10, p. 223-226,1932.

BACCHUS, M. E. Acatalogue ofthe type-specimens ofthe Scarabaeinae
(Scarabaeidae) and the smaller Lamellicom families (Coleoptera) described by
G. J. Arrow. BuUetín of the British Museum (Natural Hístory), Entomology
series, London, v. 37,n. 3, p. 97-115,1978.

BALTHASAR, V. Neue Gattungen und Arten der südamerikanischen

Coprophagen (4. Beitrag zur Kenntnis der Scarabeiden der neotropischen
Region). Entomologische Blãtter, Krefeld, v. 34, n. 4, p. 210-223,1938.

24



BALTHASAR, V. Monographieder GattungTrichillum (62. Beitrag zur

Kenntnis der Scarabeiden, Col.). Vestnik Ceskoslovenské Spolecnosti

Zoologické,Praze, v. 6-7, p. 11-26,1939.

BALTHASAR, V. Neue Scarabeiden aus Süd-Amerika. 72. Beitrag zur

Kenntnis der Scarabeiden(Col.). Casopis Ceskoslovenské Spolecnosti

Entomologícké, Praze, v. 39, p. 36-44,1942.

BARBERO, E. Scarabaeidae (Coleoptera) copronecrófagos interesantes dei

Departamento deRio San Juan, Nicarágua. Revista Nicaraguense de

Entomología, León, v. 55/58, p. 11-21,2001.

BLACKWELDER, R. E. Checklistof the coleopterous insects of México,

CentralAmerica, the West Indies, and South America.Part 2. Bulletin of the

United States National Museum, Washington, n. 185, p. 189-341,1944.

BORRE, F. P. C. A. P. de. Description d'une espèce nouvelle du genre

Trichillum Harold (Coprides; Choeridiides). CompteRendue de Ia Scéance de

IaSociété Entomologíque de Belgique (Annales de IaSodété Entomologíque

de Belgique), Bruxelles,v. 23, p. xxvii-xxviii, 1880.

BOUCOMONT, A. Coprophages d*Amériquedu Sud nouveaux ou peu connus.

Bulletin de Ia Société Entomologíquede France,Paris, v. 1928, p. 186-194,

202-207,1928.

BROWNE, D. J.; SCHOLTZ, C. H. Phylogeny of the families of Scarabaeoidea

(Coleoptera) based oncharacters of thehindwing articulation, hindwing base

andwing venation. SystematicEntomology, Oxford, v. 20,p. 145-173,1995.

25



BROWNE, D. J.; SCHOLTZ, C. H. Evolution of the scarab hindwing
articulation and wing base: acontribution toward the phylogeny ofthe
Scarabaeidae (Scarabaeoidea: Coleoptera). Systematic Entomology, Oxford, v.
23, p. 307-326,1998.

BROWNE, D. J.; SCHOLTZ, C.H. Aphylogeny ofthe families of

Scarabaeoidea (Coleoptera). Systematic Entomology, Oxford, v. 24, p. 51-84,
1999.

CARTER, H. J. Australian Coleoptera. Notes and new species. X. Proceedings
ofthe Linnean Society ofNew South Wales, Sydney, v. 61, p. 98-110,1936.

CASTELNAU, M. leComte de[F. L. N. C. deLaporte]. Histoire natureUe des

insectes coléoptères. Tome deuxíème. Paris: Terzuolo, 1840.564 p.

D*HOTMAN, D.; SCHOLTZ, C. H. Comparative morphology ofthe male

genitalia ofderived groups ofScarabaeoidea (Coleoptera). Elytron, Barcelona,
v. 4, p. 3-39,1990.

ESCOBAR, F. Diversidad ydistribución de los escarabajos dei estiércol

(Coleoptera: Scarabaeidae: Scarabaeinae) de Colômbia. In: MARTÍN-PIERA,
F.; MORRONE, J.J.; MELIC, A. Hacia un proyecto CYTED para ei
inventário y estimación deIa diversidad entomológica en Iberoamérica:
PrIBES-2000. Zaragoza: SEA, 2000. p. 197-204.

26



FERREIRA, A. M. R. M.; GALILEO, M. H. M. Revisão taxonômica do gênero

Pedaridium Harold, 1868(Coleoptera, Scarabaeidae, Scarabaeinae, Coprini).

Iheringia,Série Zoologia, Porto Alegre, v. 74, p. 3-70,1993.

GEMMINGER, M.; HAROLD, E. von. Catalogus Coleopterorum hucusque

descriptorum synonymicus et systematicus. Monachii: Sumptu E. H. Gummi,

1869. v. 4, p. 979-1364.

GÉNIER, F.; VAZ-DE-MELLO, F. Z. A Review of Arrow's Types of

Trichillum and Pedaridium (Coleoptera: Scarabaeinae, Ateuchini) with

Description ofTwo New Species ofPedaridium. ActaZoológica Cracoviensia,

Kraków, v. 45, n. 3, p. 185-196,2002.

GILL,B. D.; VAZ-DE-MELLO, F. Z. An UnusualNewSpecies of Pedaridium

Harold (Coleoptera: Scarabaeidae: Ateuchini) From Colômbia. Journal of the

Entomological Society of Ontario, Ottawa. In press.

GILLET, J. J. E. Pars 38: Scarabaeidae: Coprinae I. In: JUNK, W.;

SCHENKLING, S. Coleopterorum Catalogus. Berlin-Gravenhage: W. Junk,

1911. v. 19, p. 1-100.

HALFFTER, G.; MATTHEWS, E. G. The naturalhistoryof dung beeties of the

subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomológica

Mexicana, México, v.l2-14,p. 1-312,1966.

HAROLD, E. von.Beitráge zurKenntnis einiger coprophagen Lamellicornien.

Berliner Entomologische Zeitschrift, Berlin,v. 3, p. 193-224,1859.

27



HAROLD, E. von. Die Choeridiiden-Gattungen Uroxys und Trichillum.
Coleopterologische Hefte, München, v. 3,p.33-55,1868.

HAROLD, E. von. Miscellen. Coleopterologische Hefte, München, v. 14, p.
208-212,1875.

HOWDEN, H. F.; YOUNG, O. P. Panamanian Scarabaeinae: taxonomy,
distribution, and habits (Coleoptera, Scarabaeidae). Contributíons of the

American Entomologícal Instítute, Ann Arbor, v. 18, n. 1, p. 1-173,1981.

LAPORTE, F. L. N. C. de. Mémoire sur cinquante espèces nouvelles ou peu
connues d'insectes. Annales deIa Société Entomologíque de France, Paris, v.
1,386-415,448,1832.

LEA, A. M. Australian dung beeties ofthe subfamily Coprides. Records of the
SouthAustralian Museum, Sydney, v. 2,p. 353-396,1923.

MARTÍNEZ, A. Addenda ycorrigenda ai trabajo de Blackwelder Checklist of
the coleopterous insects ofMéxico, Central America, the West Indies, and South

America, Scarabaeidae, Coprinae. Arthropoda, Buenos Aires, v. 1, n. 1, p. 109-
114,1947.

MARTÍNEZ, A. Scarabaeidae nuevos opoço conocidos I: dos nuevas espécies
de Scarabaeinae(Coleoptera). Publicacionesde Ia Misión de Estúdios de

Patologia Regional Argentina, Buenos Aires, v. 22, n. 79, p. 31-40,1951.

28



MARTÍNEZ, A. Catálogo de losScarabaeidae argentinos (Coleoptera). Revista

dei Museo Argentino de Ciências Naturales, Buenos Aires, v. 5, n. 1,p. 1-

126,1959.

MARTÍNEZ, A.Notas para unamonografia dei gênero Trichillum Harold, 1868

(Col. Scarabaeinae-Coprini). Revista de Ia Sociedad Mexicana de Historia

Natural, México, v. 28, p. 119-147,1967/1968.

MARTÍNEZ, A.Unanueva espécie deAteuchina (Col. Scarabaeidae, Coprini).

Anales de Ia Sociedad Científica Argentina, Buenos Aires,v. 197,n. 4-6, p.

65-68,1974.

MARTÍNEZ, A. La entomofauna de Scarabaeinae de Iaprovínciade Salta (Col.,

Scarabaeidae). Anales de Ia Sociedad CientíficaArgentina, BuenosAires,v.

216, p. 45-69,1986/1987.

MARTÍNEZ, A. Unanueva espécie de Pedaridium (Col. Scarabaeidae,

Coprini). Gayana Zoológica, Concepción, v. 56, n. 1-2, p. 21-25,1992.

MATTHEWS, E. G. A revision of the Scarabaeinae dung beeties of Austrália.

BI. tribe Coprini. Australian Journal of Zoològy Supplementary Series,

Melbourne, v. 38, p. 1-52,1976.

MATTHEWS, E. G.; STEBNICKA, Z. A reviewof Demarziella Balthasar, with

a transferfromAphodiinae to Scarabaeinae (Coleoptera: Scarabaeidae).

Australian Journal of Zoology, Melbourne, v. 34,p. 449-461,1986.

29



MEDINA, CA.; et ai. Escarabajos coprófagos (Coleoptera: Scarabaeidae:
Scarabaeinae) de Colômbia. Biota Colombiana, Bogotá, v. 2, n. 2, p. 131-144,
2002.

MONTERESINO, E.; MARTÍNEZ, A.; ZUNINO, M. Los Scarabaeinae
(Coleoptera, Scarabaeidae) de Ia Província de Córdoba, Argentina. In: Dl
TADA, I. E.; BUCHER, E. H. Biodiversidad de Ia Província de Córdoba:
fauna. Córdoba: UNRC, 1996. v. 1, p. 101-117.

MONTREUIL, O. Analyse phylogénétique et paraphylie des Coprini et
Dichotomiini (Coleoptera: Scarabaeidae). Scénario Biogeographique. Annales
de Ia Société Entomologíque de France (Nouvelle Série), Paris, v. 34, n. 2, p.
135-148,1998.

NEL, A.; SCHOLTZ, C. H. Comparative morphology of the mouthparts ofadult
Scarabaeoidea. Entomology Memoirs, Pretória, v. 80, p. 1-84,1990.

OHAUS, F. Bericht über eine entomologische Studienreise in Südamerika.
Stettiner Entomologische Zeitung, Stettin, v. 70, p. 3-139,1909.

PAULIAN, R. Une nouvelle espèce de Trichillum (Col. Scarabaeidae). Bulletin
de Ia Société Entomologíque de France, Paris, v. 41, p. 205-208,1936.

PEREIRA, F. S. Escarabeídeos americanos. In: SOCIEDADE BRASILEIRA
DE ENTOMOLOGIA. Livro de homenagem aRomualdo Ferreira de
Almeida. São Paulo: Sociedade Brasileira de Entomologia, 1946. p. 289-294.

30



PEREIRA, F. S.;MARTÍNEZ, A.Notas escarabeidológicas I. Acta Biológica

Venezuelica, Caracas, v. 2, n. 33, p. 453-465,1959.

PESSOA, S. B.;LANE, F. Coleópteros necrófagos de interesse médico-legal:

ensaio monográfico sobre a família Scarabaeidae deS.Paulo e regiões vizinhas.

Arquivos deZoologia doEstado de São Paulo, São Paulo, v.2,p. 389-504,

1941.

RATCLIFFE, B. C. New species of Coprini (Coleoptera: Scarabaeidae:

Scarabaeinae) taken from thepellage of three toedsloths (Bradypus tridactylus

L.) (Edentata: Bradypodidae) in Central Amazônia with a briefcommentary on

scarab-sloth relationships. The Coleopterists Bulletin, Lawrence, v. 34,n.4, p.

337-350,1980.

RATCLIFFE, B.C. A redescription of Trichillum hirsutum Boucomont with

notes on other interesting Brazilian Scarabaeinae (Coleoptera: Scarabaeidae).

Acta Amazônica, Manaus, v. 11, n. 1, p. 183-186,1981.

ROBINSON, M.Two new species of Neotropical Scarabaeidae (Coleoptera).

Entomological News, Philadelphia, v. 59, p. 35-36,1948.

SAYLOR, L. W. New species of Trichillum (Col. Copridae). Stylops, London,

v. 4, p. 207-208,1935.

SOLÍS, A.; KOHLMANN, B. New speciesof dung beeties (Coleoptera:

Scarabaeidae: Scarabaeinae) from Costa Rica and Panamá. Zootaxa, v. 139, p.

1-14,2003.

31



VAZ-DE-MELLO, F.Z.Estado atual deconhecimento dos Scarabaeidae s. str.

(Coleoptera: Scarabaeoidea) do Brasil. In: MARTÍN-PIERA, F.; MORRONE, J.

J.; MELIC, A. Hacia un proyecto CYTED para eiinventário y estímadón de

Ia diversidad entomológica en Iberoamérica: PrIBES-2000. Zaragoza:
Sociedad Entomológica Aragonesa, 2000. p. 183-195.

VAZ-DE-MELLO, F. Z.; CANHEDO, V. L. Duas novas espécies de
Pedaridium (Coleoptera, Scarabaeidae, Coprinae, Ateuchini). Dieringia Série
Zoologia,PortoAlegre, v. 84,p. 95-100,1998.

VAZ-DE-MELLO, F. Z.; HALFFTER, G.; HALFFTER, V. ANew Species of
Pedaridium Harold, 1868 (Coleoptera: Scarabaeidae: Ateuchini) from México
and Guatemala. The Coleopterists Bulletin, Lawrence. 2003. In press.

VERDÚ, J. R.; GALANTE, E. Anew Trichillum Harold (Coleoptera:
Scarabaeidae) from Uruguay. The Coleopterists Bulletin, Lawrence, v. 51, n. 1,
p. 93-98,1997.

VERDU, J. R.; GALANTE, E. Larval morphology and breeding behavior ofthe
genus Pedaridium Harold (Coleoptera: Scarabaeidae). Annals of the

Entomologícal Society ofAmerica, Washington, v. 94, n. 4,p. 596-604,2001.

VULCANO, M. A.; PEREIRA, F.S.Sinopse dos Passalidae e Scarabaeidae s.

str. daRegião Amazônica (Insecta, Coleoptera). In: Atasdo Simpósio sobre a

BiotaAmazônica, v. 5 (Zoologia), sem local, p.533-603.1967.

32



CHAPTER 2

A Review of Arrow's Types of Trichillum and Pedaridium with Description
ofTwo New Species ofPedaridium1

1Paper submitted to Acta Zoológica Cracoviensia, Kraków, authored byFrançois
Génier and Fernando Zagury Vaz-de-Mello.

33



•1Wi*7AV,.í«»«

1 Resumo

VAZ DE MELLO, Fernando Zagury. Revisão dos tipos de Trichillum e
Pedaridium de Arrow, com descrição de duas novas espécies de Pedaridium. In:

. Espécies originalmente nos gêneros Trichillum Harold, 1868 e
Pedaridium Harold, 1868 (Coleoptera: Scarabaeidae). 2003. Cap. 2, p.33-61.
Dissertação (Mestrado em Entomologia)-Universidade Federal de Lavras,
Lavras.*

Todo o material-tipo das espécies pertencentes aos gêneros Trichillum e
Pedaridium descritas por Arrow é estudado no presente trabalho. Como
resultado, Trichillum cristatum Arrow é transferido para ogênero Pedaridium e
as seguintes sinonímias são estabelecidas (o segundo nome sendo válido):
Pedaridium mansosotoi Martínez = Pedaridium cryptops Arrow, Pedaridium
martinezi Ferreira & Galileo = Pedaridium fulgens Arrow, Pedaridium
equatoriensis Ferreira & Galileo = Pedaridium cristatum (Arrow). Para cada
espécie são fornecidas uma diagnose e informações pertinentes. Pedaridium
margareteae sp. nov. é descrito como resultado da utilização errônea do nome
Pedaridium cryptops Arrow, ePedaridium galileoae sp. nov., como resultado da
identificação errônea de P. fulgens Arrow.

* Comitê Orientador: Dr. Júlio Neil Cassa Louzada - UFLA (Orientador) Dr
Gonzalo Halffter Salas - IEcol, Dr. Sérgio Ide - IB-SP e Dr. Mario Zunino -
UniUrb.

34



2 Abstract

VAZ DEMELLO, Fernando Zagury. A Review of Arrowi Typesof Trichillum
andPedaridium With Description of TwoNew Species of Pedaridium. In:

. Species formerly in the genera Trichillum Harold, 1868 and
Pedaridium Harold, 1868 (Coleoptera: Scarabaeidae). 2003. Chap. 2.p. 33-
61.Dissertation (Master Program in Entomology)-Universidade Federal de
Lavras, Lavras.*

Ali type material belonging to the genera Trichillum and Pedaridium described
by Arrow is studied. As a result Trichillum cristatum Arrow is transferred tothe
genus Pedaridium and the following synonymies are here established (the
second name being valid): Pedaridium mansosotoi Martínez = Pedaridium
cryptops Arrow, Pedaridium martinezi Ferreira &Galileo =Pedaridium fulgens
Arrow, Pedaridium equatoriensis Ferreira & Galileo = Pedaridium cristatum
(Arrow). A diagnosis and pertinent information is given for each species.
Pedaridium margareteae sp. nov. is herein described as a result of a misuse of
the name Pedaridium cryptops Arrow, and Pedaridium galileoae sp. nov. as a
result of misidentifícation of P. fulgens Arrow.

* Guidance Committee: Júlio Neil Cassa Louzada - UFLA (Main Advisor),
Gonzalo Halffter Salas - BEcol, Sérgio Ide- IB-SP and Mario Zunino -UniUrb.

35



3 Introduction

In the past few years agrowing interest in the systematics of Neotropical
Scarabaeinae has prompted few workers to revise certain Neotropical genera as
this group of beeties is used in rapid assessment of biodiversity (Favila &
Halffter, 1997). More specifically an effort has been done to revise poorly
known groups such as Trichillum Harold, 1868 and Pedaridium Harold, 1868
(Martmez, 1968; Ferreira & Galileo, 1993). For various reasons it was not
possible for these workers to study type material deposited in European
museums at the time. As usual these revisions have instigated a greater interest
in these groups and further investigation has revealed that additional work is
required as more systematic problems have been found at the generic as well as
specific levei in the genera (pers. obs.).

This note serves to clarify the identity of Arrow's type material, which
was not studied previously by reviewers of the groups. Since ali of the species
studied here have been recently redescribed, only short diagnoses will be
presented inaddition to illustrations ofthe dorsal view of the head.

36



4 Material and Methods

Arrow's material has been borrowed from the Natural History Museum

in London. Ali of the material with the exception of one species (Pedaridium

quadridens Arrow) consists of syntypes. For each series ofsyntypes, a lectotype

and paralectotypes are designated. Species are treated in chronological order of

publication dates. Ali of the species treated herein were originally described in

three publications (Arrow, 1913; 1931; 1932). The format and terminology used

here is the same as in Génier (1996).

Abbreviations for deposition of material are as follows:

AMBC: AyrM. Bello personal collection, Rio deJaneiro, RJ,Brazil.

BDGC: Bruce D. Gillpersonal collection, Woodlawn, Ontario Canada.

BMNH:The Natural History Museum, London, England.

CMNC: Canadian Museum of Nature, Ottawa, Canada.

FVMC: Fernando Vaz-de-Mello personal collection, Lavras, MG, Brazil.

MZSP: Museu de Zoologia da Universidade de SãoPaulo,SP, Brazil.

NHMB: Naturhistorisches Museum, Basel, Switzerland.

ZMHB: Museum fíir Naturkunde der Humboldt-Universitàt, Berlin, Germany.

37



5 Results and Discussion

Pedaridium cryptops Arrow

Pedaridium cryptops Arrow, 1913:458.

Pedaridium cryptops Arrow, 1932: 226.

Pedaridium cryptops Arrow: Balthasar, 1938:220.

Pedaridium cryptops Arrow: Blackwelder, 1944:203.

Pedaridium mansosotoi Martmez, 1951: 35, new synonymy.

Pedaridium bidens Balthasar: Ferreira &Galileo, 1993:15, misidentification.

Pedaridium mansosotoi Martmez: Ferreira &Galileo, 1993: 18.

Pedaridium cryptops Arrow: Vaz-de-Mello, 2000: 194.

Pedaridium mansosotoi Martínez: Vaz-de-Mello, 2000:194.

Lectotype E (BMNH): Syntype (blue disc)/ Type (red disc, upside down)/
LECTOTYPE (red paper)/ Jahaty, Prov. Goyas. Brésil (green paper)/ Fry coll.,
1905-100/ Pedaridium cryptops type Arrow (Arrowi handwriting)/ Pedaridium
cryptops Arr. M.E. Bacchus det 1975, SYNTYPE/ LECTOTYPE,

PEDARIDIUM CRYPTOPS ARROW, dés. F. Génier &F. Vaz-de-Mello, 2001.
The lectotype is designated here in order to establish the specimen with the
diagnostic character (acute clypeal teeth) to become the type ofthe species as
the two other specimens in the syntypic series represem two different species.

Diagnosís. Clypeal teeth (HGURE 1) long, acute, dorsal surface bellow the

surface of clypeus and separated with a strong and sharp carina. Median
emargination of clypeus (FIGURE 1) moderately broad and very shallow in
dorsal view. Clypeal punctures (HGURE 1) feebly impressed, lacking sharp

38



raspy edge anteriorly and separated by more than one diameter anteriorly.

Dorsum glossy between punctures and lacking distinct opalescent sheen (in

clean specimens). Elytral striae with punctures feebly umbilicate and feebly

encroaching on intervals. Elytral interstriae 1-6 with either one or two aligned

rows of setae on disc.

Remarks. The material deposited in the Natural History Museum in London

consists of 3 specimens belonging to three different species. Two specimens

collected in Jahaty (sic) (=Jataí) (Goiás) and one specimen from Natal (Rio

Grande do Norte). The specimen from Natal, which is labeled by Arrow as

"Pedaridium cryptops" was collected by W.M. Mann. According to the original

description, two specimens have been collected by Mr. W. Mann. This is

suggesting that at least one specimen is missing from the original series. One of

the two specimens collected inJatai bear Arrowi original label, which is stating

"type". This specimen is selected as lectotype. Arrow's description clearly states

that the clypeus of his new species possesses acute teeth ("utrique acute

dentato"), and this is especially visible in this specimen. The other two

specimens have small triangular teeth on the clypeus. It is possible that Arrow

considered that they wereabraded, andincluded these two specimens in the type

series. The lectotype is identical to Pedaridium mansosotoi Martínez and we

synonymize this species herein as a result. The other specimen from the same

locality is a Pedaridium bidens Balthasar and therefore excluded from the type

series. The specimen from Natal represents consequently a new species, which

we de§cribe here.

Pedaridium fulgens Arrow

Pedaridiumfulgens Arrow, 1913:458.

39



Pedaridiumfulgens Arrow, 1932:226.

Pedaridiumfulgens Arrow: Blackwelder, 1944: 203.

Pedaridium martinezi Ferreira &Galileo, 1993: 30, new synonymy.

Lectotype E (BMNH): Syntype (blue disc)/ Type (red disc, upside down)/
LECTOTYPE (red paper)/ N. ARGENTINA, Rio Salado, Gran Chaco, E.
Wagner. 1907-384/ Pedaridium fulgens type Arrow (Arrow's handwriting)/
Pedaridium fulgens Arrow, M.E. Bacchus det 1975, SYNTYPE/ LECTOTYPE
PEDARIDIUM FULGENS ARROW, dés. F. Génier &F. Vaz-de-Mello, 2001.
Paralectotype: Same data as lectotype (1EBMNH).

The lectotype is here designated to avoid confusion if this species would
eventually reveal to be a species complex.

Diagnosís. Clypeus (HGURE 2) with three teeth on each side, a large and acute
anterior tooth and two smaller broadly triangular teeth laterally. Median
emargination of clypeus (FIGURE 2) broadly arcuate in dorsal view. Dorsum
glossy between punctures and with adistinct coppery sheen. Elytral striae with
punctures distinctly encroaching on intervals.

Remarks. The material consists of two females syntypes. The cleanest
specimen, which bear Arrow's handwritten label, is selected as lectotype. The
other specimen, with the same data, is designated as paralectotype. After a
careful examination of the closely related species we carne to the conclusion that
Pedaridium martinezi Ferreira &Galileo is identical in ali respect to P. fulgens
Arrow, consequently we synonymize herein this species. The species currently
recognized as P. fulgens is anew taxon, which is described here. In the original
description, Arrow wrote "...dentibus haud valde approximatis, denticuloque
utrinque externo...", giving rise to the interpretation of one pair of externai

40



clypeal denticles instead of two, as did Balthasar (1938), originating the

misidentification of the new species (that bear one pair of externai denticles)

with the true P.fulgens (bearing two pairs).

Pedaridium argentinum Arrow

Pedaridium argentinum Arrow, 1913:459.

Pedaridium rugiceps Arrow, 1913: 458, synonymized by Ferreira & Galileo,

1993:24-25.

Pedaridium argentinum Arrow, 1932:226.

Trichillum elongatum Balthasar, 1934: 24, synonymized by Martmez, 1987: 60.

Pedaridium argentinum Arrow: Balthasar, 1938: 220.

Pedaridium argentinum Arrow: Blackwelder, 1944: 203.

Pedaridium argentinum Arrow: Martínez, 1959:62.

Pedaridium elongatum (Balthasar): Martínez1968: 119.

Pedaridium argentinum Arrow:Martínez, 1987: 60.

Pedaridium argentinum Arrow: Ferreira& Galileo, 1993:24.

Pedaridium argentinum Arrow: Monteresino etai, 1996:107.

Lectotype T (BMNH): Syntype (blue disc)/ Type (red disc, upside down)/

LECTOTYPE (red paper)/ N. ARGENTINA, Rio Salado, Gran Chaco, E.

Wagner. 1907-384/ Pedaridium argentinum type Arrow (Arrow's handwriting)/

Pedaridium argentinum Arrow, M.E. Bacchus det 1975, SYNTYPE/

LECTOTYPE, PEDARIDIUM ARGENTINUM ARROW, dés. F. Génier & F.

Vaz-de-Mello, 2001.

Paralectotypes: Same data as lectotype (3IT BMNH); same data except: Icano

(ir, 2EE BMNH) (1EMNHN).

41



The lectotype isdesignated here inorder to select a male specimen, which bears
the diagnostic characters to separate this species from closely related species and
to establish "Rio Salado" asthetype locality.

Diagnosís. Body dark reddish brown, elongate, almost parallel sided medially.
Clypeus (HGURE 3) with two large uptumed triangular teeth. Median
emargination of clypeus (HGURE 3) broadly V-shaped. Clypeal punctures
coalescent, forming more or less transverse ridges. Dorsum glossy between
punctures lacking metallic sheen. Elytral striae narrow, almost parallel sided,
punctures feebly indicated. Elytral interstriae 2-6 with two aligned rows ofsetae
on disc.

Remarks. Arrowi type material consists of 7 specimens in BMNH and one

specimen, which was deposited at the Museum national d'Histoire naturelle in

Paris. The lectotype male is here designated and bears Arrowi "type"
handwritten label, ali ofthe rest ofthe syntype series matches the lectotype and
specimens are here designated as paralectotypes. In his identification key Arrow
is using the name rugiceps instead of argentinum. This has been noted by
Ferreira & Galileo (1993), who considered P. rugiceps as an objective
synonymy ofP. argentinum. This has been obviously a typographical mistake as

the name rugiceps in the copy of the reprint located at the CMNC has been

corrected by Arrow himself to argentinum. Martínez (1986) has synonymized P.
elongatum Balthasar without any comments. The three female type specimens of
P. elongatum have been compared to Arrow's types and the synonymy
established by Martínez is here confirmed.

Trichillum hystrix Arrow

42



Trichillum hystrixArrow, 1931: 609.

Trichillum hystrix Arrow: Paulian, 1936:206.

Trichillum hystrix Arrow: Balthasar, 1939:25.

Trichillum hystrix Arrow: Blackwelder, 1944: 204.

Trichillum (Eutrichillum) hystrixArrow: Martínez, 1968: 120.

Trichillum (Eutrichillum) hystrixArrow: Ratcliffe, 1980: 341.

Trichillum hystrix Arrow: Martínez, 1987:60.

Lectotype T (BMNH): Syntype (blue disc)/ Type (red disc, upside down)/

LECTOTYPE (red paper)/ Estância Ia Noria, Rio San Javier, Santa Fe

Argentine, G.E. Bryant, 27.XH.1911/ G. Bryant Coll., 1919-147/ Trichillum

hystrix type Arrow (Arrow's handwriting)/ Trichillum hystrix Arrow, M.E.

Bacchus det 1975, SYNTYPE/ LECTOTYPE, TRICHILLUM HYSTRIX

ARROW, dés. F. Génier & F. Vaz-de-Mello, 2001.

Paralectotypes: Same data as lectotype except: XII.1911 (ir, 1E BMNH),

10.Xn.1911 (1E CMNC), 14.XH.1911 (ir BMNH), 20.XII.1911 (2EE

BMNH), 23.XCL1911 (2EEBMNH), 3.1.1912 (ir BMNH).

The lectotype isdesignated here inorder toselect a male specimen, which bears

thedistinctive characters to separate this species from closely related species.

Diagnosís. Body reddish brown to dark brown, oval, sides broadly arcuate

medially in dorsal view. Clypeus (HGURE 4) with two large closely set

uptumed blunt teeth. Anterior margin of clypeus and gena (HGURE 4)

distinctly uptumed in non-abraded specimens. Median emargination of clypeus

(HGURE 4) U-shaped. Clypeal punctures (HGURE 4) of different sizes, more

or less well defined and in most cases separated by more than one diameter.

Eyes (HGURE 4) at the same levei as the rest of the head, elongate oval,

43



approximately three times as long as wide. Dorsum glossy between punctures
lacking metallic sheen. Elytral striae narrow, almost parallel sided, punctures
feebly indicated. Elytral interstriae 1-2 with a more orless complete row ofsetae
on externai side, interstriae 3-6 witha single row of setaeon the internai side on

disc. Anterior tarsal claws of male (similar to HGURE 11) expended into
translucent lobe basally, distai portion slender and abruptly bent forward. Apex
of large sclerite of internai sac of edeagus with a single acute dentiform
projection (two in P. hirsutum Boucomont).

Remarks. Arrow's syntype series consists of4IT and 5EE specimens (BMNH)
plus an additional female specimen ^-collection Antônio Martínez (now in
CMNC). Ali of the specimens are from the same locality, but collected on
different dates. A slightly teneral male specimen showing a complete (not
abraded) anterior margin ofthe clypeus and complete vestiture has been selected

for lectotype. The lectotype also bears Arrow's original "type" label. Trichillum
hystrix belongs to a complex of very closely related species which can be
separated with certainty only by the configuration of the sclerites of the internai

sac ofthe edeagus. Martínez (1987) considered T. hystrix undistinguishable from
Trichillum boucomonti Saylor. The relations between T. boucomonti and T.

hirsutum Boucomont will be discussed in a separate publication in preparation.
The type of T. hirsutum Boucomont has been studied and appeared to be the
closest species to T. hystrix. It can be separated from T. hystrix by the shape of
the dorsal portion of the eyes and the shape of the sclerite of the internai sac (see
diagnose) and the apicoventral portion of the tube of edeagus, which bear a
single small tubercle in T. hystrix and a broader triangular tubercle divided
medially in T. hirsutum. However, the type of T. hirsutum is a female from São

Paulo (state or city?) and the previously mentioned diagnostic characters are
based on anon type specimen dissected and collected from São Paulo deposited

44



in the CMNC. This specimen seems to match Boucomont type and it seems

reasonable here to believe that T hirsutum is in fact different from T. hystrix.

Pedaridium ohausi (Arrow)

Trichillum ohausi Arrow, 1931:610.

Trichillum ohausi Arrow: Paulian, 1936: 206.

Trichillum ohausi Arrow: Balthasar, 1939: 22.

Trichillum ohausi Arrow: Blackwelder, 1944: 204.

Pedaridium ohausi (Arrow): Martínez, 1968: 119.

Pedaridiumohausi (Arrow): Ferreira & Galileo, 1993: 12.

Lectotype T (BMNH): Syntype (blue disc)/ Type (red disc, upside down)/

LECTOTYPE (red paper)/ Loja Punzara, F. Ohs. 7.8.05/ Ecuador, F. Ohaus,

B.M. 1931-387/ Trichillum ohausi type Arrow (Arrow's handwriting)/

Trichillum ohausi Arrow, M.E. Bacchus det 1975, SYNTYPE/ LECTOTYPE,

TRICHILLUM OHAUSI ARROW, dés. F. Génier & F. Vaz-de-Mello, 2001.

Paralectotypes: Same data as lectotype except: Loja Calvário, 4.8.05 (1 sex

undetermined BMNH), Loja (1 sexundetermined BMNH), Ecuador, no locality

(2 sex undetermined BMNH).

The lectotype is designated here in order to select a malespecimen, which bears

the distinctivecharactersto separatethis species fromclosely related species.

Diagnosís. Body dark brown with feeble but distinct greenish and coppery

reflections on elytra, elongate, almost parallelsidedmedially. Clypeus(HGURE

5) with two large slightly uptumed sharp triangular teeth. Median emargination

of clypeus (HGURE 5) broadly arcuate. Clypeal punctures (HGURE 5)

separated by at least one diameter and of three different types, minute without

45



setae, larger with setae and few ill defíned punctures with asmall sharp granule
on each side of clypeus. Eyes (HGURE 5) large, oval in shape dorsally. Dorsum
with feeble but distinct microsculpture between punctures. Elytral striae narrow,
almost parallel sided, punctures feebly indicated separated by at least three
diameters on disc. Elytral interstriae 1-6 with few scattered short setae on disc,
apical declivity with regularly spaced setae, interstriae 1, 3, 5 with few distinct
sharply granulate punctures on disc.

Remarks. Arrow's syntype series consist of 5 specimens, the specimen with
Arrow's hand written type label agree with the original description and has been
selected for lectotype. Because this species does not exhibit obvious secondary
sexual characters, this specimen has been dissected and is amale. The remaining
four specimens (undetermined sex) are ali conspecifíc and designated
paralectotypes. The name P. ohausi is appropriately applied in Ferreira &
Galileo (1993).

Pedaridium cristatum (Arrow)

Trichillum cristatum Arrow, 1931: 610.

Trichillum cristatum Arrow: Paulian, 1936: 206.

Trichillum cristatum Arrow: Balthasar, 1939: 22.

Trichillum cristatum Arrow: Blackwelder, 1944: 204.

Trichillum cristatum (Arrow): Martínez 1968: 119.

Pedaridium cristatum (Arrow), new combination.

Pedaridium equatoriensis Ferreira &Galileo 1993: 14, new synonymy.

Lectotype E (BMNH): Syntype (blue disc)/LECTOTYPE (red paper)/ 19
(handwriting)/ S. ECUADOR, Piscobamba, M. Witt (recto), Lll. Lobl (verso)/

46



Trichillum externepunctatum ? Borre (handwriting)/ Trichillum cristatum

Arrow, M.E. Bacchus det 1975, SYNTYPE/ LECTOTYPE, TRICHILLUM

CRISTATUM ARROW, dés. F. Génier & F. Vaz-de-Mello, 2001.

Paralectotype: ECUADOR, Loja, Ohaus S. (recto), 2200m. malader Stadt

(verso)/ Ecuador, F. Ohaus, B.M. 1931-387/ Trichillum sp.l. unia

(handwriting)/ Trichillum cristatum Arrow, M.E. Bacchus det 1975, SYNTYPE

(1EBMNH).

The lectotype is designated here in order to select a male specimen which

matches Arrow's description and which bears the distinctive characters to

separate this species from closely related species and toestablish Piscobamba as

type locality.

Diagnosís. Body dark brown with distinct greenish and coppery reflections on

dorsum, elongate oval, sides broadly arcuate. Clypeus (HGURE 6) with two

slightly uptumed broadly triangular teeth and distinctly emarginate clypeogenal

área. Median emargination of clypeus (HGURE 6) broadly arcuate. Clypeal

punctures (HGURE 6) separated by at least one diameter and of three different

types, minute without setae on disc, larger coarse with setae near eyes and

posterior edge and few illdefíned punctures with a small sharp granule on each

side of clypeus. Eyes (HGURE 6) moderate in size, oval in shape dorsally.

Dorsum withfeeblebut distinct microsculpture between punctures. Elytralstriae

moderately large, with punctures encroaching on intervals, punctures not

umbilicate, separated byapproximately one diameter on disc. Elytral interstria 1

with a single row of setiferous punctures along externai edge, interstriae 2-6

witha rowsof regularly spacedsetiferous punctures on each side.

Remarks. Arrow's syntype series consists of 3 specimens. The specimen with

Arrow's hand written type label is a T. ohausi with abraded clypeal teeth and

47



does not agree with the original description. In order to reflect Arrow's species
concept for this taxon the male specimen, which agree with the original
description, has been selected for lectotype, even thought it was not labeled as
such by Arrow. The other specimen, which matches the original description, is a
female and differs externally from the male by its more densly punctate head
and less elevated cephalic carina. We studied the holotype ofP. equatoriensis
Ferreira &Galileo, a female, which has been collected from the same locality
(Loja), and is in ali respect identical to Trichillum cristatum Arrow. We

consequently synonymize P. equatoriensis with T. cristatum here.

Pedaridium paranense Arrow

Pedaridium paranenseArrow, 1932:224.

Pedaridiumparanense Arrow: Balthasar, 1938:219.

Pedaridium paranense Arrow: Blackwelder, 1944: 203.

Pedaridium paranensis Arrow: Ferreira & Galileo, 1993: 9.

Pedaridium paranensis Arrow: Vaz-de-Mello &Canhedo, 1998: 100.

Pedaridium paranensis Arrow: Vaz-de-Mello, 2000:194.

Lectotype T (BMNH): Syntype (blue disc)/LECTOTYPE (red paper)/ 11.248
(handwriting)/ Castro, Paraná, S. Brazil, 1926-304/ Pedaridium paranense Arr,
ME. Bacchus det 1975, SYNTYPE/ LECTOTYPE, PEDARIDIUM
PARANENSE ARROW, dés. F. Génier &F. Vaz-de-Mello, 2001.

Paralectotypes: Same data aslectotype (2IT, 1EBMNH).

The lectotype is designated here in order to select a male specimen which bear
the distinctive characters to separate this species from closely related species.

48



Diagnosís. Clypeal teeth (HGURE 7) lacking. Median emargination of clypeus

(HGURE 7) moderately broad and shallow in dorsal view. Clypeal punctures

(HGURE7) coarse, lacking sharp raspy edge anteriorly and separated by one to

two diameters anteriorly, each puncture with a long stout seta. Dorsum with

surface between punctures dull on disc of pronotum and elytra, becoming

glossier laterally, with distinct opalescent sheen and some metallic reflections

(in clean specimens). Elytral stria 1effaced on anterior half, stria 2 well defmed

on entire length, striae 3-5 effaced posteriorly, stria 6 effaced on entire length.

Elytral interstriae 1-6 with two aligned row oflong and trout setae ondisc.

Remarks. Arrow's syntype series consists of 4 specimens collected from the

same locality (Castro) in Paraná state of Brazil. The specimen, which bears

Arrow's handwritten type label, is a female and we have decided to choose a

male specimen instead, which show better diagnostic character instead, for

lectotype. Ali specimens areconspecific and therefore included in the lectotype

series. The usage of the name P. paranense is correct in Ferreira & Galileo

(1993). Ferreira & Galileo (1993) have changed, in error, the accord of the

specific epithet. Pedaridium is neutral therefore, paranense is the proper

spelling.

Pedaridium quadridens Arrow

Pedaridium quadridens Arrow, 1932: 225.

Pedaridium quadridens Arrow: Balthasar, 1938: 220.

Pedaridium quadridens Arrow: Blackwelder, 1944:203.

Pedaridium quadridens Arrow: Martínez, 1959: 62.

Pedaridium quadridens Arrow: Martínez, 1987: 60.

Pedaridium quadridens Arrow: Ferreira& Galileo, 1993: 33.

49



Holotype E(BMNH): Holotype (red disc)/ Santa Elena, Entre Rios, Argentina
G. E. Bryant, 30.1.1912/ G. Bryant Coll., 1919-147/ Pedaridium quadridens, type
Arrow (Arrow's handwriting)/ Pedaridium quadridens Arr, M.E. Bacchus det
1975, HOLOTYPE.

Diagnosís. Clypeal margin (HGURE 8) with 4 acute and uptumed teeth, two
inner teeth slightly longer and slender, teeth with anterior edge as acontinuation
of the fine, sharp and uptumed edge of the clypeus. Median emargination of
clypeus (HGURE 8) broad and very shallow in dorsal view, much wider than
distance between lateral and median teeth. Clypeal punctures (HGURE 8) ill
defined, each puncture with afine and long seta and a small and sharp granule
anteriorly, each granule separated by more than one diameter. Front with closely
set more or less well defined umbilicate punctures. Dorsum glossy between
punctures and some individual showing distinct metallic reflections (in clean
specimens). Elytral striae with punctures forming a sinuous elevated carina,
strongly encroaching on intervals. Elytral interstriae 1-6 with two aligned row of
setae on disc, each row composed of along erect brownish seta alternating with
a shorter apposed diagonally oriented andwhitish seta.

5.1 New Species

Pedaridium margareteae sp. nov.

(HGURES9,11-14)

Pedaridium cryptops sensu Balthasar, 1938:220.

Pedaridium cryptops sensu Ferreira & Galileo, 1993: 20.

Pedaridium cryptops sensu Vaz-de-Mello, 2000 (pars): 194.

50



Etymology. We dedicate this species to Ana Margarete M. Ferreira, which

recently revised the genus Pedaridium.

Diagnosís. Clypeal teeth (HGURE 9) short, triangular, dorsal surface at the

same levei as the clypeus and separated with a fine and sharp carina. Median

emargination of clypeus (HGURE 9)broad and shallow in dorsal view. Clypeal

punctures (HGURE 9) deeply impressed, raspy and separated by less than one

diameter anteriorly. Dorsum feebly glossy between punctures and with a distinct

opalescent sheen (in clean specimens). Elytral striae with punctures umbilicate

and encroaching on intervals. Elytral interstriae 1-6 with two aligned row of

setae on disc.

Holotype. T. Length 4.4 mm, greatest width 2.5 mm. Body elongate oval in

dorsal view. Color brownish with a distinct opalescent sheen (in clean

specimens). Head (HGURE 9). Anterior edge with a broad median

emargination, limited on each side by a small triangular tooth, almost straight on

a short distance laterally. Dorsal surface glossy between punctures, punctures

(HGURE 9) rounded, slightly raspy and denser anteriorly, separated by about

one diameter on anterior half, each punctures bearing an elongate squamiform

setae. Eyes (HGURE 9) with dorsal portion narrow, approximately three times

longer than wide. Pronotum. Evenly convex, surface glossy between punctures,

punctures and setae similar in size and density to those on posterior half of the

head on disc, punctures slightly larger and denser with longer setae on lateral

declivities. Elytra. Distinctiy tentiform on disc. Elytral striae 1 and 6 narrower

and shadowily impressed. Striae 2-5 wider, deeply impressed and sharply

delimited, with punctures more orless oval and umbilicate. Striae 1-3 wider and

much more deeply impressed on apical declivities. Interstriae 1-6 with two

51



aligned rows ofraspy and setose punctures on each side, setae squamiform and

longer on internai side. Legs. Anterior tarsal claws (HGURE 11) expended into
translucent lobe basally, distai portion slender and abraptly bent forward.

Posterior tibia produced into a hook on internai margin at apex. Thoracic

sterna. Mesostemum distinctiy concave, punctures coarse, umbilicate and large
smaller and more widely separated along midline. Metasternum with punctures
small on disc, lateral lobes with coarse, raspy, confluent transverse puncture
anteriorly and lacking puncture posteriorly except along metacoxal edge, median
lobe with anterior margin bluntly angulate, with few scattered setose punctures.
Abdômen. Stemites 2-5 with a single row of closely set elongate puncture
covering the entire length laterally, punctures divided in two row on a small

surface of segment 5 laterally. Segment 6 covered with more or less rounded

umbilicate punctures, punctures becoming smaller and feebly impressed
medially and posteriorly. Pygidium with more or less fused oval punctures,
punctures becoming smaller and feebly impressed at apex. Male genitalia.
Edeagus asinHGURES 13-14, sclerites ofinternai sac asinHGURE 12.

Allotype. E. Length 4.9 mm, greatest width 2.8 mm. Similar to male except:
Anterior tarsal claws slender and evenly arcuate in lateral view. Posterior tibia

lacking hook on internai margin at apex. Abdominal stemite 6 with a V-shape
sulcus posteriorly, sulcus sharply delimited anteriorly.

Material studied. 18 IT, 23 EE, 9 sex? (AMBC, BMNH, CMNC, FVMC,
MZSP)

Holotype T (MZSP, ex-FVMC): BRASIL: PI[=Piauí], S[ão]. R[aimundoJ.
Nonato, P[arque].N[acional da].S[erra]. da Capivara, 1-1999, CA Matrangolo
Allotype E(MZSP, ex-FVMC): Same data as holotype.

Paratypes: BRAZIL: Bahia: Caetité, C. Uran. Lagoa Real - INB, 8-16.1.2000 (4
FVMC); Encruzilhada, XII.1980/ A. Martínez eM. Alvarenga (3 CMNC); same

52



asbefore except XÜ-1997, Vaz-de-Mello &BeUo (4FVMC); Jequié, n.1995, C.

Sperber (1 FVMC); S. Antônio da Barra, XI-XÜ.1888, Gounelle (1 ZMHB);

Vila Nova, 1908 (1 MZSP); same as before except: X.1938, Dr. Nick, Coll.

Martínez (2 CMNC). Mato Grosso: Chapada do Guimarães, XI.1963,

Alvarenga (9 MZSP). Minas Gerais: Águas Vermelhas, XII.1997, A. BeUo (4
AMBC). Pernambuco: Pery-Pery, V-VI.1892, Gounelle (2 ZMHB); same as

before except: XI-XÜ.1892 (1 ZMHB). Piauí: São Raimundo Nonato, PNSerra

daCapivara, 1.1999, CA. Matrangolo (5 FVMC). RioGrande do Norte: Natal

(5 NHMB); same as before except: W. M. Mann, Stanford Exped., 1913-56 (1

BMNH); same as before except: Dl 1952, Alvarenga (4 MZSP); IE.1954,

Alvarengaleg, Coll.Martínez (1 CMNC).

Remarks. Paratypes vary in length from 4.0-4.5 mm. The two paratypes from

Villa Nova (Bahia) in CMNC have the clypeal teeth less widely separated and

the parameres present a less sinuous internai edge in frontal view. However, the

sclerites of the internai sac are identical and for this reason this variation is here

consider intraspecific.

Pedaridium galüeoae sp. nov.

(HGURES 10,15-17)

Pedaridium fulgens sensu Balthasar, 1938:458.

Pedaridiumfulgens sensu Martínez 1959: 62.

Pedaridiumfulgens sensu Ferreira & Galileo 1993: 37.

Pedaridiumfulgens sensu Monteresino et ai., 1996: 107.

Etymology. A patronymic in honor of Maria Helena M. Galileo, a Brazilian

cerambycidologist who was very helpful inproviding specimens for our studies.

53



Diagnosís. Clypeus (HGURE 10) with two teeth on each side, median teeth
acute, triangular and moderately large, narrower than lateral teeth at base.

Median emargination of clypeus (HGURE 10) broadly V-shaped and
moderately shallow in dorsal view. Clypeal punctures (HGURE 10) fine,
transverse and separated by two to four diameters anteriorly. Dorsum glossy
between punctures, with feeble microsculpture along margins of pronotum and
elytra, with adistinct coppery sheen (in clean specimens). Elytral striae straight
and sharply delimited throughout, with punctures feebly indicated and only
slightly umbilicate. Elytral interstriae flat, interstria 1 with a single row of
setiferous punctures along externai edge, interstriae 2-6 with a rows of
irregularly spaced setiferous punctures oneach side.

Holotype. T. Length 3.3 mm, greatest width 1.8 mm. Body elongate oval in
dorsal view. Color dark reddish brown, elytra lighter in color, dorsum with a
distinct coppery sheen. Head (HGURE 10). Anterior edge broadly V-shaped
medially, median emargination limited on each side by a small triangular tooth
and a second blunt and broader lateral tooth, edge almost straight on a short
distance laterally. Dorsal surface glossy between punctures on disc and with
feeble microsculpture along margins, punctures (HGURE 10) transverse,
separated by two to four diameters on anterior half, each punctures bearing an
elongate and feeble squamiform setae. Eyes (HGURE 10) with dorsal portion
oval, approximately two times longer than wide. Pronotum. Evenly convex,
surface glossy between punctures, punctures oval and longitudinally oriented,
distinctiy coarser than those on dorsal surface of the head, lacking on midline of
disc, setae similar in size to those on posterior half of the head on disc, punctures
slightly larger and denser with longer setae on lateral declivities. Elytra. Evenly
convex on disc. Elytral striae 1-6 simüarly impressed, narrow and sharply

54



delimited, with punctures feebly indicated. Interstria 1 with a single row of

setiferous punctures alongexternai edge, interstriae 2-6 witha row of irregularly

spaced setiferous punctures on each side. Legs. Anterior tarsal claws simply

arcuate, similarin shape and size to middle and posterior claws. Posterior tibia

lacking dentiform process on internai margin at apex. Thoracic sterna.

Mesostemum convex, punctures oval, moderately coarse, umbilicate, smaller

and more widely separated along midline. Metasternum withpunctures small on

disc, lateral lobes with coarse, raspy, confluent transverse puncture anteriorly

and lackingpuncture p