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ZOOTAXA
Synopsis of the new subtribe Scatimina (Coleoptera:
Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of
twelve new genera and review of Genieridium, new genus
FERNANDO Z. VAZ-DE-MELLO
Magnolia Press
Auckland, New Zealand
1955
VAZ-DE-MELLO
2 · Zootaxa 1955 © 2008
Magnolia Press
Fernando Z. Vaz-de-mello
Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with
descriptions of twelve new genera and review of Genieridium, new genus
(Zootaxa 1955)
75 pp.; 30 cm.
5 Dec.2008
ISBN 978-1-86977-307-6 (paperback)
ISBN 978-1-86977-308-3 (Online edition)
FIRST PUBLISHED IN 2008 BY
Magnolia Press
P.O. Box 41-383
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New Zealand
e-mail: zootaxa@mapress.com
http://www.mapress.com/zootaxa/
© 2008 Magnolia Press
All rights reserved.
No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any
means, without prior written permission from the publisher, to whom all requests to reproduce copyright
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This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose
other than private research use.
ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
Accepted by A. Smith: 2 Oct. 2008; published: 5 Dec. 2008 3
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2008 · Magnolia Press
Zootaxa 1955: 1–75 (2008)
www.mapress.com/zootaxa/
Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae:
Scarabaeinae: Ateuchini), with descriptions of twelve new genera
and review of Genieridium, new genus
FERNANDO Z. VAZ-DE-MELLO
Instituto de Ecología A.C., Departamento de Biodiversidad y Ecología Animal, Km 2.5 Carretera Antigua a Coatepec, 351, Congreg-
ación El Haya, 91070 Xalapa, Veracruz, Mexico.
Present Address: Universidade Federal de Mato Grosso, Instituto de Biociencias, Departamento de Biologia e Zoologia, Av.
Fernando Correa da Costa, s/n, CCBS II, Boa Esperanca, Cuiaba - MT 78060-900, Brazil.
E-mail: vazdemello@gmail.com
Table of contents
Abstract ...............................................................................................................................................................................3
Resúmen ..............................................................................................................................................................................4
Resumo ................................................................................................................................................................................5
Introduction .........................................................................................................................................................................6
Material and methods ..........................................................................................................................................................9
Scatimina, new subtribe ....................................................................................................................................................10
Key to the genera of Scatimina ........................................................................................................................................14
1. Besourenga, new genus ..........................................................................................................................................16
2. Bradypodidium, new genus ....................................................................................................................................18
3. Degallieridium, new genus ....................................................................................................................................20
4. Eutrichillum Martínez, 1969 new status ................................................................................................................22
5. Feeridium, new genus ............................................................................................................................................24
6. Genieridium, new genus ........................................................................................................................................26
Key to species of Genieridium ...............................................................................................................................27
7. Leotrichillum, new genus .......................................................................................................................................32
8. Martinezidium, new genus .....................................................................................................................................34
9. Nunoidium, new genus ...........................................................................................................................................36
10. Onoreidium, new genus ........................................................................................................................................37
11. Pedaridium Harold, 1868 .....................................................................................................................................40
12. Pereiraidium, new genus ......................................................................................................................................41
13. Scatimus Erichson, 1847 ......................................................................................................................................42
14. Scatrichus Génier & Kohlmann, 2003 .................................................................................................................42
15. Silvinha, new genus ..............................................................................................................................................43
16. Trichillidium, new genus ......................................................................................................................................43
17. Trichillum Harold, 1868 .......................................................................................................................................47
Acknowledgments .............................................................................................................................................................71
Literature cited .................................................................................................................................................................. 71
VAZ-DE-MELLO
4 · Zootaxa 1955 © 2008
Magnolia Press
Abstract
The new subtribe Scatimina (part of the tribe Ateuchini) is described and defined to include the following 17 genera, of
which 12 are new: Scatimus Erichson; Scatrichus Génier & Kohlmann, 2003; Trichillum Harold, 1868; Pedaridium
Harold, 1868; Eutrichillum Martínez, 1969, new status; Besourenga, new genus (type species Trichillum minutum Say-
lor); Bradypodidium, new genus (type species Trichillum bradyporum Boucomont); Degallieridium, new genus (type-
species Degallieridium lilliputanum, new species); Feeridium, new genus (type species Feeridium woodruffi, new spe-
cies); Genieridium, new genus (type species Pedaridium bidens Balthasar); Leotrichillum, new genus (type species
Pedaridium louzadaorum Vaz-de-Mello & Canhedo); Martinezidium, new genus (type species Pedaridium galileoae
Génier & Vaz-de-Mello); Nunoidium, new genus (type species Pedaridium argentinum Arrow); Onoreidium, new genus
(type species Trichillum cristatum Arrow); Pereiraidium, new genus (type species Pedaridium almeidai Pereira); Sil-
vinha, new genus (type species Silvinha unica, new species); and Trichillidium, new genus (type species Pedaridium
quadridens Arrow). The subtribe Ateuchina Laporte is here defined and includes Ateuchus Web er, Deltorhinum Harold,
Aphengium Harold and Sinapisoma Boucomont (transferred from Canthonini). The genera Scatonomus Erichson, Ano-
miopus Westwood and Hypocanthidium Balthasar are transferred from Ateuchini to Canthonini, and the genera Canthid-
ium Erichson, Parachorius Harold (formerly Ateuchini) and Holocanthon Martínez & Pereira (formerly Canthonini) are
transferred to Coprini. The genera Bdelyropsis Pereira, Vulcano & Martínez, Bdelyrus Harold, Coptorhina Hope,
Delopleurus Erichson, Demarziella Balthasar, Onychothecus Boucomont, Paraphytus Harold, Pedaria Laporte, Plero-
nyx Lansberge, Pseuduroxys Balthasar, Sarophorus Erichson and Uroxys Westwood are considered as incerta sedis in the
Ateuchini, not fitting into Ateuchina or Scatimina. A key is presented to the genera of Scatimina, and another for species
of the genus Genieridium, which are also diagnosed. Each genus of Scatimina is diagnosed, has a complete species list,
and includes remarks on affinities, composition and distribution. All genera except Scatimus and Scatrichus are
(re)described, and examined material is listed for each species. The following three new species are described: Degallie-
ridium lilliputanum, new species; Feeridium woodruffi, new species and Silvinha unica, new species. The following 32
new combinations are established (original genus in parenthesis): Besourenga minutus (Saylor) (Trichillum); B. vej-
dovskyi (Balthasar) (Trichillum); B. amarillai (Aguilar) (Pedaridium); B. horacioi (Martínez) (Trichillum); Bradypodid-
ium adisi (Ratcliffe) (Trichillum); B. bradyporum (Boucomont) (Trichillum); B. venezuelense (Ferreira & Galileo)
(Pedaridium); Eutrichillum arcus (Solís & Kohlmann) (Trichillum); E. hirsutum (Boucomont) (Trichillum); E. hystrix
(Arrow) (Trichillum); Genieridium bidens (Balthasar) (Pedaridium); G. bordoni (Martínez) (Pedaridium); G. cryptops
(Arrow) (Pedaridium); G. margareteae (Génier & Vaz-de-Mello) (Pedaridium); G. medinae (Gill & Vaz-de-Mello)
(Pedaridium); G. paranense (Arrow) (Pedaridium); G. zanunciorum (Vaz-de-Mello & Canhedo) (Pedaridium); Leotri-
chillum louzadaorum (Vaz-de-Mello & Canhedo) (Pedaridium); Martinezidium fulgens (Arrow) (Pedaridium); M. gali-
leoae (Génier & Vaz-de-Mello) (Pedaridium); M. martinsi (Ferreira & Galileo) (Pedaridium); M. maya (Vaz-de-Mello,
Halffter, & Halffter) (Pedaridium); Nunoidium argentinum (Arrow) (Pedaridium); Onoreidium bottimeri (Howden &
Young) (Pedaridium); O. cristatum (Arrow) (Trichillum); O. howdeni (Ferreira & Galileo) (Pedaridium); O. ohausi
(Arrow) (Trichillum); Pereiraidium almeidai (Pereira) (Pedaridium); Trichillidium brevisetosum (Howden & Young)
(Pedaridium); T. caingua (Martínez) (Pedaridium); T. pilosum (Robinson) (Trichillum); Trichillidium quadridens
(Arrow) (Pedaridium). Distribution maps are presented for the newly described species, for Nunoidium argentinum, for
Pereiraidium almeidai, and for every species of Genieridium.
Key words: dung beetles; Neotropical region; new species; new genus
Resúmen
Se describe a la nueva subtribu Scatimina (parte de la tribu Ateuchini), que consiste de los siguientes 17
géneros, de los cuales 12 son nuevos: Scatimus Erichson; Scatrichus Génier & Kohlmann, 2003; Trichillum
Harold, 1868; Pedaridium Harold, 1868; Eutrichillum Martínez, 1969, n. status; Besourenga, n. g. (especie-
tipo Trichillum minutum Saylor); Bradypodidium, n. g. (especie-tipo Trichillum bradyporum Boucomont);
Degallieridium, n. g. (especie-tipo Degallieridium lilliputanum, n. sp.); Feeridium, n. g. (especie-tipo Feerid-
ium woodruffi, n. sp.); Genieridium, n. g. (especie-tipo Pedaridium bidens Balthasar); Leotrichillum, n. g.
(especie-tipo Pedaridium louzadaorum Vaz-de-Mello & Canhedo); Martinezidium, n. g. (especie-tipo Pedar-
Zootaxa 1955 © 2008 Magnolia Press · 5
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
idium galileoae Génier & Vaz-de-Mello); Nunoidium, n. g. (especie-tipo Pedaridium argentinum Arrow);
Onoreidium, n. g. (especie-tipo Trichillum cristatum Arrow); Pereiraidium, n. g. (especie-tipo Pedaridium
almeidai Pereira); Silvinha, n. g. (especie-tipo Silvinha unica, n. sp.); y Trichillidium, n. g. (especie-tipo
Pedaridium quadridens Arrow). La subtribu Ateuchina Laporte es diagnosticada y incluye a Ateuchus Weber,
Deltorhinum Harold, Aphengium Harold y Sinapisoma Boucomont (transferido desde Canthonini). Se transfi-
ere a los géneros Scatonomus Erichson, Anomiopus Westwood y Hypocanthidium Balthasar de Ateuchini a
Canthonini, y los géneros Canthidium Erichson, Parachorius Harold (Ateuchini) y Holocanthon Martínez &
Pereira (Canthonini) a Coprini. Los géneros Bdelyropsis Pereira, Vulcano & Martínez, Bdelyrus Harold, Cop-
torhina Hope, Delopleurus Erichson, Demarziella Balthasar, Onychothecus Boucomont, Paraphytus Harold,
Pedaria Laporte, Pleronyx Lansberge, Pseuduroxys Balthasar, Sarophorus Erichson y Uroxys Westwood se
consideran como incerta sedis en Ateuchini. Se presenta una clave para los géneros de Scatimina, y otra para
las especies del género Genieridium, que son también diagnosticadas. Se presentan diagnosis para cada
género de Scatimina, así como un listado de las especies incluidas, con observaciones sobre afinidades, com-
posición y distribución. Se diagnostica a cada género de Scatimina y sus especies ya descritas son enlistadas,
con observaciones sobre afinidades, composición y distribución. Todos los géneros, con la excepción de Sca-
timus y Scatrichus, son (re)descritos, y se enlista todo el material examinado de cada especie ya descrita. Se
describen a las siguientes tres nuevas especies: Degallieridium lilliputanum, n. sp.; Feeridium woodruffi, n.
sp. y Silvinha unica, n. sp. Se establecen las siguientes 32 nuevas combinaciones (género original entre
paréntesis): Besourenga minutus (Saylor) (Trichillum); B. vejdovskyi (Balthasar) (Trichillum); B. amarillai
(Aguilar) (Pedaridium); B. horacioi (Martínez) (Trichillum); Bradypodidium adisi (Ratcliffe) (Trichillum); B.
bradyporum (Boucomont) (Trichillum); B. venezuelense (Ferreira & Galileo) (Pedaridium); Eutrichillum
arcus (Solís & Kohlmann) (Trichillum); E. hirsutum (Boucomont) (Trichillum); E. hystrix (Arrow) (Trichil-
lum); Genieridium bidens (Balthasar) (Pedaridium); G. bordoni (Martínez) (Pedaridium); G. cryptops (Arrow)
(Pedaridium); G. margareteae (Génier & Vaz-de-Mello) (Pedaridium); G. med i n a e (Gill & Vaz-de-Mello)
(Pedaridium); G. paranense (Arrow) (Pedaridium); G. zanunciorum (Vaz-de-Mello & Canhedo) (Pedarid-
ium); Leotrichillum louzadaorum (Vaz-de-Mello & Canhedo) (Pedaridium); Martinezidium fulgens (Arrow)
(Pedaridium); M. galileoae (Génier & Vaz-de-Mello) (Pedaridium); M. martinsi (Ferreira & Galileo) (Pedar-
idium); M. maya (Vaz-de-Mello, Halffter & Halffter) (Pedaridium); Nunoidium argentinum (Arrow) (Pedar-
idium); Onoreidium bottimeri (Howden & Young) (Pedaridium); O. cristatum (Arrow) (Trichillum); O.
howdeni (Ferreira & Galileo) (Pedaridium); O. ohausi (Arrow) (Trichillum); Pereiraidium almeidai (Pereira)
(Pedaridium); Trichillidium brevisetosum (Howden & Young) (Pedaridium); T. caingua (Martínez) (Pedarid-
ium); T. pilosum (Robinson) (Trichillum); Trichillidium quadridens (Arrow) (Pedaridium). Se presentan
mapas de distribución para las especies nuevas, para Nunoidium argentinum, para Pereiraidium almeidai, y
para cada especie de Genieridium.
Resumo
A nova subtribo Scatimina (parte da tribo Ateuchini) é descrita, e inclui os seguintes 17 gêneros, dos quais 11
são novos: Scatimus Erichson; Scatrichus Génier & Kohlmann, 2003; Trichillum Harold, 1868; Pedaridium
Harold, 1868; Eutrichillum Martínez, 1969, n. status; Besourenga, n. g. (espécie-tipo Trichillum minutum
Saylor); Bradypodidium, n. g. (espécie-tipo Trichillum bradyporum Boucomont); Degallieridium, n. g.
(espécie-tipo Degallieridium lilliputanum, n. sp.); Feeridium, n. g. (espécie-tipo Feeridium woodruffi, n. sp.);
Genieridium, n. g. (espécie-tipo Pedaridium bidens Balthasar); Leotrichillum, n. g. (espécie-tipo Pedaridium
louzadaorum Vaz-de-Mello & Canhedo); Martinezidium, n. g. (espécie-tipo Pedaridium galileoae Génier &
Vaz-de-Mello); Nunoidium, n. g. (espécie-tipo Pedaridium argentinum Arrow); Onoreidium, n. g. (espécie-
tipo Trichillum cristatum Arrow); Pereiraidium, n. g. (espécie-tipo Pedaridium almeidai Pereira); Silvinha, n.
VAZ-DE-MELLO
6 · Zootaxa 1955 © 2008
Magnolia Press
g. (espécie-tipo Silvinha unica, n. sp.); y Trichillidium, n. g. (espécie-tipo Pedaridium quadridens Arrow). A
subtribo Ateuchina Laporte é diagnosticada e inclui Ateuchus Weber, Deltorhinum Harold, Aphengium Harold
e Sinapisoma Boucomont (transferido de Canthonini). Os gêneros Scatonomus Erichson, Anomiopus West-
wood y Hypocanthidium Balthasar são transferidos de Ateuchini a Canthonini, e os gêneros Canthidium
Erichson, Parachorius Harold (Ateuchini) e Holocanthon Martínez & Pereira (Canthonini) a Coprini. Os
gêneros Bdelyropsis Pereira, Vulcano & Martínez, Bdelyrus Harold, Coptorhina Hope, Delopleurus Erichson,
Demarziella Balthasar, Onychothecus Boucomont, Paraphytus Harold, Pedaria Laporte, Pleronyx Lansberge,
Pseuduroxys Balthasar, Sarophorus Erichson e Uroxys Westwood são considerados incerta sedis em Ateu-
chini. É apresentada uma chave para os gêneros de Scatimina, e outra para as espécies de Genieridium, que
são também diagnosticadas. Cada gênero de Scatimina é diagnosticado e é acompanhado por uma lista de suas
espécies já descritas, com observações sobre afinidades, composição e distribuição. Todos os gêneros, exceto
Scatimus e Scatrichus, são (re)descritos, e todo o material examinado das espécies já descritas é listado. As
seguintes três novas espécies são descritas: Degallieridium lilliputanum, n. sp.; Feeridium woodruffi, n. sp. e
Silvinha unica, n. sp. As seguintes 32 novas combinações são estabelecidas: (gênero original entre parênte-
ses): Besourenga minutus (Saylor) (Trichillum); B. vejdovskyi (Balthasar) (Trichillum); B. amarillai (Aguilar)
(Pedaridium); B. horacioi (Martínez) (Trichillum); Bradypodidium adisi (Ratcliffe) (Trichillum); B. brady-
porum (Boucomont) (Trichillum); B. venezuelense (Ferreira & Galileo) (Pedaridium); Eutrichillum arcus
(Solís & Kohlmann) (Trichillum); E. hirsutum (Boucomont) (Trichillum); E. hystrix (Arrow) (Trichillum);
Genieridium bidens (Balthasar) (Pedaridium); G. bordoni (Martínez) (Pedaridium); G. cr ypt o ps (Arrow)
(Pedaridium); G. margareteae (Génier & Vaz-de-Mello) (Pedaridium); G. med i n a e (Gill & Vaz-de-Mello)
(Pedaridium); G. paranense (Arrow) (Pedaridium); G. zanunciorum (Vaz-de-Mello & Canhedo) (Pedarid-
ium); Leotrichillum louzadaorum (Vaz-de-Mello & Canhedo) (Pedaridium); Martinezidium fulgens (Arrow)
(Pedaridium); M. galileoae (Génier & Vaz-de-Mello) (Pedaridium); M. martinsi (Ferreira & Galileo) (Pedar-
idium); M. maya (Vaz-de-Mello, Halffter & Halffter) (Pedaridium); Nunoidium argentinum (Arrow) (Pedar-
idium); Onoreidium bottimeri (Howden & Young) (Pedaridium); O. cristatum (Arrow) (Trichillum); O.
howdeni (Ferreira & Galileo) (Pedaridium); O. ohausi (Arrow) (Trichillum); Pereiraidium almeidai (Pereira)
(Pedaridium); Trichillidium brevisetosum (Howden & Young) (Pedaridium); T. caingua (Martínez) (Pedarid-
ium); T. pilosum (Robinson) (Trichillum); Trichillidium quadridens (Arrow) (Pedaridium). Mapas de distri-
buição são apresentados para as espécies novas, para Nunoidium argentinum, para Pereiraidium almeidai, e
para cada espécie de Genieridium.
Introduction
The tribe Ateuchini, as defined by Montreuil (1998), includes most genera of Scarabaeinae with apically
expanded mesotibiae and metatibiae not assignable to the other tribes with similarly expanded tibiae: namely
Coprini, Phanaeini, Oniticellini, Onitini and Onthophagini. All genera presently included in Ateuchini are
tropical with a few species distributed north to the Nearctic Region, and the tribe is presently defined mainly
on the absence of the defining characters of the other paracoprid (tunneler) tribes. In other words, Ateuchini as
presently recognized, lacks a unique set of defining characters (synapomorphies).
The purpose of this work is to define a new subtribe for some Neotropical genera now included in Ateu-
chini. It is a contribution toward an ongoing study of the suprageneric classification of the subfamily by
François Génier and me.
History of the genus Pedaridium Harold and relatives
The genus Pedaria was erected by Laporte (1832), for a single African species, P. nigra, from Senegal. In
1859, Harold described the first non-African species of this genus, Pedaria hirsuta, based on a number of
Zootaxa 1955 © 2008 Magnolia Press · 7
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
specimens from Brazil from his own and Sturm’s collections.
In 1868, Harold erected a new scarab genus, Trichillum, including in it, T. heydeni Harold, and established
in his key the genus Pedaridium for the New World species of Pedaria. He distinguished it from Trichillum by
the shorter basal metatarsomere and unexpanded epipleuron. He did not explicitly designate a type species for
Pedaridium; however, as only Pedaria hirsuta was described from the New World, it is considered a designa-
tion by monotypy. Harold (1869a) first published the combination Pedaridium hirsutum, and Preudhomme de
Borre (1880) described a second species of Trichillum.
Arrow (1913) described three more species of Pedaridium, Boucomont (1928) described two more spe-
cies of Trichillum. Arrow (1931) revised the genus Trichillum and described three new species, not mention-
ing species described in 1928 by Boucomont. One year later Arrow (1932) described two more species of
Pedaridium, gave an updated key for the genus Pedaridium, and made comments about Boucomont’s (1928)
descriptions of Trichillum. Saylor (1935) described three more species of Trichillum.
Paulian (1936) described the homonymous T. arrowi based on a single specimen from Pará, Brazil, giving
a new key for the species included in this genus, except for those of Saylor (1935). Balthasar (1938) described
one more species of Pedaridium and presented a key for all species of Pedaridium known at that time. The
same author (Balthasar 1939a) reviewed the genus Trichillum and described two new species, and later (Bal-
thasar 1942) one more species of Trichillum. Pereira (1946) described another species of Pedaridium. Robin-
son (1948) described one more Trichillum. Martínez (1951) described another Pedaridium.
Martínez (1969) erected the subgenus Eutrichillum within Trichillum, transferred T. bradyporum, T.
ohausi and T. elongatum to Pedaridium, hypothesized that T. hirsutum could also be a Pedaridium, and con-
sidered T. hirsutum and T. cristatum as incertae sedis. In the same paper the author reviewed the subgenus Tri-
chillum, describing three new species in this subgenus and one new species without a subgeneric designation.
The same author (Martínez 1974) described “Pedaridium (?) caingua,” and stated that this new species,
together with P. quadridens, should be included in a new genus under study by him.
Ratcliffe (1980) described adults, larvae, and pupae of a new species of Trichillum; gave keys for the sub-
genera of Trichillum and for species in the subgenus Eutrichillum (including T. hirsutum Boucomont). The
same author (Ratcliffe 1981) redescribed the type of Trichillum hirsutum.
Howden & Young (1981) described two species of Pedaridium from Panama, assigned Trichillum pilosus
Robinson to Pedaridium based on characters used by Paulian (1936) and Martínez (1969) to distinguish these
genera, and commented on the confusion regarding distinguishing characters between Trichillum and Pedar-
idium and the need of comprehensive revisions of both.
Martínez (1992) described a Venezuelan species of Pedaridium, and in the abstract of this paper, in both
Spanish and English, mentioned an undescribed species from Mexico.
Ferreira & Galileo (1993) revised the genus Pedaridium, with the exception of P. bordoni Martínez, which
was probably unknown to them. In this revision, they transferred one species of Trichillum to Pedaridium,
described six new species, and chose to emphasize the shape of pseudepipleuron (they called this the epipleu-
ron) to distinguish Trichillum and Pedaridium. It is important to point out that the authors did not examine
most of the type specimens, which are in European museums.
Verdú & Galante (1997) described a new species of Trichillum from Uruguay, and gave a new key for spe-
cies in the subgenus Trichillum.
Montreuil (1998), in a paper on phylogenetic relationships between Ateuchini and Coprini, examined
Pedaridium (the type species and possibly some additional unspecified species) and placed it in an unresolved
clade including Demarziella, Pedaria, Sarophorus Erichson, and the clade Bdelyrus Harold + Bdelyropsis
Pereira, Vulcano, & Martínez. This paper did not include Trichillum.
Vaz-de-Mello & Canhedo (1998) described two more Pedaridium species, and included those and P. bor-
doni Martínez in the Ferreira & Galileo (1993) key to species. Aguilar-Julio (2001) described a new Para-
guayan Pedaridium. The author did not mention the revision by Ferreira & Galileo (1993), and stated that the
VAZ-DE-MELLO
8 · Zootaxa 1955 © 2008
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new species had sinuate epipleura (sic), a character that would place the species in Trichillum according to
Ferreira & Galileo (1993).
Verdú & Galante (2001) described larvae and breeding behavior of two species of Pedaridium, and con-
cluded that 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 behavior.
Génier & Vaz-de-Mello (2002) diagnosed and designated lectotypes for those species described by Arrow
in both Trichillum and Pedaridium, synonymizing three previously described species, and described two new
species for species previously misidentified. Solís & Kohlmann (2003) described a species of Trichillum
(Eutrichillum) and Gill & Vaz-de-Mello (2003) and Vaz-de-Mello et al. (2004) described two other Pedarid-
ium species. Vaz-de-Mello (2003) presented a preliminary overview of the entire group, without nomenclato-
rial validity (this work contains a disclaimer), but containing an exhaustive review of literature and
specimens. Finally, Vaz-de-Mello & Génier (2005) synonymized three previously described species in Trichil-
lum and Pedaridium, and described three new previously misidentified species in the genus Trichillum.
Since very early, both Trichillum and Pedaridium have been placed near Ateuchus (for instance by Harold
[1869a] under the name Choeridium). Morphological differences between Pedaridium and Pedaria have
never been discussed in the literature, besides the difference in the geographical distribution.
The differences between Trichillum and Pedaridium have been discussed in many papers. In the original
description of the genera (for Pedaridium, in the key), Harold (1868) stated that they differ by the shape of
“epipleuron” and length of the basal metatarsomere. After that, only the second character was taken into
account for assigning new species to each of those genera (Arrow 1913, 1931, 1932; Balthasar 1939b; Mar-
tínez 1969). Howden & Young (1981) used the unique or double setose puncture rows in the interstriae to dif-
ferentiate genera, resulting in the placement of all Panamanian species in Pedaridium. Ferreira & Galileo
(1993) used the shape of the epipleuron, pointing out that Pereira (1946) had described a Pedaridium with
basal metatarsomere longer than the second metatarsomere. During the 20th century, some species described
in Trichillum have been transferred to Pedaridium: Martínez (1969) transferred three species, Howden &
Young (1981) transferred one species, and Ferreira & Galileo (1993) transferred one species.
Verdú & Galante (2001), citing larval morphology of three species of the genus Pedaridium, as well as
information provided by Vaz-de-Mello in litteris on adult morphology, considered that this genus is probably
a polyphyletic group.
All the confusion regarding generic assignments indicates that the two genera are not both monophyletic
groups, and that a global revision of this group is necessary in order to establish consistent genus-level taxa.
Moreover, the enormous variability in external characters such as dorsal hairs, punctures, legs and clypeal
form, also indicate that other characters must be used in order to reach a more accurate generic system for this
group.
Vaz-de-Mello (2003) proposed the division of this group into 20 separate genera (without formally
describing them) based on an extensive morphological phylogenetic analysis. This classification, with many
modifications based on subsequent unpublished analysis, is very similar to the one adopted here, and this
work is referred as the reference for the phylogenetic support of the new genera proposed in this work.
Until now, the genus Pedaridium has included 28 valid species. The genus Trichillum includes two sub-
genera: the nominotypical subgenus contains ten valid species and Trichillum (Eutrichillum) contains six
valid species. One species of Trichillum is considered as incerta sedis, without subgeneric placement.
The former genus Scatimus Erichson
Erichson (1847) erected the genus for the new species Scatimus cucullatus, which he distinguished from
Choeridium LePeletier and Serville (= Ateuchus Weber) based primarily on the presence (in Scatimus) of car-
inae on the outer edge of the mesotibiae and metatibiae. Other species have been added to the genus by Harold
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SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
(1862, 1869b), Preudhomme de Borre (1886), Balthasar (1939a), Martínez (1988), and Kohlmann & Solís
(1996). The phylogenetic position of this genus has been debated, some considering it to be a close relative of
Ateuchus (Erichson 1848, Lacordaire 1856, Harold 1867, Bates 1887, Balthasar 1939a, Edmonds & Halffter
1978, Halffter & Edmonds 1982) while others place it closer to Copris Geoffroy (Luederwaldt 1931, Pereira
1954) and yet others as a “bridge” between Ateuchus and Copris (Martínez 1988).
Scatimus was first reviewed by Balthasar (1939a). Génier & Kohlmann (2003) split the genus, creating
Scatrichus for S. bicarinatus (Harold) and two new species. In Scatimus, they synonymized two species and
described four as new. Génier & Kohlmann (2003) also proposed the phylogenetic sisterhood between (Scati-
mus + Scatrichus) and Pedaridium.
Material and methods
This study is based on examination of about fifteen thousand specimens belonging to more than one hundred
species (many yet undescribed) from the following collections (institutional curators’ names in parenthesis):
ABC Alberto Ballerio personal collection, Brescia, Italy.
AMBC Ayr M. Bello personal collection, Rio de Janeiro, RJ, Brazil.
BDGC Bruce D. Gill personal collection, Woodlawn, Ontario, Canada.
BMNH The Natural History Museum, London, England (Malcolm Kerley).
BRBA Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires (A. Bachmann).
CAMC Claudia A. Medina personal collection, Cali, Colombia.
CMNC Canadian Museum of Nature, Ottawa, Canada (Henry Howden and François Génier).
CNCI Canadian National Collection of Insects, Ottawa, Canada (Anthony Davies and Patrice Bou-
chard).
EMOC Enrique Montes de Oca personal collection, Xalapa, Mexico.
ESAP Escola Superior de Agricultura “Luiz de Queirós”, Universidade de São Paulo, Piracicaba, SP,
Brazil (Roberto A. Zucchi).
FEIS Faculdade de Engenharia de Ilha Solteira da Universidade Estadual Paulista, Ilha Solteira, SP,
Brazil (Carlos A. H. Flechtmann).
FMLT Fundación Miguel Lillo, San Miguel de Tucumán, Argentina (A. Terán).
FMNH Field Museum of Natural History, Chicago, IL, USA (Alfred Newton)
FVMC Fernando Vaz-de-Mello collection, Universidade Federal de Mato Grosso, Departamento de Bio-
logia e Zoologia, Cuiaba, MT, Brazil (Fernando Vaz-de-Mello)
FZRS Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre, RS, Brazil (Maria Helena M. Gali-
leo).
GVHC Gonzalo and Violeta Halffter personal collection, Coatepec, México.
IAHC Instituto Alexander von Humboldt, Villa de Leyva, Colombia (Fernando Fernández).
IBSP Coleção Entomológica “Adolph Hempel”, Instituto Biológico, São Paulo, SP, Brazil (Sergio Ide).
IEX Instituto de Ecología, A.C., Xalapa, México (Miguel Ángel Morón).
IRSN Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium (Marcel Cludts).
LDC Leonardo Delgado personal collection, México, DF, México.
LEMQ Lyman Entomology Museum, McGill University, Ste. Anne de Bellevue, Canada.
MAMC Miguel A. Morón personal collection, Xalapa, México.
MAPA Museu Anchieta, Porto Alegre, RS, Brazil (Fernando R. Meyer).
MEUA Museo Entomológico, Universitad d’Alicant, Spain (José Verdú and Eduardo Galante).
MNHN Muséum National d’Histoire Naturelle, Paris (Yves Cambefort and Olivier Montreuil).
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MNRJ Museu Nacional da Universidade Federal do Rio de Janeiro, RJ, Brazil (Miguel A. Monné and
Paulo R. Magno).
MTMB Magyar Természettudomány Muzéum, Budapest, Hungary (Otto Merkl).
MZSP Museu de Zoologia da Universidade de São Paulo, SP, Brazil (Ubirajara R. Martins, Cleide
Costa, Carlos Campaner, Sergio Ide, and Sonia Casari).
NHMB Naturhistorisches Museum, Basel, Switzerland (Eva Sprecher).
NMM Natuurhistorisch Museum Maastricht, The Netherlands (F. Dingemans-Backels and Alexey
Tishechkin).
NMP Narodní Muzeum, Prague (Josef Jelínek).
PSC Paul Schoolmeesters private collection, Herent, Belgium.
PUCE Pontifícia Universidad Católica del Ecuador, Quito, Ecuador (Giovanni Onore and Carlos Car-
pio).
UNSM University of Nebraska State Museum, Lincoln, NE, USA (Brett Ratcliffe and Mary Liz Jame-
son).
URRJ Universidade Federal Rural do Rio de Janeiro, Seropédica, RJ, Brazil (Francisco Racca Filho and
Paschoal Grossi)
USNM United States National Museum, Washington, USA (Nancy Adams)
UVG Universidad del Valle de Guatemala (Enio Cano and Jack Schuster).
WDEC W.D. Edmonds personal collection, Marfa, TX, USA.
ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany (Hella Wendt).
Species already described (except for Scatimus and Scatrichus, treated by Génier & Kohlmann [2003])
and those described in the present paper have a mention of the material examined up to date, which will be
referred in subsequent papers.
Biogeographical divisions are according to Morrone (2006).
Scanning Electron Microscope photos have been taken with a LEO variable pressure scanning electron
microscope, at the Núcleo de Apoio à Pesquisa em Microscopia Eletrônica Aplicada à Agricultura, at Escola
Superior de Agricultura Luiz de Queirós, Universidade de São Paulo, Piracicaba, Brazil.
Scatimina, new subtribe
Diagnosis: The subtribe Scatimina can be separated readily from all remaining Scarabaeinae by the following
combination of characters: (1) hypomeron anteriorly deeply excavated, excavation bordered posteriorly by
transverse carina; (2) trochantofemoral pit present; and (3) pseudoepipleuron present lateral to 8th elytral
stria, delimited by folding or convexity of 9th interstria, and separated from epipleuron by stria distinctly visi-
ble at least basally (Figs. 5–11).
Description: Clypeus centrally with an emargination, two or more teeth, or both, never simply rounded;
external genal angle clearly obtuse; clypeal ventral process forming transverse carina, sometimes angled or
tuberculated in middle, united to transversal supraepipharyngeal ridge by angle or short longitudinal carina.
Separation between submentum and gula simply curved to obtusely V-shaped, sometimes indistinct, gula lat-
erally glabrous, and anteriorly slightly pointed. Antennae with 9 antennomeres, first antennomere of club
completely tomentose. Mentum apically not emarginated, or only slightly so, with medial dorsal, pointed pro-
longation (serving as "base" for membranous dorsally-positioned ligulae). Labial palpi with 3 palpomeres;
third much narrower, smaller than second. Side of pronotal disc with indistinct to distinct callosity, without
longitudinal sulcus or distinct fovea; centrally without elevations, sulci, or tubercles, simply convex. Pronotal
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SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
osterior border unbeaded or very feebly beaded. Hypomeron anteriorly deeply excavated, excavation sepa-
rated from posterior part by transverse carina. Mesosternum anteriorly with transverse sulcus. Metasternum
anteriorly with transverse sulcus parallel and very close to (often coincident with) middle of mesometasternal
suture, prolonged parallel to mesocoxa laterally.
Protibial apices mesally truncated, straight (except for mesal apical tooth in some males); protibiae apical
border straight, continuous (i. e., without angle) with anterior border of apical tooth, such that two margins
(protibial body and apical tooth) lie in straight line (Figs. 1–4). Protibiae with three teeth or less, with tarsal
insertion near the mesal apical border. Protrochantofemoral pit present. Mesocoxal axis very slightly angled,
almost parallel to longitudinal body axis. Mesotibial apex obliquely truncated (mesal angle acute). Metatibial
apex entire below tarsal insertion. Mesotibiae and metatibiae with longitudinal external carina indicated on
disc, and dorso-external longitudinal carina well defined, while ventro-external carina is absent or indicated
by row of setae. Mesotibial and metatibial apical fimbriae alternate in size and thickness, with short, thick
setae co-mingled with long, narrow setae (sparse short, thick setae also present on the external surface of
mesotibiae and metatibiae). Elytra with nine striae. Pseudoepipleura present lateral to 8th elytral stria, delim-
ited by fold or convexity of 9th interstria and separated from epipleuron by epipleural stria well defined at
least in basal half (Figs. 5–11).
Sixth abdominal ventrite never shortened medially. Basal pygidial sulcus present. Male genitalia with
genital segment transverse, conformed of a central rounded plate and a lateral rectangular elongated plate at
each side. Parameres almost symmetrical (at least in size and shape of sides). Transverse parietal lamella of
internal sac (magnifying glass-shaped lamella or circular sclerite of authors) with basal arc very reduced, so
that one side of the circle is obliterated or completely absent. Female genitalia with coxites present, some-
times fused ventrally. Spermathecal base simply rounded or pointed, never bilobed or expanded; apex simply
pointed.
Type genus: Scatimus Erichson, 1847
Remarks. This new subtribe includes the following Neotropical genera: Besourenga, new genus; Brady-
podidium, new genus; Degallieridium, new genus; Eutrichillum Martínez, new status; Feeridium, new
genus; Genieridium, new genus; Leotrichillum, new genus; Martinezidium, new genus; Nunoidium, new
genus; Onoreidium, new genus; Pedaridium Harold; Pereiraidium, new genus; Scatimus Erichson; Scatri-
chus Génier & Kohlmann; Silvinha, new genus; and Trichillidium, new genus; Trichillum Harold.
Besides the characters mentioned above, the following are putative apomorphies supporting Scatimina:
the presence of a pit in the external face of first and second antennal lamellae (these pits appear to have been
secondarily lost in Scatrichus), the presence of an anterior transverse carina on mesoepimeron (lost in the
group formed by Trichillum, Besourenga, Degallieridium, Feeridium, Eutrichillum, and in most species of
Scatimus), the spatulate form of the protibial spur in males (modified in one species of Trichillum and at least
some species of Scatimus), the sinuate form of mesotibial spurs, the absence of a basal raspula and the pres-
ence of an elongated flagelliform sclerite in the internal sac.
The Scatimina are distributed in the entire Neotropical region as defined by Morrone (2006) (except for
the Antillean Dominion), and in the entire Mexican Transition Zone, in a province of the Nearctic Region
(Sonora), and in part of the South American Transition Zone (Monte and perhaps Prepuna provinces).
The trochanterofemoral pit is not unique to the new subtribe and is also present in the following four gen-
era of Ateuchini: Bdelyropsis Pereira, Vulcano & Martínez, Demarziella Balthasar, Pedaria Laporte, and
Uroxys Westwood, and several genera of Canthonini. These genera were previously considered related to
members of the Scatimina, but are herein viewed as Ateuchini incertae sedis or transferred to Canthonini (see
below).
Although Pedaria and Demarziella have been regarded as relatives of Trichillum and Pedaridium (based
on the presence of dorsal setae and fused abdominal sternites), they show the following important differences:
(1) elytra with ten striae (as opposed to nine in the Scatimina – the ninth and tenth striae are often partially
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fused in the middle, but well differentiated both anteriorly and posteriorly); (2) parameres distinctly asymmet-
rical; and (3) presence of well-developed magnifying glass shaped lamella in the internal sac. These charac-
ters of Pedaria and Demarziella are shared with Coptodactyla Burmeister, Thyregis Blackburn, and
Microcopris Balthasar, formally belonging to the tribe Coprini, which lack fused abdominal sternites. These
genera could be regarded as a separate tribe (Coptodactylini Paulian), but that is beyond the scope of the
present paper.
It is tempting to include the genus Uroxys in Scatimina, but the heterogeneity, presumed polyphyletic ori-
gin of the genus (Halffter & Matthews 1966, Halffter & Edmonds 1982, Martínez 1988), and various taxo-
nomic and nomenclatorial difficulties make it inadvisable at the present time. One difficulty is the uncertain
position of the type species of Uroxys. Some species presently included in Uroxys are here excluded from Sca-
timina because of the presence of lateral longitudinal sulci on each side of the pronotum and the complete
magnifying glass shaped sclerite in the internal sac. However, it seems that at least one of the species groups
now placed in Uroxys belongs or is very close to Scatimina.
Finally, Bdelyropsis has the distinct pseudoepipleuron originating from a fold in the 8th interstria, and
appearing related to a group of genera currently in the Canthonini that includes Zonocopris Arrow and Cryp-
tocanthon Balthasar, among others (Vaz-de-Mello 2007, see below on Bdelyrus).
A new diagnosis for the tribe Ateuchini greatly depends on the reassignment of the genera here consid-
ered as incertae sedis to other tribes and will be presented in a subsequent paper. The proposal of a new sub-
tribe in Ateuchini results in the recognition of the subtribe Ateuchina, which is provisionally diagnosed as
follows: (1) medial part of clypeal surface concave; (2) clypeal ventral process in the form of a transverse car-
ina; (3) pronotum simply convex, at most with discal longitudinal sulcus; (4) lateral pronotal pit well-defined;
(5) hypomeron deeply excavated anteriorly and with a transverse carina (as in Scatimina); (6) fore legs with
trochantofemoral pit absent; (7) protibia with anterior border completely straight (as in Scatimina), except for
anteroapical tooth in male; (8) nine elytral striae (with pseudoepipleuron delimited by a fold or convexity in
the 9th interstria); (9) parameres symmetrical; (10) internal sac sclerites represented by few small flagelliform
lamellae, all apical in position.
As defined herein, the subtribe Ateuchina is Neotropical and Nearctic and includes the following genera:
Ateuchus Weber (with the exclusion of some species to be assigned to other genera in the near future), Del-
torhinum Harold, Aphengium Harold, and Sinapisoma Boucomont (formerly placed in Canthonini), as well as
taxa yet to be described.
Most other ateuchine genera (all lacking trochantofemoral pit) are herein considered to be Ateuchini
incertae sedis.
The genera Canthidium Erichson (based on examination of the type species, Canthidium lentum Erichson)
and Parachorius Harold have 10 elytral striae, the 9th being easily discernible from the 8th only at elytral apex,
and both have hypomera very feebly excavated anteriorly, although the transverse carina is present; they also
show an angle between the anterior borders of the apical protibial teeth and the mesal apical margin of tibia
(i.e., entire anterior tibial border is not straight). Those characters point to a close relationship between these
genera and Holocanthon Martínez & Pereira (presently placed in Canthonini), and justify their inclusion in the
tribe Coprini. Within the Coprini, the most closely related group would be the one including Dichotomius
Hope and its allies. The subgenus Eucanthidium Martínez & Halffter (based on examination of the type spe-
cies, Canthidium cupreum Blanchard), shares all those characters except that only nine elytral striae are
present; Eucanthidium may later turn out to be a taxonomic isolate.
Bdelyrus Harold, has a double pseudoepipleuron originated from folds in both 8th and 9th interstriae, lacks
the developed trochantofemoral pit, and has a very peculiar assemblage of internal sac sclerites. However, its
unusual abdominal shape is similar to that of Paracryptocanthon, and the supraunguicular spines (found oth-
erwise only in Zonocopris) appears to relate Bdelyrus to the yet unnamed group that includes those genera
(Vaz-de-Mello 2007, see above also on Bdelyropsis). Furthermore, internal sac characters appear to relate
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SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Bdelyrus to Onychothecus Boucomont and Paraphytus Harold, both possessing ten elytral striae.
Scatonomus Erichson and Anomiopus Westwood (and the related Hypocanthidium Balthasar) appear to be
true Canthonini, in fact closely related to Canthon. This relationship was suggested by Canhedo (2006), who
did not make any formal classification changes, and I here assign Scatonomus and Anomiopus to the tribe
Canthonini.
Finally, Sarophorus Erichson, Coptorhina Hope, Frankenbergerius Balthasar, and Delopleurus Erichson
have a unique lateral sinuosity on the elytra; a longitudinal groove on the gula; a very simple internal sac, in
some cases lacking sclerites; and a very distinctive spermatheca with flat basal paired lobes found otherwise
only in some Canthonini. These genera should be considered as a group distinct from Ateuchini (Zunino
1983, Frolov & Scholtz 2003, Frolov & Scholtz 2005).
A summary of the present considerations on subtribal or tribal position of above-mentioned is given on
Table 1.
TABLE 1. Tribal and subtribal position of genera formerly or here included in the Ateuchini and some close relatives, as
proposed here. An * indicates genera not examined.
The present necessity to split into 15 two previously very heterogenous genera, Trichillum and Pedarid-
ium, is yet another consequence of the tremendous increase in data and material during the remarkable surge
in field and museum work on dung beetles during the past 30 or so years. The number of new species known
to me in these two groups alone has increased by about 80 (200%) in the last decade; while some are
described here, many will be described in subsequent papers. Likewise, in this paper and elsewhere, the recent
increase in the number of new taxonomic characters has permitted taxonomic and cladistic studies of higher
resolution. Other recent cases of extensive splitting of previously heterogeneous genera are those of Panelus
Lewis (Scholtz & Howden 1987; Ochi et al. 1998; Paulian 1976, 1985; Matthews 1974), Epilissus Reiche
Ateuchini
Ateuchina Scatimina Incertae sedis
Aphengium Harold, 1868 Besourenga, new genus Bdelyropsis Pereira, Vulcano & Mar-
tínez, 1960
Ateuchus Weber, 1801 Bradypodidium, new genus Bdelyrus Harold, 1869
Deltorhinum Harold, 1867 Degallieridium, new genus Coptorhina Hope, 1830
Sinapisoma Boucomont, 1928 Eutrichillum Martínez, 1969, new status Delopleurus Erichson, 1847
Feeridium, new genus Demarziella Balthasar, 1961
Genieridium, new genus Onychothecus Boucomont, 1912
Leotrichillum, new genus Paraphytus Harold, 1877
Martinezidium, new genus Pedaria Laporte, 1832
Nunoidium, new genus *Pleronyx Lansberge, 1874
Onoreidium, new genus *Pseuduroxys Balthasar, 1938
Pedaridium Harold, 1868 Sarophorus Erichson, 1847
Pereiraidium, new genus Uroxys Westwood, 1842
Scatimus Erichson, 1847
Scatrichus Génier & Kohlmann, 2003
Transferred to Canthonini Silvinha, new genus Transferred to Coprini
Anomiopus Westwood, 1842 Trichillidium, new genus Canthidium Erichson, 1847
Hypocanthidium Balthasar, 1938 Trichillum Harold, 1868 Holocanthon Martínez & Pereira, 1956
Scatonomus Erichson, 1835 Parachorius Harold, 1875
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(Olsoufieff 1935, 1947; Paulian 1939), and the “group” Stiptopodius Harold (Branco 1989, 1990, 1991,
1992a, 1992b, 1995). My personal opinion is that, if these genera (including those treated here) were larger in
size and more abundant in collections (it must be said that most collections are very poor in representatives of
the diversity of those groups, most having a very small number of species of few subgroups of it), then they
would have been split (or independently described one by one) much earlier.
Key to the genera of Scatimina (partly taken from Génier & Kohlmann [2003])
1. Length of last abdominal sternite (measured along midline) about equal to lengths of other sternites or
not longer than two of them combined................................................................................................... 2
-. Last abdominal sternite elongate (measured along midline), extending from pygidium to metacoxae,
remaining sternites only discernible laterally......................................................................................... 3
2(1). Head with two parallel transverse carinae; lateral edge of pronotum crenulate, with setose punctures;
apical declivity of elytron with setiferous punctures on interstriae; trochantofemoral pit of anterior leg
oval, transverse; lateral declivity of pronotum with accessory setiferous punctures. Southeastern and
central Brazil ..........................................................................14. Scatrichus Génier & Kohlmann, 2003
-. Head with one transverse carina, or with one conical tubercle; lateral edge of pronotum simple, lacking
setose punctures; apical declivity of elytron lacking setiferous punctures on interstriae; trochantofemo-
ral pit of anterior leg rounded; lateral declivity of pronotum lacking setiferous punctures. Northwestern
Mexico to northern South America ........................................................... 13. Scatimus Erichson, 1847
3(1). Elytral pseudoepipleuron forming two sinuosities, the posterior one (near metacoxa) partially covering
true epipleuron, and sometimes angled (Figs. 5, 9) .............................................................................. 4
- Elytral pseudoepipleuron at most forming one long sinuosity on anterior half that does not fold over
true epipleuron (sometimes true epipleuron with excavation near metacoxa) (Figs. 6–8, 10–11) ........5
4(3). Clypeal teeth clearly arising below dorsal clypeal margin (Fig. 91); pseudoepipleuron without anterior
longitudinal carina on basal half; posterior pseudoepipleuron sinuosity bent over anteriorly, posterior
half of pseudoepipleuron not distinguishable from true epipleuron (i.e., epipleural stria indistinct on
posterior half) (Fig. 9); male with apical tarsomere of anterior tarsi excavated dorsally to receive
claws; parameres strongly divergent in the middle and convergent apically (Fig. 92). Southeastern Bra-
zil ....................................................................................................................... 15. Silvinha new genus
-. Clypeal teeth continuous with dorsal clypeal margin (Figs. 98, 100, 102); pseudoepipleuron with a
sharp anterior longitudinal carina extending up to posterior pseudoepipleuron sinuosity, which is bent
over true epipleuron and angled, posterior half of pseudoepipleuron completely distinguishable poste-
riorly from true epipleura by epipleural stria (Fig. 5); male with apical tarsomere of anterior tarsi not
modified; parameres flattened, truncated apically, without external lobes. South America east of the
Andes excluding southern Argentina ......................................................... 17. Trichillum Harold, 1868
5(3). Pseudoepipleuron posteriorly abruptly narrowed near metacoxa, margin forming an angle (Fig. 10). 6
-. Pseudoepipleuron gradually reduced in width, margin straight near metacoxa (Figs. 6, 8, 11)............ 7
6(5). Clypeo-genal suture clearly visible, extending from frontoclypeal suture to clypeogenal border; cly-
peo-genal border incised, with clypeus and gena separately rounded, clypeus evenly rounded between
clypeal teeth and clypeo-genal suture (Fig. 49). Northern Argentina, central and southeastern Brazil,
Paraguay, eastern Bolivia, Amazonia (except north), one isolated species in Costa Rica.......................
................................................................................................................ 4. Eutrichillum Martínez, 1969
-. Clypeogenal and frontoclypeal sutures indistinct; clypeal margin lateral to clypeal teeth strongly
curved or angled, becoming straight anteriorly to clypeogenal border, itself straight or slightly sinuate
(Figs. 39, 41). Paraguay, eastern Bolivia, central Brazil, and western Amazonia ...................................
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SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
........................................................................................................................ 1. Besourenga new genus
7(5). Head with two horns (males) (Fig. 88) or two feeble separate convexities (females) (Fig. 90) on fronto-
clypeal region; anterior margin of pronotum beaded. Southern Brazil ... 12. Pereiraidium, new genus
-. Head without horns or tubercles on frontoclypeal region; anterior margin of pronotum unbeaded .... 8
8(7). Protibia with two or three lateral teeth, distributed along one-half or less of tibial length; mesotibia not
strongly widened apically, apicolateral area ventrally covered by very long setae............................... 9
-. Protibia with three lateral teeth, distributed along at least apical three-fifths of tibial length, if occupy-
ing less than three-fifths of tibial length, then, mesotibia with strong lateroventral tooth and apex
abruptly extended laterally, and with sparse apical ventral setae ....................................................... 10
9(8). Head flat to very feebly evenly convex, lacking evident shallow concavities in front of eyes; clypeus
laterally straight to slightly curved outward; gena not protruding beyond clypeus (Figs. 43, 45); body
elongated (Fig. 17); elytral striae non-moniliform (strial punctures small and well separated); claws
and apical tarsomere of protarsi of males strongly modified (Fig. 46). Costa Rica to southeastern Brazil
..................................................................................................................2. Bradypodidium, new genus
-. Head evenly convex mesally, with distinct shallow concavities in front of eyes; clypeal margin later-
ally curved mesad; gena protruding beyond clypeus (Figs. 93, 95, 97); body strongly rounded (Figs.
35–36); elytral striae moniliform at least on apical half (strial punctures twice the width of stria, punc-
tures weakly separated or contiguous in posterior half); claws and tarsomeres of male not modified.
Nicaragua to Ecuador; Brazil, Argentina, Paraguay, Bolivia ................... 16. Trichillidium, new genus
10(8). Sides of pronotum, in dorsal view, sinuous (Figs. 29, 87); elytra distinctly tectiform, sutural (first)
interstriae distinctly elevated, elytral surface flat (Fig. 29). Southern Brazil...........................................
.................................................................................................................. 11. Pedaridium Harold, 1868
-. Sides of pronotum, in dorsal view, approximately straight to uniformly curved; elytra at most slightly
tectiform; sutural (first) interstria not distinctly elevated; elytral surface evenly convex ................... 11
11(10). Eyes dorsally as wide as long; dorsal interocular distance less than two times width of one eye (Figs.
20, 52); pronotum separated from hypomeron by row of punctures, not by carina. Amazon River bor-
der and French Guyana ................................................................................... 5. Feeridium, new genus
-. Eyes dorsally longer than wide; dorsal interocular distance at least five times width of a single eye;
pronotum separated of hypomeron by sharp longitudinal carina or at least weak carina between punc-
tures (Fig. 59) ...................................................................................................................................... 12
12(11). Head disc strongly convex, sometimes with a transverse carina; clypeal emargination indistinct; cly-
peal teeth upturned and widely separated, area between teeth shallow, very widely U-shaped (Figs. 79,
82–83); head central surface strongly convex, sometimes with a transverse carina; mesotibiae (and
metatibiae to lesser degree), in ventral view, with strong tooth near mid-length of lateral border, tooth
bearing a very thick and short seta (Fig. 81); mesotibiae with ventral transverse ventral apical carina of
the same tibia)with stout setae similar to one on ventral lateral tooth. Ecuador, Panama, and Costa Rica
.................................................................................................................... 10. Onoreidium, new genus
-. Head disc flat or anterior margin in the middle concave or clypeal emargination distinct; if clypeal
teeth absent, then emargination widely V-shaped (narrowly angled in the middle); mesotibiae and
metatibiae, in ventral view, with very reduced external tooth at mid-length; mesotibiae and metatibiae
transverse apical ventral carina bearing long setae ............................................................................. 13
13(12). Pseudoepipleuron with longitudinal angle making it approximately vertical on anterior half and hori-
zontal on posterior half; glabrous; width of eye (dorsal view) greater than one-half length (Figs. 47,
72); body length under 2.8 mm............................................................................................................ 14
-. Pseudoepipleuron entirely at the same plane, with distinct row of setae along most of length (rarely
lacking on anterior third only); width of eye (dorsal view) equal to or less than one half length; body
length over 2.8 mm .............................................................................................................................. 15
VAZ-DE-MELLO
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14(13). Hypomeron with posterior longitudinal carina; elytral interstriae uniformly flat, not distinctly more
convex apically; discal interstriae with single row of setose punctures. Central Brazil, Paraguay,
Argentina ................................................................................................... 7. Leotrichillum, new genus
-. Hypomeron without posterior longitudinal carina; elytral interstriae more convex apically; discal inter-
striae with two rows of punctures and one row of setae. Central Brazil .................................................
................................................................................................................. 3. Degallieridium, new genus
15(13). Mesotibiae and metatibiae slender, length at least four times apical width. Mexico, Guatemala, Argen-
tina, western Paraguay............................................................................... 8. Martinezidium, new genus
-. Mesotibiae and metatibiae strongly widened posteriorly, length at most three times apical width (in
some cases metatibiae strongly modified laterally) .............................................................................16
16(15). Eyes dorsally minute or absent, when present separated by more than 10 times their width (Figs. 55,
57, 60, 62, 64, 68, 70); pronotum unbeaded posteriorly, separated from hypomeron by weak lateral car-
ina between punctures; males with modified claws (Figs. 60, 62), length of mesotibiae and metatibiae
more than three times apical width; pygidium approximately 45 degrees from vertical plane, in ventral
view, its horizontal projection spanning at least 2/5 of abdominal length. Colombian Andes, southern
Venezuela, Bolivia, Paraguay, Brazil and Argentina .................................. 6. Genieridium, new genus
-. Eyes dorsally oval and well developed, separated by seven to eight times their width (Fig. 77); prono-
tum beaded posteriorly, separated from hypomeron by strong and smooth carina; males with claws
unmodified, mesotibiae and metatibiae less than three times as long as wide at apex; pygidium almost
vertical, in ventral view, its horizontal projection spanning less than 1/5 of abdominal length. Central
Argentina ....................................................................................................... 9. Nunoidium, new genus
1. Besourenga new genus
Diagnosis: Size small (2.0–3.5 mm); clypeus with two teeth separated by narrow U- or V-shaped emargin-
ation; clypeogenal edge straight or sinuate, with lateral border of clypeus angled or strongly curved lateral to
clypeal teeth (Figs. 39, 41); elytra with only one row of setose punctures on each discal interstria; pseudoepi-
pleuron strongly narrowed near metacoxa, extending to apex (Fig. 10).
Description: Size small, body oval (Figs. 15–16), color grey or brown to black, always lacking metallic
sheen. Clypeus anteriorly bearing two strong teeth, separated by narrow U- or V-shaped emargination. Cly-
peogenal and frontoclypeal sutures indistinct. Clypeal margin externally to clypeal teeth strongly curved or
angled, becoming straight anteriorly to clypeogenal border, which is straight or slightly sinuate. Genal margin
straight or slightly curved between clypeus and lateral angle (Figs. 39, 41). Pronotum lacking anterior or pos-
terior beads. Pronotal disc laterally with glabrous indistinct callosity, centrally covered by large setose punc-
tures. Hypomeron posteriorly with longitudinal carina. Mesoepimeron with transverse anterior carina.
Metasternum with setiferous punctures on the disc. Elytra with discal interstriae with single row of setiferous
punctures, mesal striae slightly to strongly widened and deepened at apical declivity. Pseudoepipleura
strongly narrowed near metacoxa extending to apex; with setiferous punctures present only at base. Protibiae
with three strong lateral teeth distributed along apical four fifths of tibial length; ventrally with scale-like setae
on teeth. Parameres shorter than one half of the length of the phallobase, simply elongated and flattened (Fig.
40). Coxites symmetrical well separated. Spermatheca spiral shaped.
Sexual dimorphism: Males bear a digitiform projection of mesal apical angle of protibia, and apical tar-
somere of protarsi excavated dorsally to receive proclaws that are strongly angulated at mid-length (Fig. 42).
Metasternal disc is somewhat concave in males, but very flat in females.
Type species: Trichillum minutum Saylor, 1935 (present designation) = Besourenga minutus (Saylor,
1935), new combination.
Zootaxa 1955 © 2008 Magnolia Press · 17
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Etymology: Combination of besouro (Brazilian Portuguese word for “beetle”), and Alvarenga, proper
name, in honor to Moacyr Alvarenga, good friend and amateur coleopterist responsible for collecting much of
the material studied here and specialist in the taxonomy of the family Erotylidae. Besourenga was his nick-
name in the Brazilian Air Force, where his passion for collecting beetles was widely celebrated. Masculine in
gender.
Distribution: Bolivia east of the Andes, southern Amazonia in Bolivia, Peru and Brazil, eastern Paraguay
and central Brazil including western parts of São Paulo, Minas Gerais, and Bahia states. This distribution
comprises the Pantanal and Tapajós-Xingu provinces of the Amazonian subregion, and the Cerrado and
Chaco provinces of the Chacoan subregion (Morrone 2006).
Remarks: A clade comprising Trichillum, Besourenga, Degallieridium, Feeridium and Eutrichillum is
putatively supported by the absence of the transvese carina on mesoepimeral disc, while the sister-genus rela-
tionship of Besourenga and Trichillum is supported by the spiral spermathecal shape and the form of clypeal
border sides. The monophyly of the genus is supported by: (1) slight expansion of the lateral genal angle in
relation to other groups closely related; (2) smaller size; and (3) absence of a lateral pronotal callosity. The
optimizations of the presence of setiferous punctures on the pronotal disc (also present in one species of Tri-
chillum) and modification of proclaws in males are ambiguous and could be merely symplesiomorphies.
Except for B. horacioi from forests in southern Amazonia, all species of Besourenga are from open habitats in
the Chaco and Cerrado biomes. Specimens of this genus are usually collected with flight intercept traps, at
light, or more rarely with dung-baited pitfall traps.
Composition: Besides the type species, the new genus includes Besourenga vejdovskyi (Balthasar, 1939),
new combination (described as Trichillum), Besourenga amarillai (Aguilar-Julio, 2001), new combination
(described as Pedaridium), Besourenga horacioi (Martínez, 1969), new combination, and several new spe-
cies currently under study.
Material examined:
1.1. Besourenga minutus (Saylor, 1935), new combination
Trichillum minutum: Saylor 1935: 207; Balthasar 1939b: 13, 20, 24; Blackwelder 1944: 204; Hamel et al. 2006: 12.
Trichillum (Eutrichillum) minutum: Martínez 1969: 120–121; Ratcliffe 1980: 341
Type series: Holotype
&
: PARAGUAY: Concepción: Horquetá (USNM).
Non-type material examined: BOLIVIA: Santa Cruz: Ichilo, Buenavista, Tacu, III-1951, Martínez (1
CMNC); PARAGUAY: Concepción: Horquetá, IV-1934, Schultze (2 CMNC, 1 FVMC), Horquetá, 4-XII-
1934 (1 CMNC).
1.2. Besourenga vejdovskyi (Balthasar, 1939), new combination
Trichillum vejdovskyi: Balthasar 1939b: 20, 23–24; Martínez 1947: 110; Vulcano & Pereira 1967: 578; Hamel et al.
2006: 12.
Trichillum (Eutrichillum) vejdovskyi: Martínez 1969: 120–121; Ratcliffe 1980: 341
Type series: Holotype
%
: BOLIVIA: Santa Cruz: Umg. Buenavista, 450 m, Steinbach (NMP).
Non-type material examined: BOLIVIA: Santa Cruz: Gutiérrez, Portachuelo, II-1950, Martínez (1
CMNC); Río Piray, XI-1950, Martínez (1 CMNC); Santa Cruz, XI-1955, Zischka (1 FVMC).
1.3. Besourenga amarillai (Aguilar-Julio, 2001), new combination
Pedaridium amarillai: Aguilar-Julio 2001: 1–3
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Type series: Holotype
&
not seen (Museo de História Natural del Paraguay, Asunción).
Non-type material examined: BRAZIL: Distrito Federal: RECOR-IBGE, XII-1999, M Milhomem,
campo sujo (2 FVMC); Goiás: Goiânia, 21-XII-1984, MJ Ferreira (2 FVMC); Jataí, Faz. Nova Orlândia, I-
1964, Martins, Morgante & Silva (1 IBSP).
1.4. Besourenga horacioi (Martínez, 1969), new combination
Trichillum (?) horacioi: Martínez 1969: 142–145
Trichillum horacioi: Hamel et al. 2006: 12.
Type series: Holotype
&
: BOLIVIA: Santa Cruz: Sara, Nueva Moka, XII-1960, A Martínez (BRBA).
Non-type material examined: BRAZIL: Pará: Serra Norte, Piste N1 km 22, 1-XI-1984 (2 CMNC);
Redenção, XI-1999, P Scheffler (2 FVMC); X-1998, P&T Scheffler (10 FVMC); Rondônia: 62 km SW
Ariquemes, nr Faz. Rancho Grande, 8-20-XI-1994, J Eger, C O'Brien, black light (4 BDGC); PERU: Madre
de Dios: Río Palma Real Grande, Limon Camp, 220 m, X-1999, T Larsen (10 CMNC).
2. Bradypodidium, new genus
Diagnosis: Small oval to elongate species (Fig. 17), hairs very fine, clypeus with none, two or four teeth
(Figs. 43, 45), elytral striae deeper and wider posteriorly, protibial teeth concentrated in the apical one-half of
tibia; males with proclaws and last tarsomere strongly modified (Fig. 46); lacking apicomesal tooth in
protibia; parameres elongated, flat and wider at apex, always with mesal apices superposed (Fig. 44).
Description: Size 2.3–3.7 mm. Body oval, with elytra clearly wider than prothorax (Fig. 17); color tan to
black, rarely with traces of metallic sheen. Clypeus with none, two, or four teeth (interspecific and sexually
dimorphic in at least some species related to a wide variety of general head forms: transverse, rounded, or tri-
angularly elongated). Clypeal border lateral to teeth straight to feebly rounded, continuous with genal border.
Clypeofrontal and clypeogenal sutures indistinct, head dorsally flat except for anteromedian part of clypeus.
Eyes, in dorsal view oval; interocular width 8 to 10 times eye width (Figs. 43, 45). Pronotum lacking anterior
or posterior beads, lateral callosity weakly defined. Disc centrally with setose punctures. Hypomera laterally
separated from pronotal disc by strong carina and bearing a strong posterior longitudinal carina parallel to the
pronotal-hypomeral delimitation. Mesoepimera with anterior transverse carina. Metasternum with setose
punctures on the disc. Elytral interstriae almost flat, each with two well-separated rows of setose punctures;
striae wider and deeper on the posterior declivity. Pseudoepipleura gradually narrowed from anterior third to
end of elytron, with setiferous punctures all along. Protibiae with three teeth, basal one frequently reduced;
teeth concentrated in apical two-fifths of tibia; lateral tibial margin denticulate between base and basal tooth;
ventral scale-like setae of tibial teeth absent. Mesotibiae and metatibiae elongate, much longer than five times
their apical width. Metatarsus with first tarsomere flattened and oval, longer than second. Parameres blade-
like, tips slightly overlapped, apically wider than at mid-length (Fig. 44), about as long as half phallobase
length or shorter. Coxites present, large and symmetrical. Spermatheca C-shaped, gradually narrowed posteri-
orly, basally globular and strongly modified in conical point at duct insertion.
Sexual dimorphism: Apical tarsomere of protarsi of male excavated dorsally to receive proclaws; claws
strongly angulate at mid-length (Fig. 46). Metasternal disc somewhat concave in males, and very flat in
females. Protibial teeth much wider and robust in females; males frequently with very reduced basal tooth. In
species with variable clypeal shape, males often with more elongated clypeus when teeth are present, and
reduced or absent lateral clypeal teeth; while females almost always have transverse or rounded head with
four well developed clypeal teeth. Pygidium much more transverse and vertical in females than in males.
Zootaxa 1955 © 2008 Magnolia Press · 19
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Type species: Trichillum bradyporum Boucomont, 1928 (present designation) = Bradypodidium brady-
porum (Boucomont, 1928), new combination.
Etymology: Combination of Bradypus (generic name of the most common sloths [Mammalia: Edentata:
Bradypodidae], in reference to the phoretic relation these scarabs have with them) and Pedaridium, a genus
name. Bradypodidium is neuter in gender.
Distribution: Widely co-distributed with sloths in the Neotropical region from Costa Rica to southeastern
Brazil.
Remarks: Collection records suggest an obligatory relationship with sloths. Bradypodidium is closely
related to Trichillidium, sharing with the latter protibial teeth concentrated in the apical half of tibia, slender
dorsal setae, among many other characters. Monophyly of Bradypodidium is supported by the relatively slen-
der male tibia in relation to female, laminate overlapped parameres, relatively elongated body, transverse
pygidium in females, and the association with sloths. See under Onoreidium for more remarks on Trichillid-
ium-Bradypodidium.
Composition: Besides the type species, the genus Bradypodidium includes B. adisi (Ratcliffe, 1980), new
combination, B. venezuelense (Ferreira & Galileo, 1993), new combination, and at least three new species
presently under study.
Material examined:
2.1. Bradypodidium adisi (Ratcliffe, 1980), new combination
Trichillum (Eutrichillum) adisi: Ratcliffe 1980: 337–341
Pedaridium adissi [sic]: Ferreira & Galileo 1993: 28–29
Pedaridium adisi: Vaz-de-Mello & Canhedo 1998: 100; Vaz-de-Mello 2000: 194; Verdú & Galante 2001: 601–602
Type series: Holotype not seen (Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil).
Paratypes: BRAZIL: Amazonas: Ilha do Curari, 25-I-1978, R Best, Br tridactylus (8 CMNC, 2 CNCI);
Manaus, 2-VI-1977, J Adis, B. tridactylus (1 BDGC, 1 CMNC, 1 CNCI, 3 FVMC, 2 FMNH); Paricatuba nr.
Manaus, 14-VI-1977, J. Adis, col. from Brad. tridactylus (1 FMNH).
Non-type material examined: BRAZIL: Amazonas: Ilha do Curari, 2-VI-1977, J Adis, Sloth (1 CMNC,
1 FVMC); Pará: Ilha de Marajó, distr. Caldeirão, II-1924, Wilh. Ehrhardt ("Typus" Pedaridium setulosum
Balthasar i. litt. NMP); PERU: Loreto: R Yaropa, Puerto Miguel, 200 m, 16-23-XII-1994, T Hácz & G Holz-
inger (2 HMNH); Río Ucayali, Yarinacocha, 27-V-1945, J Schunke, pele de preguiça (1 CMNC, 1 FVMC).
2.2. Bradypodidium bradyporum (Boucomont, 1928), new combination
Trichillum bradyporum: Boucomont 1928: 188; Balthasar 1939b: 15–17, 26; Blackwelder 1944: 204; Costa-Lima 1953:
22
Pedaridium bradyporum: Martínez 1969: 119; Ferreira & Galileo 1993: 36; Vaz-de-Mello et al. 2002: 676; Solís &
Kohlmann 2003: 9–11
Type series: Holotype not sexed: COSTA RICA: Limón: Hamburgfarm, Reventazón, Ebene Limón, 21-VIII-
1925, F Nevermann, am Affer im Pelz von 3 zeilig Faulfier (MNHN).
Non-type material examined: COSTA RICA: Cartago: Catie, 3 km SE Turrialba, 600 m, 13-16-V-1985,
J Doyen (1 BDGC); Turrialba, Catie, 600 m, 16-V-1979, H&A Howden (1 CMNC); Turrialba, III-1952, A
Trejos, on Bradypus griseus (1 CNCI); Limón: Hamburgfarm, Reventazón, Ebene Limón, 27-X-1931, Nev-
ermann, am anus von Bradypus infuscatus (1 BDGC, 1 CMNC, 1 MZSP, 7 NHMB, 3 FVMC); 1936 (1
CMNC); 1936-39 (1 CNCI); ECUADOR: Esmeraldas: La Chiquita, 5 m, 11 km SE San Lorenzo, 3-16-VI-
1975, S&J Peck (1 CMNC).
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2.3. Bradypodidium venezuelense (Ferreira & Galileo, 1993), new combination
Pedaridium venezuelensis [sic]: Ferreira & Galileo 1993: 27–28
Type series: Holotype not seen.
Paratype: VENEZUELA: Carabobo: San Esteban, 100 m, 01-VII-1975, Martínez (1 FZRS).
Non-type material examined: VENEZUELA: Aragua: Rancho Grande 1100 m, 16-V-1967, H&A
Howden (1 CMNC); 18-19-II-1971, H&A Howden (1 CMNC); 24-25-II-1971, 800 m (1 FVMC).
3. Degallieridium new genus
Diagnosis: Body short, oval, convex (Fig. 18) and very small size (length 2.0 mm or less). Dorsum brown;
dorsal setae thick, as wide as lateral elytral striae. Clypeus with two teeth, laterally rounded; eyes dorsally as
long as twice their width (Fig. 47). Hypomeron without lateral longitudinal carina. Elytral interstriae with two
rows of punctures, only one of them bearing setae. Pseudoepipleuron medially inflexed, but gradually nar-
rowed posteriorly (Fig. 11). Parameres strongly divergent apically (Fig. 48).
Description: Length 1.8–2.0 mm; body short (Fig. 18), oval, bright brown, with no traces of metallic
sheen. Clypeus with two triangular teeth separated by V-shaped emargination, laterally simply rounded and
continuous with genal border. Clypeofrontal and clypeogenal sutures indistinct. Eyes, in dorsal view, twice as
long as wide; interocular width four times eye width (Fig. 47). Pronotum lacking anterior or posterior beads;
lateral callosity present, elongated and darker than remaining parts of pronotum; disc centrally with large set-
ose punctures. Pronotum and hypomeron separated by simple complete carina, with its posterior two-thirds
placed ventrally (anterior third lateral). Hypomeron laterally lacking longitudinal carina; mesoepimeron ante-
riorly lacking transverse carina. Metasternum with small glabrous punctures on the disc. Elytral interstriae
with two rows of punctures on disc, inner one bearing setae; interstriae flat on disc, apically convex; striae
apically clearly wider and deeper than on disc. Pseudoepipleura medially inflexed (posteriorly almost in a
transverse plane in relation to anterior part), but without sharp angle and gradually narrowed posteriorly (Fig.
11), completely glabrous. Protibiae laterally with three strong teeth distributed along apical seven-tenths, not
denticulate basally, and ventrally lacking scale-like setae on teeth. Width of apices of mesotibiae and metatib-
iae about one-third length of tibia. Length of parameres shorter than one-half that of phallobase, right angled
in relation to phallobase axis, apically flattened and strongly divergent with sharp lateral apical angles (Fig.
48). Internal sac with a long helicoidal pseudoflagellum occupying most of phallobase length, and secondary
pseudoflagellum flattened all along. Coxites small and well defined, glabrous. Spermatheca C-shaped,
strongly narrowed apically, basally somewhat bulbous; spermathecal duct not sclerotized.
Sexual dimorphism: Males have proclaws strongly bent in the middle with last tarsomere hollowed to
receive them, an apical mesal tooth in protibia directed downward, and less transverse pygidium compared to
females.
Type species: Degallieridium lilliputanum new species (monotypy).
Etymology: After Nicolas Degallier, friend, histeridologist and collector of part of the type series of the
species described below. Gender neuter.
Distribution: Central Brazil (Minas Gerais state and Distrito Federal) and Bolivia, corresponding to scat-
tered localities in Cerrado and Chaco provinces of the Chacoan subregion.
Remarks: This genus appears closely related to Feeridium, with which shares the absence of the
hypomeral lateral longitudinal carina, pseudoepipleural longitudinal fold, and parameral and spermathecal
general form. It is most easily distinguishable by size (Feeridium is about five times longer) and body form
(very elongated in Feeridium), although many other differences do exist, for instance, Feeridium lacks modi-
Zootaxa 1955 © 2008 Magnolia Press · 21
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
fication in male proclaws. Both genera belong to the clade that includes also Trichillum, Eutrichillum, and
Besourenga (see discussion above). A strong overall similarity exists between Degallieridium and Leotrichil-
lum, however relationships of the latter with other genera are obscure.
Composition: Besides the type species, the genus contains at least one new species currently in the pro-
cess of being described.
3.1. Degallieridium lilliputanum new species
Type series: Holotype
%
: BRAZIL: Distrito Federal: Brasilia, 1100 m, XI-2001, N. Degallier (MZSP ex-
FVMC).
Paratypes: BRAZIL: Distrito Federal: Brasilia, 1100 m, XI-1999 (1 CMNC); N. Degallier, at light (3
CMNC, 2 FVMC); X-2000, N. Degallier (1 FVMC); Minas Gerais: Paracatu, XII-1996, S. Lourenço (2
FVMC).
Description: Holotype
%
. Size, 1.9 mm. Color tan to light brown, dorsal setae claviform, pale yellow.
Head as long as wide, gently and regularly curved from the rounded genal lateral angle to clypeal teeth. Cly-
peal teeth triangular, short, separated by wide V-shaped emargination with rounded bottom. Head covered by
round simple punctures separated by 1.5 to two diameters; punctures variable in size, with smaller punctures
concentrated in anterior half and larger ones on posterior half; larger punctures up to 1.5 times as large as
smaller ones, some bearing a seta (Fig. 47). Pronotum evenly covered by large oval setose punctures, sepa-
rated by less than one to 1.5 diameters, some punctures bearing a seta. Posterior two-thirds of pronotal lateral
margin bent ventrally and hidden from dorsal view; anterior third visible from above. Prosternum and mesos-
ternum covered by oval, ocellated, glabrous punctures, separated by less than half their diameter; hypomeron
with similar but slightly smaller and more rounded punctures, more sparse anteriorly. Mesometasternal suture
rounded at midlength, more strongly arched than laterally, resulting in a broad rounded triangular metasternal
lobe. Posterior two-thirds of anterior metasternal lobe with sides nearly parallel; slightly divergent; lobe and
metasternal disc covered by simple punctures, with very sparse ocellate punctures posteriorly and laterally,
similar to those of metasternal sides, where glabrous ocellated punctures are much smaller and sparser than in
prosternum and mesosternum. Elytral mesal striae with regular punctures separated by about three diameters,
each puncture as wide as 1.5 strial widths on disc; striae on posterior fourth much deeper and wider, with
punctures nearly confluent. Interstrial setae wider and denser laterally than mesally. Length of apical tooth of
protibia approximately three times that of basal tooth, and 1.5 times that of middle tooth; spur triangular, flat-
tened dorsally. Length of mesofemora and metafemora about three times their maximum width. Metatarsi
with basal tarsomere rectangular, length about 1.25 times that of second tarsomere. Abdominal sternites cov-
ered by punctures similar to those of lateral metasternal lobes. Pygidium about 1.5 times as wide as long, with
very sparse simple punctures; basal punctures setiferous. Length of parameres slightly less than half that phal-
lobase, with lateroapical angles directed ventrally (Fig. 48). Internal sac with helicoidal pseudoflagellum
occupying most of phallobase length, base of which strongly curved; length of secondary pseudoflagellum
two thirds that of primary one.
Variation: Paratypes vary only in sexual features (females lack modifications in protarsi and claws, lack
the mesoapical tooth of protibia, have smaller protibial teeth, more transverse pygidium, and abdominal stern-
ites more convex and with longer disc) and slightly in size (1.8–2.0 mm).
Etymology: Reference to Lilliput, the island of small people, the Lilliputians, in Jonathan Swift’s 1726
novel, Gulliver’s Travels.
Distribution: Central Brazil (northwestern Minas Gerais and Distrito Federal), corresponding to the cen-
tral part of the Cerrado province in the Chacoan subregion (Fig. 103).
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4. Eutrichillum Martínez, 1969 new status
Trichillum (Eutrichillum): Martínez 1969: 121; Ratcliffe 1980: 340–341; Martínez 1989: 60; Solís & Kohlmann 2003:
10, 12.
Trichillum (Eutrichillum) Martínez, 1967 [sic]: Vaz-de-Mello, 2000: 195.
Diagnosis: Head with clypeogenal margin strongly incised (causing clypeus and genae to be separately
rounded) and clypeogenal suture strongly marked, in some cases joining the clypeofrontal suture and forming
a very feeble acute carina, insconspicuous at midlength (Fig. 49). Pseudoepipleura strongly narrowed posteri-
orly.
Description: Length 2.7–4.6 mm, body short oval (Fig. 19), color grey to black with feeble metallic
sheen, or completely metallic green to red. Clypeus with two strong teeth, separated by narrow U-shaped
emargination, mesal border of tooth almost parallel; clypeus external to teeth regularly rounded, separated of
genal border by incision, genal border being separately strongly and regularly curved. Clypeofrontal and cly-
peogenal sutures strongly indicated and united, being frequently cariniform at least near junction. Eyes dor-
sally as wide as third to fifth of its length, interocular width 8 to 12 times eye width (Fig. 49). Pronotum
lacking anterior or posterior beads; lateral pronotal callosity well defined and elongate; disc centrally with
uniform elongate setose punctures; pronotum separated of hypomeron by complete carina. Hypomeron with
strong lateral longitudinal carina; mesoepimeron anterior transverse carina present; metasternal disc with
round simple setose punctures. Elytral interstriae with only one row of setose punctures, situated laterally in
interstriae one and two, and mesally in the remaining interstriae; interstriae flat all along, slightly deeper and
widened posteriorly. Pseudoepipleura strongly narrowed posteriorly from about metacoxa, forming strong
marginal angle; glabrous all along. Protibiae with three very strong teeth, distributed along apical three-fifths,
denticulate basally, with ventral scale-like setae. Mesofemur and metafemur as wide as third to fourth of their
length, tibiae apically as wide as third to fourth of their length. Metatarsi with basal tarsomere longer than sec-
ond tarsomere. Parameres with apex strongly deflexed, forming a strong dorsal hump at apical third, with
strong outer subquadrate flat apical lobes directed ventrally (Fig. 50). Internal sac with pseudoflagellum
highly sclerotized, positioned spirally inside phallobase, but straightened when extruded, with wide triangular
base with one to three strong teeth and pointed apex, with the entire body canaliculate longitudinally, and
large triangular plate accessory lamella; coxites not heavily sclerotized, large. Spermatheca C-shaped, sinuate
in the middle, but with strongly narrowed and spiral-shaped base and apex (Fig. 14), duct very long, spirally
shaped and not sclerotized.
Sexual dimorphism: Males have proclaws strongly bent mesally and apical tarsomere modified in order
to receive them (Fig. 51); protibial apicomesal tooth present in males and apically directed; abdominal disc
flatter, pygidium less transverse, and metasternum flatter than in females.
Type species: Trichillum boucomonti Saylor, 1935 = T. hirsutum Boucomont, 1928 (original designation)
= Eutrichillum hirsutum Boucomont, 1928, new combination.
Distribution: A group of species (of which two already described) inhabit South American lowlands, east
of the Andes as far south as Buenos Aires in Argentina; one species in Costa Rica. This is a very disjunct dis-
tribution, corresponding on one side the entire Chacoan, Parana and Amazonian subregions, and on the other,
some localities in Eastern Central America province of the Caribbean subregion.
Remarks: Species of this genus are mostly necrophagous and frequently attracted to lights. The genus
belongs to a clade that includes Feeridium, Degallieridium, Besourenga and Trichillum (see above under
Besourenga), and is characterized by the following synapomorphies: clypeus and gena separately rounded;
clypeogenal and clypeofrontal sutures strongly indicated; parameres forming dorsal hump at apical third;
pseudoflagellum hosted by internal sac spirally and straightened when extruded; spermatheca with base and
apex strongly narrowed and spiral shaped.
Composition: Apart of the type species, the genus contains Eutrichillum hystrix (Arrow, 1931), new
Zootaxa 1955 © 2008 Magnolia Press · 23
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
combination (described as Trichillum), E. arcus (Solís & Kohlmann, 2003), new combination (described as
Trichillum), and at least five new species currently being described.
Material examined:
4.1. Eutrichillum arcus (Solís & Kohlmann, 2003), new combination
Trichillum (Eutrichillum) arcus: Solís & Kohlmann 2003: 10, 12–14
Type series: Holotype not seen (Instituto Nacional de Biodiversidad, San José, Costa Rica).
Paratypes: COSTA RICA: Alanjuela: Sect. San Ramón de los Ríos, 1.5 km NO Hda. Nueva Zelandia,
620 m, 12-21-VI-1996, FA Quesada (3 FVMC); Guanacaste: Est. Pitilla, 9 km S Sta. Cecilia, P.N. Guana-
caste, 700 m, V-1994, C Moraga (2 FVMC); Heredia: Est. Biol. La Sielva, 21-VI-1998, C Carlton & A
Tishechkin (1 FVMC).
4.2. Eutrichillum hirsutum (Boucomont, 1928), new combination
Trichillum hirsutum: Boucomont 1928: 187; Arrow 1932: 226; Paulian 1936: 206–207; Balthasar 1939b: 13, 19, 22–23;
Pessôa & Lane 1941: 447; Blackwelder 1944: 204; Martínez 1969: 119; Vaz-de-Mello & Génier 2005: 44–45;
Hamel et al. 2006: 12.
Trichillum boucomonti Saylor, 1935: Saylor 1935: 208; Balthasar 1939b: 13, 18, 22; Blackwelder 1944: 204
Trichillum (Eutrichillum) boucomonti; Martínez 1969: 120–121; Ratcliffe 1980: 341; (pars) Ratcliffe 1981: 185; Mar-
tínez 1987: 60; Monteresino et al. 1996: 107; Vaz-de-Mello 2000: 195; Génier & Vaz-de-Mello 2002: 189
Trichillum (Eutrichillum) hirsutum; Ratcliffe 1980: 341; (pars) Ratcliffe 1981: 183–184; Vaz-de-Mello 2000: 195.
Type series: Trichillum hirsutum Boucomont, 1928: Holotype
&
: BRAZIL: São Paulo: no locality (MNHN).
Trichillum boucomonti Saylor, 1935: Holotype
&
: PARAGUAY: Concepción: Horquetá (USNM).
Non-type material examined: ARGENTINA: Corrientes: Sto Tomé, XI-1945, Martínez (2 CMNC); Mis-
iones: PN Iguazu, 180 m, XII-1990-I-1991, S Peck (2 CMNC); BOLIVIA: Ichilo: Buenavista, XI-XII-1948,
L. Peña (1 FMNH); Santa Cruz: Gutiérrez, Portachuelo, II-1950, Martínez (2 CMNC); Santa Cruz, XI-1955,
Zischka (5 CMNC); BRAZIL: Bahia: Barreiras, XII-1991 (1 FVMC); Encruzilhada, XII-1980, Alvarenga &
Martínez (2 CMNC, 1 FVMC); Encruzilhada, 980 m, XI-1972, Alvarenga (3 MZSP); Distrito Federal:
Brasília 1100 m, II-2001, N Degallier (2 FVMC); X-2000 (4 FVMC); XI-1999 (1 CMNC); XII-1997, P.
Grossi (1 FVMC); Est. Florestal Cabeça do Veado, 1100 m, 27-X-1971, EG, I & EA Munroe (1 CNCI);
RECOR-IBGE, 27-V-1997, I Diniz (1 FVMC); XI-1999, M Milhomem, campo sujo (1 FVMC); XII-1999 (1
FVMC); Espírito Santo: Pque Sooretama, Linhares, 12-27-X-1962, FS Pereira (1 MZSP); Venda Nova do
Imigrante, X-1998, Falqueto & Vaz-de-Mello (2 FVMC); Goiás: Bom Jardim de Goiás, II-1997, FZ Vaz-de-
Mello (2 FVMC); Goiânia, 03-XII-2001, SS Silva (3 FVMC); Jataí, Faz Nova Orlândia, I-1964, Martins,
Morgante & Silva (2 MZSP, 1 IBSP); Jataí, I-1955 (2 CMNC, 10 MZSP); Rio Verde, II-1998, J Carlos (1
AMBC); Mato Grosso do Sul: Costa Rica, 17-XII-1993, S Ide (5 MZSP); Nova Andradina, II-1996, Louzada
& Vaz-de-Mello (1 FVMC); Selvíria, UNESP farm:01-IV-1999, CAH Flechtmann, ex black light, Brachiaria
decumbens pasture (1 FEIS); 20-I-1999 (1 FEIS); Mato Grosso: Barra do Tapirapé, 2-16-I-1966, B Malkin (1
IBSP); XI-1964, Malkin (1 CMNC); Diamantino, Alto Rio Arinos, XI-1998, E Furtado (1 FVMC); Pq Nac.
Xingu, Jacaré, XI-1961, Alvarenga & Bokermann (116 MZSP); Xingu:XI-1947 (1 MNRJ); XI-1961 Alva-
renga & Werner (89 MZSP); XI-1961, Alvarenga (3 CMNC); Minas Gerais: Açucena, II-1952, Pereira (1
CMNC); Águas Vermelhas, XII-1998, Bello & Vaz-de-Mello (1 FVMC); Belo Horizonte, X-1950 (1 MZSP);
Buritis (Rib. Confins), X-1964, Exp. Dep. Zool. (1 CMNC); Caxambu, 02-XII-1990, Bello (1 FVMC); Cord-
isburgo, Faz Pontinha, I-1992, FZ Vaz-de-Mello (1 FVMC); Ipatinga, XI-1994, E Grossi (1 FVMC); Mar-
tinho Campos, X-1991 (1 FVMC); Montes Claros, I-2000, JNC Louzada (3 FVMC); Nova Era, I-1993
(1FVMC); Paracatu, II-1997, S Lourenço (2 FVMC); XII-1996 (3 FVMC); Vespasiano, I-1952, Pe. Pereira (1
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24 · Zootaxa 1955 © 2008
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BDGC, 6 CMNC); XII-1945, Pereira (4 CMNC); XII-1951 (1 MZSP); Viçosa, mata do Paraíso, I-1995, JNC
Louzada (1 FVMC); I-1996, Louzada, Sperber & Vaz-de-Mello (2 FVMC); Viçosa, 02-II-1994, JNC Louzada
(1 FVMC); 03-II-1994 (4 FVMC); 04-I-1994 (3 FVMC); 07-I-1994 (2 FVMC); 10-I-1992, Lopes & Louzada
(1 FVMC); 10-I-1994, JNC Louzada (2 FVMC); 12-I-1992, Lopes & Louzada (1 FVMC); 21-XI-1991, Lopes
& Louzada (1 FVMC); 25-I-1994, JNC Louzada (1 FVMC); Paraná: Curitiba, II-1944, Hatsbach (1 MZSP);
Londrina, :XI-1998, J Lopes (3 AMBC); XI-XII-1998, IM Medri (1 FVMC); Vila Velha, II-1945 (1 MZSP);
Rio de Janeiro: 17 km E Nova Friburgo, 750 m, 21-I-2000, Génier & Ide (1 CMNC); Itatiaia, 20-I-1993, CL
Godinho Jr (1 FVMC); Nova Friburgo, 1000 m, I-2002, P&E Grossi (2 FVMC); I-2001 (1 AMBC); III-1998,
P Grossi (1 FVMC); XI-1996, FZ Vaz-de-Mello (1 FVMC); Rio Grande do Sul: Near Cachoeira, Henninger
(1 CMNC); São Paulo: no locality, Mráz leg. (2 NHMB); Aclimação, XII-1958 (1 CMNC); Barueri, 22-II-
1956, K. Lenko (1 MZSP); XII-1955 (1 CMNC); Botucatu, 22-II-1955, Werner (3 MZSP); Cerqueira César,
XII-1999, J Carlos (1 AMBC); Cidade, Ipiranga, XII-1958, A Martínez (3 BDGC, 5 CMNC); XII-1962, Mar-
tínez (2 CMNC); Itu, Faz. Pau d' Alho, 12-15-XI-1960, Martins (1 MZSP); II-1969, Martins (1 MZSP); Mogi
Guaçu, Faz. Campininhas, I-8-I-1970, JM & BA Campbell (2 CNCI); Pirassununga, Usina – luz, 9-XI-1945,
Schubart (2 MZSP); XI-1952 (2 MZSP); Pirassununga, XI-1952 (1 CMNC); XI-1956, Martínez (3 CMNC);
Ribeirão Preto (Fac. Medicina), I-1955, Barretto (1 MZSP); Rio Claro, XII-1942, Claretiano (17 MZSP); São
Paulo, Ipiranga (1 MZSP); Ypiranga, XI-1929, Spitz (1 MZSP); PARAGUAY: Do.?: Chaco, XII-1934,
Schultze (1 CMNC); Alto Paraná: Limoy, XI-1990, G Arriágada (3 FVMC); RB Itabo, X-1989, G Arriágada
(2 FVMC); Canindeyú: Est. Pozuelo, XII-1990, G Arriágada (3 FVMC); Central: Asunción, IX-1944 (5
MZSP); XI-1944 (1 CMNC); Concepción?: Mariscal Estigarribia, I-1990, G Arriágada (1 FVMC); Con-
cepción: Horquetá, I-1934, Schultze (1 CMNC); I-1943 (1 CMNC); IV-1934 (1 CMNC); Guayrá: Villarrica,
X-1934, Köller (1 MNHU); XI-1955, Schade (2 CMNC); Paraguary: Naranjo, 09-14-XII-1998 (1 FVMC);
Sapucay, 05-XI-1991, U Drechsel (12 ABC); San Pedro: Río Ypané, Cororó, XI-1979, Martínez (17
CMNC); San Pedro?: Peribebuy, XI-1946 (1 CMNC).
4.3. Eutrichillum hystrix (Arrow, 1931), new combination
Trichillum hystrix: Arrow 1931: 609-610; Arrow 1932: 226; Paulian 1936: 206, 207; Balthasar 1939b: 13, 21, 25; Black-
welder 1944: 204; Martínez 1959: 64.
Trichillum (Eutrichillum) hystrix: Martínez 1969: 120–121; Ratcliffe 1980: 341; Martínez 1987: 60; Génier & Vaz-de-
Mello 2002: 188–190.
Type series: Lectotype
%
(designated by Génier & Vaz-de-Mello, 2002): ARGENTINA: Santa Fé: Estancia la
Noria, Rio San Javier, G.E. Bryant, 27.XII.1911 (BMNH). Paralectotypes: Same data as lectotype except:
XII.1911 (1
%
, 1
&
BMNH), 10.XII.1911 (1
&
CMNC), 14.XII.1911 (1
%
BMNH), 20.XII.1911 (2
&&
BMNH),
23.XII.1911 (2
&&
BMNH), 3.I.1912 (1
%
BMNH).
Non-type material examined: ARGENTINA: Buenos Aires: Belgrano, XII-1941 (1 CMNC); Capital
Federal, Villa Devoto, XII-1925, Bridarolli (2 CMNC); Gral. Sarmiento, JC Paz, I-1952, Martínez (5 CMNC);
San Isidro, Casa, I-1960, Martínez (2 CMNC); Tigre, V-1945, MJ Viana (2 CMNC, 1 FVMC); Buenos Aires,
Richter (3 IRSN); Córdoba: Cruz Alta; II-1946; JP Duret (4 CMNC); Santa Fé: Do. Capital, Piquete; I-
1942; Martínez (2 CMNC); Estancia La Noria, Río San Javier, 23-XII-1911, GE Bryant (1 NHMB); Santa
Fé?: Carcarana (10 UNSM, 4 FVMC); PARAGUAY: Alto Paraná: Puerto Stroessner; 6-I-1966; Hungarian
Soil-Zool. Exp. (1 HMNH); San Pedro?: Peribebuy:V-1946, Williener (1 CMNC); NO DATA (4 FMLT).
5. Feeridium new genus
Diagnosis: Size large (4.5–5.2 mm), very elongated (Fig. 20). Mesofemora and metafemora very strong and
rounded (Fig. 54), eyes extremely large dorsally, interocular width narrower than twice eye width (Fig. 52).
Eytral interstriae sparsely punctate, with uniseriate setae only at apex and sides.
Zootaxa 1955 © 2008 Magnolia Press · 25
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Description: Length 4.5–5.2 mm, body very elongated, with subparallel sides (Fig. 20); color brown,
opaque, without metallic sheen. Clypeus with two strong teeth, separated by narrow U-shaped emargination,
clypeus laterally and gena continuously rounded. Clypeofrontal and clypeogenal sutures not discernible. Eyes,
in dorsal view, enormous, as long as wide; interocular width less than 1.5 times eye width (Fig. 52). Pronotum
lacking anterior and posterior beads, lateral pronotal callosity rounded and darkened; disc covered by simple
strong punctures, glabrous; separated of hypomeron by row of punctures, without carina. Hypomeron without
lateral longitudinal carina; mesoepimeron anterior transverse carina strong and distinct; metasternal disc with
minute impressed punctures. Elytral interstriae flat, with unorganized punctures on disc, sparse uniseriate
setae present only apically and laterally; striae slightly deeper posteriorly, not widened. Pseudoepipleura grad-
ually narrowed posteriorly, with slight longitudinal fold near metacoxa, completely glabrous. Protibia with
three very strong teeth, distributed along apical three-fifths, basally not denticulate, lacking ventral scale-like
setae. Mesofemora wider than third of their length, metafemora about as wide as half of their length (Fig. 54);
mesotibiae apically wider than one of their length, metatibiae apically quite as wide as half of their length
(Fig. 54). Metatarsi with basal tarsomere as long as 1.5 times the following tarsomere. Parameres as long as
two-thirds of phallobase, flattened and truncate, apically divergent (Fig. 53). Internal sac with pseudoflagel-
lum relativelly short and bent at mid-length, rounded by flat large accessory lamella. Coxites triangular, very
sclerotized, and pointed. Spermateca C-shaped, bulbous basally (narrower at duct insertion), drastically nar-
rowed apically, with duct short and not sclerotized.
Sexual dimorphism: Males have central portion of last abdominal sternite flat to slightly concave, pygid-
ium more elongated, and a protibial apicomesal tooth, ventrally directed.
Type species: Feeridium woodruffi new species (monotypy).
Etymology: After François Feer, who collected and sent me the first specimens I saw of this genus. Gen-
der neutral.
Composition: The new genus currently includes solely the new species.
Distribution: French Guyana and Amazonian Brazil. Known from Humid Guiana, Roraima and Varzea
provinces of the Amazonian subregion.
Remarks: The condition of the parameres and pseudepipleura relates this genus to Degallieridium. How-
ever, it superficially resembles both Genieridium and Nunoidium, from which it can be immediately distin-
guished by the condition of pseudepipleuron, head, and elytra. Synapomorphies defining this genus include
lack of row-organized elytral disc punctation, absence of pronotal-hypomeral carina, lack of sexual differ-
ences in protarsi, size of eyes and psammophiliform development of legs.
5.1. Feeridium woodruffi new species
Type series: Holotype
%
: BRAZIL: Amazonas: Tabatinga, XI-1956, FM Oliveira (BRBA).
Paratypes: BRAZIL: Amazonas: 70 km N Manaus, Fazenda Esteio, 07-VI-1986, B Klein, human dung,
nature forest, AM (1 FVMC); Tabatinga, XI-1956, FM Oliveira (1 BRBA); FRENCH GUYANA: Cayenne:
Nouragues, III-1997, F Feer (1 FVMC); V-1995 (1 FVMC).
Description: Holotype
%
. Length 5.0 mm. Body elongated, aphodiiform (Fig. 20). Head wider than long,
clypeus with two rounded triangular teeth, slightly rounded laterally, with margin continuous to that of genae.
Head convex, concave anteriorly and transversely concave anteriorly to each eye, forming a diagonal lump at
each side, indicating the location of the indistinct clypeogenal suture. Head surface with mixed simple setose
and simple smaller glabrous punctures, the setose ones denser on clypeus. Eyes about as wide as long (Fig.
52); interocular width about 1.3 times eye width. Pronotal disc convex, covered by simple glabrous punctures
distanced by 1 to 3 diameters, setae present only on anterolateral and lateral margins. Hypomeral posterior
punctures larger but more sparse than at pronotal disc. Mesosternum covered by large almost coalescent ocel-
late diagonally elongated punctures. Mesometasternal suture strongly angled with a rounded tip, metasternal
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26 · Zootaxa 1955 © 2008
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anterior lobe clearly narrower posteriorly, rhomboid. Metasternum with disc with rhomboid, feebly delimited
concavity, covered by sparse glabrous simple punctures. Lateral metasternal lobes with diagonally elongated
ocellated glabrous punctures, distanced by about one diameter. Elytra with discal striae sinuate-punctate,
punctures indistinct and almost contiguous or separated by less than one diameter; discal interstriae with unor-
ganized small punctures, distanced by two to four diameters; setae present only apically, and in one well
delimited row in the eighth interstria, mesally. Protibiae with apicomesal tooth short, triangular and downward
directed; spur flattened, ogival with acute tip. Mesotibiae and metatibiae with apex with apical carina accom-
panied by row of very dense and long setae, longer than basitarsomere (Fig. 54). Metatibial basal tarsomere
flattened and trapezoidal. Abdomen with rounded ocellate punctures on disc, distanced by about 2 diameters,
with lateral row of setose punctures posteriorly on the last sternite. Pygidium with simple sparse setose punc-
tures. Parameres as in Fig. 53.
Variation: Paratypes vary in size (4.5–5.2 mm) and in sexual features: females lack the protibial api-
comesal tooth and the metasternal disc concavity.
Etymology: After Dr. Robert Woodruff, scarabeidologist. Although the specific epithet was used previ-
ously as an unpublished name for Martínez’s material deposited at BRBA, I extend my own homage to Dr.
Woodruff.
Distribution: French Guyana and Brazil (Amazonas state), as for the genus (Fig. 103).
6. Genieridium new genus
Diagnosis: Moderate to large-sized species (only rare specimens of G. c r y pto p s measure less than 3.7 mm),
oval-elongate shape (Figs. 21–22); clypeal teeth two (Figs. 55, 60, 62, 64, 70) or none, in the latter case
clypeus widely emarginated (Figs. 57, 68). Eyes, in dorsal view, absent or very small, distanced by more than
12 times eye width. Pronotum separated from hypomeron by weak carina (Fig. 59). Elytra convex at least in
the anterior half. Length of mesotibiae and metatibiae at least three times apical width. Males with mesal api-
cal protibial teeth; all claws bent (mesoclaws and metaclaws less modified than proclaws); and last tarsomere
of prolegs modified to receive claws. Length of last abdominal sternite along midline equal to or less than
medial width of metafemur; pygidium, in ventral view, occupying more than two-fifths of abdominal horizon-
tal length.
Description: Length 3.5–6.5 mm, body oval elongated, convex (Figs. 21–22), color grey to black, brown
in teneral specimens, shiny or opaque, lacking any traces of metallic sheen. Clypeus with two teeth or none,
always with central anterior concavity coincident with emargination, which is also present in toothless species
(Figs. 57, 68). Clypeus laterally regularly rounded, continuous with genal border. Clypeofrontal and clypeoge-
nal sutures indistinct, head lacking any traces of carinae or tubercles. Eyes, in dorsal view, absent or very
small, when present elongated, more than three times their width; interocular width at least 12 times eye width
(Figs. 55, 57, 60, 62, 64, 68, 70). Pronotum lacking anterior or posterior beads, simply convex, with lateral
callosities distinguished only by lack of punctation; disc covered by simple setose punctures and separated of
hypomeron by weak but distinct carina interrupted by regular setose punctures. Hypomeron ventrally with
strong lateral longitudinal carina; mesoepimera with transverse anterior carina stronger mesally. Metasternum
with simple setose punctures on disc. Elytra with one or two rows of setose punctures, discal interstriae flat all
along; striae apically deeper and slightly wider than on disc. Pseudoepipleura gradually narrowed posteriorly,
superiorly with complete row of setose punctures. Protibiae with three strong teeth distributed along at least
apical three-fifths, basally denticulate, with ventral scale-like setae. Mesotibiae and metatibiae strongly wid-
ened apically, apical width more than third of tibial length. Parameres flattened, right-angled in relation to
phallobase, shorter than half of phallobase length. Spermatheca simply C-shaped, in one case (G. cryptops)
with elongated base and apex. Coxites triangular and symmetrical.
Sexual dimorphism: Males have proclaws bent at midlength, and mesoclaws and metaclaws also modi-
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SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
fied, either bent at midlength or larger than in females and with stronger basal angle; protarsus with last tar-
somere modified to receive claws; metasternal disc strongly concave (less concave or flat in females),
pygidium narrower than in females, and each tibia with apicomesal tooth, sometimes modified in laminate
anteriorly directed apodeme in metatibia.
Type species: Pedaridium bidens Balthasar, 1942 (present designation) = Genieridium bidens (Balthasar,
1942), new combination.
Etymology: After François Génier, good friend and scarabeidologist, companion in so many works and
ideas. Gender neutral.
Distribution: Colombia, Venezuela, non-Amazonian Brazil, eastern Bolivia, Paraguay, and northeastern
Argentina. The genus shows a disjunct distribution, with one group in the Caribbean subregion (Northwestern
South American dominion), occupying the Cauca, Maracaibo, Venezuelan Coast and Venezuelan Llanos
provinces, and the other in the Chacoan, Parana and Amazonian subregions, occupying the Pantanal, Caat-
inga, Cerrado, Chaco, Brazilian Atlantic Forest, Parana Forest, and Araucaria angustifolia Forest provinces
(Figs. 104–107).
Remarks: The position of Genieridium was not resolved, and it was recovered in a trichotomy with
Pedaridium and Nunoidium. The new genus main external synapomorphies are the elongation and inclined
orientation of pygidium, and the punctation interrupting the pronotal-hypomeral carina. Apart from characters
mentioned as synapomorphies of Genieridium, it can be distinguished from Pedaridium by the much shorter
phallobase, differently shaped internal sac sclerites (Pedaridium lacks all but the pseudoflagellum), lack of
basal “tooth” in the spermatheca, elytra conjointly convex (Pedaridium has clearly tectiform elytra, separately
flattened), lack of longer setae on the anterior part of pronotum and lack of a pronotal lateral expansion. From
Nunoidium it is readily distinguished by the lack of a posterior pronotal bead, reduction of lateral bead,
reduced eyes, and presence of sexually dimorphic claws. This genus includes two clearly delimited mono-
phyletic, biogeographically-defined groups: species from Venezuela and Colombia are characterized by
strong modification of male metalegs (with flattened, blade-like apicomesal tooth) and southern species by the
reduction of the seventh elytral stria.
Composition: Besides the type species, the genus is presently defined to include G. cryptops (Arrow,
1913), new combination; G. paranense (Arrow, 1932), new combination; G. bordoni (Martínez, 1992), new
combination; G. zanunciorum (Vaz-de-Mello & Canhedo, 1998), new combination; G. margareteae (Génier
& Vaz-de-Mello, 2002), new combination; and G. medinae (Gill & Vaz-de-Mello, 2003), new combination
(all described as Pedaridium). Because no undescribed species of this genus are known to me, I present a
small synopsis below.
Key to species of Genieridium
1. Eyes hidden dorsally (Fig. 64). Central Colombia ......... 6.5. G. medinae (Gill & Vaz-de-Mello, 2004)
-. Eyes small but visible dorsally ............................................................................................................. 2
2(1). Clypeal teeth lacking (Figs. 57, 68) ...................................................................................................... 3
-. Two clypeal teeth present...................................................................................................................... 4
3(2). Elytral striae without evident punctures. Venezuela .......................... 6.2. G. bordoni (Martínez, 1992)
-. Elytral striae with ocellated punctures. Southern Brazil ..................... 6.6. G. paranense (Arrow, 1932)
4(2). Clypeal teeth very acute, sides almost parallel in the apical part (Figs. 60, 70).................................... 5
-. Clypeal teeth obtuse, short, and equilateral (Figs. 55, 62).................................................................... 6
5(4). Clypeal teeth arising from below clypeal margin (Fig. 60). Central Brazil, Northern Argentina, Para-
guay and Bolivia .................................................................................... 6.3. G. cryptops (Arrow, 1913)
-. Clypeal teeth at same plane as clypeal margin (Fig. 70). Southeastern Brazil ........................................
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28 · Zootaxa 1955 © 2008
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.......................................................................... 6.7. G. zanunciorum (Vaz-de-Mello & Canhedo, 1998)
6(4). Elytral striae straight, without punctures. Central Brazil and Paraguay .................................................
............................................................................................................... 6.1. G. bidens (Balthasar, 1938)
-. Elytral striae marked by inconspicuous punctures that make striae appear sinuated. Northeastern and
Central Brazil ....................................................... 6.4. G. margareteae (Génier & Vaz-de-Mello, 2002)
Material examined:
6.1. Genieridium bidens (Balthasar, 1938), new combination
Pedaridium bidens: Balthasar 1938: 218–220; Blackwelder 1944: 203; Vaz-de-Mello & Génier 2005: 45–46
Pedaridium hirsutum: Pessôa & Lane 1941: 437
Pedaridium brasiliensis Ferreira & Galileo, 1993: Ferreira & Galileo 1993: 5–7, 21–23, 54; Koller et al. 1999: 405, 407;
Vaz-de-Mello 2000: 194; Verdú & Galante 2001: 599–601
Type series: Pedaridium bidens Balthasar, 1938. Lectotype (designated by Vaz-de-Mello & Génier, 2005):
%
,
Paraguay (NMP). Paralectotypes: BRAZIL: Goiás: Jatahy (1 NMP, 1 NHMB)
Pedaridium brasiliensis Ferreira & Galileo, 1993: Holotype: BRAZIL: Bahia: Encruzilhada, 980 m, XI-
1972, Alvarenga (MZSP)
Paratypes: BRAZIL: Bahia: Encruzilhada, 980 m, XI-1972, Alvarenga (8 MZSP); Mato Grosso do Sul:
Murtinho (luz), I-1930, R Spitz (2 MZSP); Três Lagoas, Faz. Dr. José Mendes, 15-30-V-1964, Exp. Dep.
Zool. (1 MZSP); Faz. Retiro das Telhas, 15-30-V-1964, Exp. Dep. Zool. (2 CMNC, 11 MZSP); Mato Grosso:
Chapada dos Guimarães, XI-1963, Alvarenga (3 MZSP); Minas Gerais: Arinos, 06-08-XI-1964, Exp. Dep.
Zool. (2 MZSP); Paraná: Vila Velha, XI-1944, Hatsbach (1 CMNC, 1 MZSP); São Paulo: Castilho, marg.
esq. Rio Paraná, X-1964, Exp. Dep. Zool. (1 MZSP); Franca, VIII-1910, Garbe (2 MZSP); Itu, Faz Pau
d'Alho, 28-29-X-1965, Martins & Biasi (1 CMNC); Itú, 27-XII-1957, U. Martins (4 MZSP); XI-1958, U.
Martins (2 MZSP); Pirassununga, 09-X-1945, Schubart (1 MZSP).
Non-type material examined: BRAZIL: Bahia: no data, Bondar (1 MNRJ); Encruzilhada, 980 m, XI-
1972, Alvarenga (1 IBSP); XI-1974, Alvarenga (1 CMNC); XII-1980, Alvarenga & Martínez (8 CMNC);
Vitória da Conquista, I-1993 (2 MZSP); Distrito Federal: Est. Florestal Cabeça do Veado, 1100 m, X-1971,
EG, I & EA Munroe (1 CNCI); Goiás: Bom Jesus, X-1996, J Carlos (1 FVMC); Niquelândia, X-1993, arm
luminosa (1 FVMC); Rio Verde, 17-XI-1984, Bello (1 FVMC); X-1992, J Carlos (2 AMBC); XI-1993 (1
AMBC); XI-1995 (2 AMBC); Mato Grosso do Sul: Costa Rica, 17-XII-1993, S Ide (2 MZSP); Murtinho,
XII-1929, Malkin (1 MNRJ); Selvíria, UNESP farm, 01-V-1990, CAH Flechtmann, ex Guzerá bovine drop-
ping, Brachiaria decumbens pasture (1 FEIS); 01-VI-1991 (2 FEIS); 06-VII-1991 (1 FEIS); 06-VIII-1991 (1
FEIS); 13-XII-1992 (1 FEIS); 15-VI-1991 (2 FEIS); 16-II-1991 (1 FEIS); 18-III-1999, ex black light (2
FEIS); 18-V-1991, ex Guzerá bovine dropping (1 FEIS); 20-I-1999, ex black light (24 FEIS); 20-VIII-1991,
ex Guzerá bovine dropping (1 FEIS); 20-X-1991 (3 FEIS); 22-IV-1992 (2 FEIS); 23-II-1991 (3 FEIS); 23-II-
1992 (1 FEIS); 27-IV-1991 (1 FEIS); 31-X-1993 (1 FEIS); Terenos, 01-IV-1994, WW Koller (1 FVMC);
Mato Grosso: Chapada dos Guimarães, XI-1963, Alvarenga (1 MZSP); Minas Gerais: Águas Vermelhas,
XII-1997, Bello (3 AMBC); XII-1998, Bello & Vaz-de-Mello (1 FVMC); Cordisburgo, Faz Pontinha, X-
1993, FZ Vaz-de-Mello (1 CMNC, 2 BDGC); Ipatinga, XI-1992, E Grossi (1 FVMC); Montes Claros, I-2000,
JNC Louzada (1 FVMC); Três Marias, X-1989 (2 FVMC); Pará: Belém, IX-1964, E Dente (1 MZSP); Can-
indé (Rio Gurupi), X-1964, Malkin (1 CMNC); São Paulo: Bálsamo, 12-XI-1987, C Bergmann, seringueira
(1 IBSP); 12-XII-1987 (1 IBSP); 13-X-1988 (1 IBSP); 19-II-1987 (1 IBSP); 29-X-1987 (1 IBSP); Castilho,
marg. esq. Rio Paraná, 15-22-IX-1962, Exp. Dep. Zool. (1 IBSP); Itirapina, 15-IX-1996, JR Verdú (2 FVMC);
Itu, Faz Pau d'Alho, I-1959, Martins (1 CMNC); Mirante do Paranapanema, 09-X-1991, J Rodrigues, black
light, pasture area (2 FEIS); 13-XI-1991 (2 FEIS); Pradópois, XII-1976, PM Botelho (1 FVMC, 4 ESAP); São
Zootaxa 1955 © 2008 Magnolia Press · 29
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Carlos, 02-XII-1993, J Mendes (1 FVMC); 11-XI-1993 (1 FVMC); PARAGUAY: no data: (1 HMNH);
Anisits (1 HMNH); Boquerón: Pto Casado, XI-1950, Martínez (2 CMNC); Caaguazú: Caaguazú, Piscicul-
tura, II-1986, A Martínez (1 BDGC, 6 CMNC); Central: Asunción, 10-X-1904, Vezényi (1 HMNH); 28-IX-
1904 (1 HMNH); Concepción: Horquetá, XII-1950, Martínez, a la luz (6 CMNC); Paraguary: Sapucay, 05-
XI-1991, U Drechsel (9 ABC); San Pedro: Cororó, XI-1999, G Arriágada (1 FVMC); Rio Ypane, Cororó,
XI-1979, A Martínez (1 BDGC, 19 CMNC).
Diagnosis: 3.8–5.3 mm. Surface dull. Clypeal teeth obtuse and equilateral, continuous (at same plane)
with clypeal margin, separated by wide V-shaped emargination. Eyes, in dorsal view, visible and very small
(Fig. 55). Elytral striae lacking punctures. Very similar to G. margareteae, but lacking punctures in striae, and
anterior clypeal carina. Parameres as in Fig. 56.
Distribution: The entire Cerrado province of the Chacoan subregion, with invasion of pasturelands in
neighboring provinces (Fig. 104).
Remarks: This is a common pastureland species in Brazil, and may be related to G. margareteae.
6.2. Genieridium bordoni (Martínez, 1992), new combination
Pedaridium bordoni Martínez, 1992: Martínez 1992: 22–23; Vaz-de-Mello & Canhedo 1998: 100
Type series: Holotype
%
and allotype
&
: VENEZUELA: Barinas: Santa Bárbara, IV-1981, G&H Martínez
(BRBA).
Paratypes: VENEZUELA: Anzoátegui: Pariaguán, 09-VIII-1967, J&B Bechyné (1 CMNC); 12-VIII-
1967 (1 CMNC); Barinas: Santa Bárbara, IV-1981, G&H Martínez (1 BDGC, 9 CMNC); Guárico: Cal-
abozo, VI-1963, Bordón & Martínez, (1 CMNC); 15-VII-1962, Estación Biol. leg. (2 CMNC); Monagas:
Jusepín, IX-1965, F.Fernandez & CJ Rosales (1 BDGC); 500 m, 08-VIII-1966, CJ Rosales & F Fernández Y
(1 CMNC); 17-IX-1965 (1 CMNC); 21-IX-1965 (1 CMNC); 04-X-1965 (1 CMNC); 07-VIII-1966 (2
CMNC).
Non-type material examined: VENEZUELA: Anzoátegui: Aramina (Santa Clara), I-195, R Lichy (1
BDGC); Bolívar: 15 km E Caicara, 12-VI-1996, B Gill (20 BDGC); 12-13-VI-1996, H&A Howden (38
CMNC, 4 FVMC); 20 km SW Ciudad Bolívar, 19-VI-1987, S&J Peck, woodland on sand UV (1 CMNC); 25
km SW Pto. Ordaz, 21-VI-1987, S&J Peck, sandy woodland UV (1 CMNC); 8 km SW Caicara, 16-VI-1987,
S&J Peck, woodland UV (2 CMNC); Delta Amacuro: 15 km E Los Castillos, 4.5 km NE Ciudad Guyana,
26-IV-1987, MA Ivie, at light (1 BDGC, 11 CMNC); Guárico: Hato Masajuaral (44 km S Calabozo), 3-10-V-
1985, Menko & Carpenter (3 BDGC).
Diagnosis: 4.7–5.5 mm. Surface dull. Clypeal teeth lacking, but clypeal emargination clearly rounded,
widely V-shaped and with bottom declivitous (Fig. 57). Elytral striae not punctured. Male apical tooth of
metatibia flattened. Parameres as in Fig. 58.
Distribution: Maracaibo, Venezuelan Coast and Venezuelan Llanos provinces of the Caribbean subregion
(Fig. 107).
Remarks: This species is undoubtedly related to G. medinae, based on secondary sexual characters, but
shares with G. paranense the absence of clypeal teeth.
6.3. Genieridium cryptops (Arrow, 1913), new combination
Pedaridium cryptops: Arrow 1913: 458; Arrow 1932: 226; Balthasar 1938: 220; Blackwelder 1944: 203; Vulcano &
Pereira 1967: 577; Génier & Vaz-de-Mello 2002: 186–187; Hamel et al. 2006: 12.
Pedaridium mansosotoi Martínez, 1951: Martínez 1951: 35–40; Martínez 1959: 62; Ferreira & Galileo 1993: 7, 18–20;
Forsyth et al. 1998: 371; Koller et al. 1999: 405, 407; Vaz-de-Mello 2000: 194
VAZ-DE-MELLO
30 · Zootaxa 1955 © 2008
Magnolia Press
Pedarium [sic] cryptops: Bacchus 1978: 101.
Pedaridium bidens (misidentification, not Balthasar, 1938): Ferreira & Galileo 1993: 7, 15–16; Vaz-de-Mello 2000: 194
Type series: Pedaridium cryptops Arrow, 1913: Lectotype
&
(designated by Génier & Vaz-de-Mello, 2002):
BRAZIL: Goiás: Jatahy (BMNH)
Pedaridium mansosotoi Martínez, 1951: Holotype
%
: ARGENTINA: Formosa: Clorinda, 12-XII-1950,
A Martínez (BRBA); allotype
&
: PARAGUAY: ?: km 50 de Río Paraguay entre Asunción y B. Negal, 30-XI-
1950, A Martínez (BRBA).
Non-type material examined: BRAZIL: Bahia: Barreiras, X-1991 (1 FVMC); XII-1991, luz (3 FVMC);
Encruzilhada, 980 m, XI-1972, Alvarenga (11 MZSP, 3 IBSP, 2 FVMC); XI-1974, Alvarenga (1 CMNC);
XII-1980, Martínez & Alvarenga (2 CMNC); Distrito Federal: Brasília – Aeroporto, I-1964, Martínez (1
CMNC); Brasília 1100 m, III-2001, N Degallier (1 FVMC); XI-2000 (5 FVMC); XII-2000, luz (2 FVMC);
XI-1999 (1 CMNC); Est. Florestal Cabeça do Veado, 1100 m, 27-X-1971, EG, I & EA Munroe (11 CNCI); X-
1971, EG, I & EA Munroe (5 CNCI); RECOR-IBGE, 09-XII-1997, I Diniz (2 FVMC); XI-1999, M Mil-
homem, campo sujo (1 FVMC); Goiás: Aruanã, Rio Araguaia, II-1961, Dirings (1 MZSP); Bom Jardim de
Goiás, II-1997, FZ Vaz-de-Mello (1 FVMC); Campinas, XII-1925, Borgmeier & Lopes (1 MZSP); Goiatuba,
I-1953, J Guérin (1 IBSP); Jataí, Faz. Nova Orlândia, I-1964, Martins, Morgante & Silva (1 MZSP); Rio
Verde, XI-1993, J Carlos (1 AMBC); Mato Grosso do Sul: Costa Rica, 17-XII-1993, S Ide (10 MZSP);
Selvíria, UNESP farm, 02-II-1993, CAH Flechtmann, ex Guzerá bovine dropping, Brachiaria decumbens pas-
ture (1 FEIS); 09-XI-1994 (1 FEIS); Três Lagoas, margem esq. Rio Sucuriú, Faz. Canaã, I-1967, Lane (1
MZSP); Mato Grosso: Barra do Tapirapé, XI-1964, B. Malkin (4 MZSP); Chapada dos Guimarães, XI-1963,
Alvarenga (1 MZSP); Macaúba, XII-1966, R Schmitz (1 CMNC); Virapuru (sic) 160 km S, 8-10-III-1979, CR
Owen (2 HMNH); Minas Gerais: Águas Vermelhas, XII-1997, Bello (2 AMBC); XII-1998, Bello & Vaz-de-
Mello (3 FVMC); Cordisburgo, Faz Pontinha, I-1994, FZ Vaz-de-Mello (5 FVMC); I-1999 (1 FVMC); XII-
1993 (6 AMBC, 5 FVMC); Ibitira, XI-1988, luz (1 FVMC); Montes Claros, I-2000, JNC Louzada (30
FVMC); XII-1999 (2 FVMC); Paracatu, II-1997, S Lourenço (3 FVMC); XII-1996, S Lourenço (190 FVMC);
Paraopeba, 03-XI-1992, UV (1 FVMC); Serra do Caraça, 27-XI-05-XII-1972, Exp. Mus. Zool. (1 MZSP);
Três Marias, III-1990 (1 FVMC); X-1989 (3 FVMC); XII-1993 (1 FVMC); Unaí, Faz. Bolívia, 22-24-X-
1964, Exp. Dep. Zool. (1 MZSP); Piauí: São Raimundo Nonato - PN Serra da Capivara, I-1999, CA
Matrangolo (2 FVMC); São Paulo: Agudos, Duraflora SA, 07-XII-1993, CAH Flechtmann, P car. v. baha-
mensis log-baited tent trap, P oocarpa stand (1 FEIS); Bálsamo, 10-XII-1987, EC Bergmann, seringueira (1
IBSP); Boa Esperança do Sul, Faz. Itaquerê, 27-I-1964, K. Lenko (3 MZSP); Botucatu, 17-XI-1963, Manto-
vani (2 IBSP); Itu, Faz. Pau d' Alho, 1-5-I-1961, Martins (1 MZSP); Itu, II-1959, Martins (1 MZSP); Osasco,
Fca Fósforos, XII-1962 (2 CMNC); Pirassununga, 03-XI-1996, MA Ruiz Díaz (2 ESAP); Teodoro Sampaio,
Morro do Diabo State Reservation, 03-III-1993, CAH Flechtmann, ex bovine dropping baited pitfall trap (1
FEIS); 16-II-1993 (1 FEIS); 29-IX-1993 (1 FEIS); Toc an ti n s: Pium, XI-1971, J da Silva (2 CMNC); PARA-
GU AY: Amambay: Srra. Amambay, I-1960, Schultz (2 CMNC); Caaguazú: Caaguazú, XII-1977, Martínez
(1 CMNC); Concepción: Horquetá, XI-1950, Martínez (2 CMNC); San Pedro: Cororó, Rio Ypane, II-1979,
A Martínez (2 BDGC, 12 CMNC); II-1974 (1 CMNC); III-1979 (4 CMNC); XI-1979 (1 CMNC); XI-1979
(11 CMNC); Cororó, XI-1999, G Arriágada (1 FVMC).
Diagnosis: Surface shiny. Clypeal teeth long and acute, located bellow the surface of clypeal margin and
separated of clypeal disc by strong and sharp carina (the clypeal central margin itself, which is centrally
widely curved inwardly) (Fig. 60). Elytral striae with weak punctures. Elytral discal interstriae with either one
or two rows of setose punctures. Parameres as in Fig. 61.
Distribution: Cerrado, Caatinga and Chaco provinces of the Chacoan subregion (Fig. 105).
Remarks: This species is unusual in having either one or two rows of punctures on the discal interstriae;
it is the most variable in size within the genus. It appears to be related to G. zanunciorum, with which it shares
Zootaxa 1955 © 2008 Magnolia Press · 31
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
elongated clypeal teeth and shiny dorsal surface. The internal sac is unique by having a helicoid pseudoflagel-
lum. Like G. bidens, this species is a common pastureland species, easily attracted to lights.
6.4. Genieridium margareteae (Génier & Vaz-de-Mello, 2002), new combination
Pedaridium cryptops (misidentification, not Arrow, 1913): Ferreira & Galileo 1993: 7, 20–21; Vaz-de-Mello 2000: 194
Pedaridium margareteae: Génier & Vaz-de-Mello 2002: 192–193
Holotype
%
and allotype female: BRASIL: Piauí: São Raimundo Nonato, Parque Nacional da Serra da Capiv-
ara, I-1999, CA Matrangolo (MZSP).
Paratypes: BRAZIL: Bahia: Caetité, C. Uran. Lagoa Real - INB, 8–16.I.2000 (4 FVMC); Encruzilhada,
XII.1980/ A. Martínez e M. Alvarenga (3 CMNC); same as before except XII-1997, Vaz-de-Mello & Bello (4
FVMC); Jequié, II.1995, C. Sperber (1 FVMC); S. Antonio da Barra, XI–XII.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. Bello
(4 AMBC). Pernambuco: Pery-Pery, V–VI.1892, Gounelle (1 ZMHB, 1 FVMC); same as before except: XI–
XII.1892 (1 ZMHB). Piauí: São Raimundo Nonato, PN Serra da Capivara, I.1999, C.A. Matrangolo (5
FVMC). Rio Grande do Norte: Natal (5 NHMB); same as before except: W. M. Mann, Stanford Exped.,
1913-56 (1 BMNH); same as before except: III. 1952, Alvarenga (4 MZSP); III.1954, Alvarenga leg, Coll.
Martínez (1 CMNC).
Diagnosis: This species is almost identical to G. bidens, but differs by having the elytral discal striae
strongly punctured and the clypeal teeth weakly transversely carinated basally (Fig. 62). Parameres as in Fig.
63.
Distribution: Caatinga province of the Chacoan subregion, with one record from northeastern Pantanal
province of the Amazonian subregion (Fig. 106).
Remarks: This species is probably the sister to G. bidens, but apparently occupies a more restricted habi-
tat.
6.5. Genieridium medinae (Gill & Vaz-de-Mello, 2003), new combination
Pedaridium medinae: Gill & Vaz-de-Mello 2003: 47.
Type Material: Holotype
%
: COLOMBIA: Risaralda: PNR Ucumari, La Pastora, 2400 m Aliso, 07-V-1995,
C. Medina (IAHC); allotype female: COLOMBIA: Risaralda: PNR Ucumari 1800m, La Suiza, 29-III-1995,
C. Medina (IAHC).
Paratypes: COLOMBIA: Cundinamarca: Tecadama (sic) [Tequendama?] Falls, 30 km SW Bogotá, 27-
II-6-III-1972, S&J Peck, forest dung trap (1 CMNC); Quindio: 5 km E Salento, 1800 m, 9-XII-1995, BD Gill,
dung trap (9 BDGC; 1 CMNC; 4 FVMC); R Herencia Verde, 1800 m, 12-XII-1995, Medina & Gill, excr hum.
(1 CAMC); Risaralda: Pereira, SFF Otún Quimbaya, Est. La Suiza, 1850 m, 25-IV-04-V-1997, A Vitolo (1
FVMC); Pque. Nat. Reg. Ucumari, La Suiza 1800 m, CA Medina (1 CMNC); Pque. Nat. Reg. Ucumari, La
Suiza 1800 m, 29-III-1995, F Escobar, excr hum. (2 CAMC).
Diagnosis: Clypeal teeth triangular, obtuse, equilateral and continuous with clypeal margin. Eyes, in dor-
sal view, not visible. Elytral striae composed of contiguous large umbilicate punctures, elytral disc weakly
tectiform in the posterior half. Male metatibiae strongly modified (Figs. 66–67). Parameres as in Fig. 65.
Distribution: Central Colombia (Fig. 107).
Remarks: This is the only known species of Scatimina whose eyes are not exposed dorsally.
VAZ-DE-MELLO
32 · Zootaxa 1955 © 2008
Magnolia Press
6.6. Genieridium paranense (Arrow, 1932), new combination
Pedaridium paranense: Arrow 1932: 224–226; Balthasar 1938: 219; Blackwelder 1944: 203; Bacchus 1978: 106; Génier
& Vaz-de-Mello 2002: 191–192
Pedaridium paranensis [sic]: Ferreira & Galileo 1993: 9; Vaz-de-Mello & Canhedo 1998: 100; Vaz-de-Mello 2000: 194
Type series: Lectotype
%
(designated by Génier & Vaz-de-Mello, 2002): BRAZIL: Paraná: Castro, 1926
(BMNH)
Paralectotypes: Same data as lectotype (3 BMNH).
Non-type material examined: BRAZIL: Minas Gerais: Belo Horizonte, XI-1950, A Machado (1 MZSP);
Paraná: Curityba, XI-1941, Claretiano (1 CMNC); Ponta Grossa, VIII-1942, F Justus (5 CMNC); Castro (5
MZSP, 1 FVMC); 1907, E Garbe (4 MZSP); Londrina, XII-1935, B Pohl (1 FVMC); Rio de Janeiro: Estr
Rio-São Paulo km 47, XI-1944, Wygod (1 MNRJ); São Paulo: Cerqueira César, 12-X-1992, J Carlos (1
AMBC).
Diagnosis. Clypeal teeth absent (Fig. 68), as in G. bordoni. First elytral stria inconspicuous on anterior
half, stria 2 well defined for entire length, striae 3–5 inconspicuous posteriorly, stria 6 completely inconspicu-
ous. Parameres as in Fig. 69.
Distribution: Southeastern Brazil, Paraná Forest and Araucaria angustifolia Forest provinces of the
Parana subregion (Fig. 104).
Remarks: This rare species appears to be the sister taxon to G. bidens-margareteae; the clypeal structure
shared with G. bordoni is probably homoplasic.
6.7. Genieridium zanunciorum (Vaz-de-Mello & Canhedo, 1998), new combination
Pedaridium zanunciorum: Vaz-de-Mello & Canhedo 1998: 98–100; Vaz-de-Mello 2000: 194
Type series: Holotype
%
: BRAZIL: Minas Gerais: Santa Bárbara, 17-XI-1994, armadilha UV, Zanúncio
(MZSP).
Paratype: BRAZIL: Minas Gerais: Santa Bárbara, 22-X-1993, JC Zanúncio (FVMC).
Non-type material examined: BRAZIL: Goiás: Goiatuba, 1941, J Guérin (1 IBSP); Minas Gerais: Ara-
guari, II-1970, H Martínez (1 CMNC); São Paulo: Ypiranga, F. Ohaus (1 MNHU); Ipiranga, II-1927, Spitz (1
CMNC).
Diagnosis: Length 5.3–6.3 mm. Dorsum black, shiny. Clypeal teeth very acute, at same plane as lateral
margins and separated from them by weak emarginations (Fig. 70). In other aspects similar to G. cryptops,
except that interstriae always have biseriate setose punctures. Parameres as in Fig. 71.
Distribution: Scattered localities in Central and Southeastern Brazil (Fig. 106).
Remarks: This species appears to be associated with some special habitat; the few known specimens
were collected at light.
7. Leotrichillum new genus
Diagnosis: Small (2.2–2.8 mm), oval, and elongated body (Fig. 23); color tan to brown; clypeus with two
teeth and evenly rounded laterally (Fig. 72). Pronotum separated of hypomeron by sharp carina, hypomeron
with sharp lateral longitudinal carina posteriorly. Elytral interstriae with uniseriate setose punctures; pseu-
doepipleura glabrous and gradually narrowed posteriorly. Mesofemora and metafemora oval; mesotibiae and
metatibiae apical width of tibia much longer than third of tibial length. Metatarsi with basal tarsomere slightly
longer than second. Parameres with an apicolateral invagination (Fig. 73).
Zootaxa 1955 © 2008 Magnolia Press · 33
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Description: Length 2.2–2.8 mm, body oval, elongated, convex (Fig. 23); tan to brown, shiny, lacking
metallic sheen, dorsal setae slightly claviform. Clypeus with two short triangular teeth, separated by wide V-
shaped emargination, evenly curved laterally, continuous with genal margin. Clypeofrontal and clypeogenal
sutures indistinct. Eyes, in dorsal view, rounded, each about as wide as seven-tenths of its length; interocular
width five to six times eye width (Fig. 72). Pronotum lacking without anterior or posterior beads; lateral
pronotal callosity indicated by indistinct dark spot; disc covered by simple setose punctures, separated from
hypomeron by very weak carina. Hypomeron with strong longitudinal lateral carina; mesoepimera with weak
anterior transverse carina, stronger mesally; metasternal disc covered by simple punctures. Elytral interstriae
with one row of setose punctures, flat. Elytral striae not distinctly widened or deeper on declivity. Pseudoepi-
pleura glabrous, gradually narrowed to apex, with longitudinal fold near metacoxa, causing anterior part to be
almost vertical in position, and posterior part almost horizontal. Protibiae with three strong teeth distributed
along apical three-fifths of lateral margin, which is denticulate from base to basal tooth, with ventral scale-like
setae on teeth. Mesofemora and metafemora oval; mesotibiae and metatibiae apically wider than third of tibial
length. Pygidium nearly vertical. Parameres shorter than half length of phallobase, flattened and apically
rounded, laterally incised (Fig. 73). Pseudoflagellum helicoidal and thick, with flat elongated auxiliary
lamella. Coxites large, oval and symmetrical. Spermatheca simply C-shaped, gradually narrowed apically,
bulbous at base, with duct insertion at ventral part of the base of the spermathecal body.
Sexual dimorphism: Males with proclaws bent in the middle, protarsi with apical tarsomere modified to
receive claws; mesotarsi and metatarsi with claws bigger and stronger than in females. All tibiae with apico-
mesal tooth in males, mesotibial and metatibial teeth smaller than protibial one. Metasternum slightly concave
in males, flat in females; pygidium slightly longer in males than females.
Type species: Pedaridium louzadaorum Vaz-de-Mello & Canhedo, 1998 (monotypy) = Leotrichillum
louzadaorum (Vaz-de-Mello & Canhedo, 1998), new combination.
Etymology: After my son, Léo Falqueto Vaz de Mello. Gender neutral.
Distribution: Northern Argentina and southern Paraguay, central and northeastern Brazil, occupying the
Chaco, Caatinga and Cerrado provinces of the Chacoan subregion.
Remarks: The pygidial sulcus of L. louzadaorum is, in fact, not divided as described originally (Vaz-de-
Mello & Canhedo 1998); it is entire. The relationship of Leotrichillum to other genera is uncertain, as it has
several characters relating it both to Genieridium (e.g. oval and elongated body, presence of mesotibial and
metatibial mesoapical teeth in males, slightly modified mesotibial and metatibial male claws, weak pronotal-
hypomeral carina) and to Degallieridium (e.g. very small size, color tan to brown with darkened pronotal cal-
losities, claviform dorsal setae, longitudinally bent pseudoepipleura, large eyes). Unambiguous synapomor-
phies for Leotrichillum are the laterally incised parameres and the ventral position of spermathecal duct
insertion, apart of others that depend on the resolution of its phylogenetic position.
Composition: Besides the type species, the genus contains at least one undescribed species.
Material examined:
7.1. Leotrichillum louzadaorum (Vaz-de-Mello & Canhedo, 1998)
Pedaridium louzadaorum: Vaz-de-Mello & Canhedo 1998: 96–97, 100; Vaz-de-Mello 2000: 194
Type series: Holotype
%
: BRAZIL: Minas Gerais: Três Marias, XII-1993, Zanúncio (MZSP)
Paratypes: BRAZIL: Minas Gerais: Três Marias, X-1989 (1 BDGC); XII-1990, Zanúncio (1 CMNC);
XII-1990, Zanúncio (3 FVMC); XII-1993 (1 FVMC); IX-1994 (1 FVMC).
Non-type material examined: BRAZIL: Minas Gerais: Três Marias, X-1989 (1 FVMC); X-1994 (1
AMBC); XII-1990, Zanúncio (2 FVMC); XI-1993 (2 FVMC); Águas Vermelhas, XII-1997, Bello (1 AMBC),
Montes Claros, XII-1992, Zanúncio (1 FVMC); I-2000, JNC Louzada (2 FVMC); XII-1990 (1 FVMC);
Piauí: São Raimundo Nonato - PN Serra da Capivara, I-1999, CA Matrangolo (1 FVMC).
VAZ-DE-MELLO
34 · Zootaxa 1955 © 2008
Magnolia Press
8. Martinezidium new genus
Diagnosis: Clypeus with two, four or six teeth (Figs. 74, 76). Males without modified proclaws or apicomesal
protibial tooth. Elytral interstriae either with one or two rows of setose punctures; pseudoepipleura gradually
narrowed to apex and with row of setae. Mesotibiae and metatibiae longer than four times tibial apical width.
Parameres conical (Fig. 75).
Description: Length 2.8–5.0 mm. Body oval, elongated (Figs. 24–25), dorsally brown or black, some-
times with metallic sheen. Clypeus rounded, with two to six triangular teeth separated by wide U- or V-shaped
emargination. Clypeofrontal suture indistinct, clypeogenal suture very weak. Eyes, in dorsal view, as wide as
four- to seven-tenths eye length; interocular width about six to eight times eye width (Figs. 74, 76). Pronotum
lacking anterior or posterior beads, both anterior and posterior angles obtuse; separated from hypomeron by
strong lateral carina, hypomeron with strong posterior carina parallel to lateral border. Both anterior and pos-
terior pronotal angles obtuse, anterior margin not beaded. Mesoepimera with anterior transverse carina.
Protibiae with three strong lateral teeth, middle one broader, all acute; teeth distributed along apical three-
fifths; laterally denticulate from base to basal tooth; with ventral scale-like setae on teeth. Proclaws small,
evenly curved in both sexes. Metatarsi with basal tarsomere equal or shorter than second one. Interstriae flat
to feebly convex on disc, with one to two rows of setiferous punctures. Pseudoepipleura gradually narrowed
posteriorly, with setiferous punctures all along the mesal margin (poorly preserved or worn specimens lacking
in anterior third). Striae much deeper and with larger punctures apically. Aedeagus with each paramere coni-
cal (Fig. 75). Coxites small and symmetrical. Spermatheca C-shaped, with very long and swollen unsclero-
tized duct.
Sexual dimorphism: Females differ from males by having broader protibiae, lateral teeth slender and less
conspicuous than in males, protibial spur shorter and basally broader than in males; abdominal sternite VI
longer medially than in males, pygidium wider than in males.
Type species: Pedaridium galileoae Génier & Vaz-de-Mello, 2002 (present designation) = Martinezidium
galileoae (Génier & Vaz-de-Mello, 2002), new combination.
Etymology: Named after Antonio Martínez, from whose collection a great part of the examined
material came, and who contributed much to the study of Trichillum and Pedaridium from the 1950’s
to just before his death in 1993. Gender neutral.
Distribution: The genus has a very widely disjunct distribution, with one group in Southern Paraguay and
Argentina, from Jujuy to Chubut, occupying the Chaco and Pampa provinces of the Chacoan subregion in the
Neotropical region, and the Monte province in the South American transition zone; and the other in Mexico
(Jalisco, San Luis Potosí, Veracruz, Yucatán, Quintana-Roo, Chiapas, Campeche) and Guatemala (Petén),
equivalent to Mexican Gulf and Yucatán Peninsula provinces of the Caribbean subregion of the Neotropical
region, and the western part of Transmexican Volcanic Belt province of the Mexican Transition Zone.
Remarks: The relationship of this genus to other genera are unclear. Its disjunct distribution is correlated
to morphological differences indicating that the geographic groups likely correspond to species-group clades.
Northern species are black, without traces of metallic sheen, and clearly flatter than southern species, and with
slightly convex discal elytral striae (Fig. 25). Southern species are brown, with slight to very strong cupreous
to green metallic sheen, and have unique modifications in parameres mesally, making them asymmetrical
(Fig. 75). Apomorphies for this genus are the conical parameres and the lack of sexual differences in tarsi and
mesally in protibia.
Composition: Besides the type species, the genus contains also M. martinsi (Ferreira & Galileo, 1993),
new combination, M. fulgens (Arrow, 1913), new combination, M. maya (Vaz-de-Mello, Halffter, &
Halffter, 2004), new combination (all described as Pedaridium) and at least five new species currently being
described.
Material examined:
Zootaxa 1955 © 2008 Magnolia Press · 35
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
8.1. Martinezidium fulgens (Arrow, 1913)
Pedaridium fulgens: Arrow 1913: 458; Arrow 1932: 226; Blackwelder 1944: 203; Génier & Vaz-de-Mello 2002: 187–
188
Pedaridium martinezi Ferreira & Galileo, 1993: Ferreira & Galileo 1993: 30
Type series: Pedaridium fulgens Arrow, 1932: Lectotype and paralectotype
&&
(designated by Génier & Vaz-
de-Mello, 2002): ARGENTINA: Gran Chaco: Río Salado, E. Wagner (BMNH).
Pedaridium martinezi Ferreira & Galileo, 1993: Holotype not seen (not located).
Paratypes: ARGENTINA: Córdoba: Do. Santa Maria, Diquecito, XII-1965, Martínez (1 CMNC); La
Rioja: Olta, II-1934, González (1 CMNC).
Non-type material examined: PARAGUAY: Boquerón: km 145 a Puerto Casado, XI-1950, Martínez (1
CMNC).
8.2. Martinezidium galileoae (Génier & Vaz-de-Mello, 2002)
Pedaridium fulgens (misidentification, not Arrow, 1913): Balthasar 1938: 458; Martínez 1959: 62; Ferreira & Galileo
1993: 37; Monteresino et al. 1996: 107
Pedaridium galileoae: Génier & Vaz-de-Mello 2002: 195–196
Type series: Holotype
%
: ARGENTINA: Corrientes, Ituzaingó, Arenal de la Costa, Set. 975, Coll. Martínez;
allotype
&
, same data (CMNC).
Paratypes: ARGENTINA: Córdoba: Ciudad, I.1945, Coll. Martínez (1 CMNC); Do. Cruz del Eje, Gua-
naco Muerto, I.1977, Coll. Martínez (1 CMNC). Corrientes: Alto Paraná, Ituzaingó, XI.1975, Coll. Martínez
(1 FVMC); Ituzaingó, Arenal de la Costa, IX. 1975, Coll. Martínez (1 BDGC, 6 CMNC); Dº Ituzaingo, Villa
Olivari, Coll. Martínez (8 CMNC); same as before except: XII.1982 (1 BDGC). La Rioja: no locality,
XI.1959, M.J. Viana (1 CMNC); Olta, II.1934, M. Gómez leg., Coll. Martínez (1 CMNC). Mendoza: no
locality, Bruch (1 CMNC). San Luis: 18 km S. Arizona, 18–23.I.1982, 250m, H.& A. Howden (1 CMNC);
Desaguadero, II.2000, G. Arriágada (1 FVMC); San Geronimo, II.1974, M. Viana (2 CMNC).
Non-type material examined: ARGENTINA: San Luis: Desaguadero, 20-II.2000, G. Arriágada (1
FVMC).
8.3. Martinezidium martinsi (Ferreira & Galileo, 1993)
Pedaridium martinsi: Ferreira & Galileo 1993: 7, 23–24
Type series: not seen.
Non-type material examined: ARGENTINA: Neuquén: Ag. Florencio, Ruta 40, XII-1967, A Martínez (2
BDGC, 5 CMNC); Bajada Marucho, XII-1966, Gentili (1 CMNC); XII-1966, Martínez (4 CMNC); C Cura -
Buitres - 650 m, X-1968, Gentili (1 CMNC); Covunco, II-1976, Martínez (3 CMNC); La Pintada, XI-1957,
Schajovskoy (2 CMNC, 1 FVMC); Lotena-Granito, 800 m, X-1971, Gentili (1 CMNC); P del Águila, X-
1961, Grai (1 CMNC); Pampa del Saldo, III-1964, Gentili (1 CMNC).
8.4. Martinezidium maya (Vaz-de-Mello, Halffter, & Halffter, 2004), new combination
Pedaridium maya: Vaz-de-Mello, Halffter, & Halffter 2004: 247; Deloya & Peraza 2006: 341; Morón 2006: 122
Type series. Holotype
%
: MEXICO: Quintana-Roo, 5 km N Carrillo Puerto, IX-1984, A. Martínez (IEX),
Allotype
&
: México: Quintana-Roo, Reserva de Sian Ka'an, VIII-1984, NTP4A, M. A. Morón (IEX).
VAZ-DE-MELLO
36 · Zootaxa 1955 © 2008
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Paratypes: GUATEMALA: Petén, San Andrés, San Miguel La Palotada, 06-VI-1999, M. Tolon, EX1-L4-
KK3 (1 UVG); Tikal, 23-26-VIII-1972, S. & J. Peck (1 Halffter, 1 Vaz-de-Mello, 1 CMN); MEXICO:
Campeche, Chicana, 10 km W Xpujil, 300 m, 12-14-VII-1983, S. & J. Peck (1 CMN); Chiapas, Pque. Nal.
Sumidero, Coyote Mirador, 1700 m, 19-VI-1989, H. Howden (1 CMN); Pque. Nal. Sumidero, 26-V-1-VI-
1990, B. Gill, dung (2 BDGC); Pque. Nal. Sumidero, 21-VI-1989, E. Zuccaro & P. K. Lago (2 BDGC); Quin-
tana-Roo, 100' 20 mi S Felipe C Puerto, 13-VIII-1971, A. Newton (1 GVHC, 1 FVMC); 5 km N Carrillo
Puerto, IX-1984, Martínez (1 GVHC); Reserva de Sian Ka'an 1 km al NNE del Rancho "E124", 23-VIII-
1985, J. F. Camal, Trampa NTP 80 (1 GVHC); Reserva de Sian Ka'an 2 km al W del Rancho "El Ramonal",
23-VIII-1985, O. Canul, Trampa NTP 80 (1 Halffter); Reserva de Sian Ka'an 5 km del Crucero del camino
nuevo a Vigía Chico, 23-VIII-1985, O. Canul, Trampa NTP 80 (1 GVHC); Reserva de Sian Ka'an Bosque
Tropical Subcaducifolio, 10 m, 19-VI-1984, O. Canul & S. F. Camal, Trampa NTP 80 (2 GVHC, 1 FVMC, 1
MAMC); Reserva de Sian Ka'an km 12 Carretera nueva al Rancho "El Ramonal", 19-VI-1984, J. F. Camal,
Trampa NTP 80 (1 GVHC); 5 km N Puerto, IX-1984, Martínez (2 CMN); Reserva de Sian Ka'an 300 m S de
la unión de caminos nuevo y antiguo al Rancho Yuras, 10 m, 19-VI-1984, O Canul & SF Camal, selva tropical
subcaducifolia, NTP80 (1 LDC).
Non-type material examined: MEXICO: Quintana-Roo, Carrillo Puerto, IX-1984, A. Martínez (2
FVMC); Ver ac ru z: Apazapan, 280 m, IX-1999, E. Montes de Oca & Q. Santiago (1 EMOC); Jalcomulco,
450 m, IX-2000, E. Montes de Oca & Q. Santiago (1 EMOC).
9. Nunoidium, new genus
Diagnosis: Body dark, elongated (Fig. 26). Clypeus with two short triangular teeth separated by wide V-
shaped emargination. Eyes, in dorsal view, as wide as two-fifths of their length, interocular width seven to
eight times eye width (Fig. 77). Pronotum and hypomeron separated by strong carina, pronotum with posterior
bead. Mesofemora and metafemora oval, mesotibiae and metatibiae strongly widened apically. Male claws
and tarsi unmodified.
Description: Length 3.2–5.0 mm, body elongated, parallel sided (Fig. 26), color dark brown to black,
shiny, without metallic sheen. Clypeus with two triangular upturned teeth, separated by wide V-shaped emar-
gination, laterally rounded, margin continuous with that of gena. Clypeofrontal and clypeogenal sutures indis-
tinct. Eyes, in dorsal view, as wide as two-fifths of their length; interocular width seven to eight times eye
width (Fig. 77). Pronotum lacking anterior bead, but with uninterrupted posterior one; separated from
hypomeron by smooth carina; lateral callosity weakly defined, elongated; disc covered by setose punctures.
Hypomeron with strong lateral longitudinal carina; mesoepimera with anterior transverse carina; metasternum
covered by simple setose punctures on disc. Elytral discal interstriae (except sutural interstria) with biseriate
setose punctures, interstriae flat on disc, mesal ones slightly convex on declivity; striae deeper and wider on
declivity than on disc. Pseudoepipleura gradually narrowed posteriorly, with continuous row of punctures.
Protibiae with three strong teeth, distributed along apical three-fifths of length, denticulate at base, with ven-
tral scale-like setae on teeth. Mesofemora elongated, oval, as long as three times their medial width; metafem-
ora short, oval, as long as twice its medial width; mesotibiae and metatibiae strongly widened apically;
mesotibiae apically as wide as third of its length, metatibiae as long as twice apical width. Pygidium vertical,
transverse. Parameres flattened (Fig. 78), shorter than half length of phallobase, right-angled in relation to
phallobase. Internal sac with helicoid pseudoflagellum and three flattened elongated acessory lamellae. Cox-
ites symmetrical, rectangular, mesally toothed. Spermatheca very elongated, C-shaped, apically slightly spi-
raled.
Sexual dimorphism: Males with apicomesal tooth in all tibiae; metasternum slightly concave (flat in
females), abdominal disc flat (slightly concave in females) and pygidium less transverse.
Zootaxa 1955 © 2008 Magnolia Press · 37
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Type species: Pedaridium argentinum Arrow, 1913 (= Nunoidium argentinum (Arrow, 1913), new com-
bination).
Etymology: Named after my son, Nuno Falqueto Vaz de Mello. Gender neutral.
Distribution: Central, northern and northwestern Argentina, occupying about the Southern Chaco prov-
ince of the Chacoan subgregion (Fig. 108).
Remarks: Nunoidium appears to be closely related to Genieridium, however, the completeness of the car-
ina separating pronotal disc from hypomeron, vertical pygidial position, and presence of posterior pronotal
bead, among other characters, will readily separate it. Apomorphies of this genus are the presence of posterior
pronotal bead, absence of modification in male claws, elongation of spermatheca, and psammophilous modifi-
cations of metalegs.
Composition: Only the type species is known.
Material examined:
9.1. Nunoidium argentinum (Arrow, 1913)
Pedaridium rugiceps [lapsus]: Arrow 1913: 458
Pedaridium argentinum: Arrow, 1913: 459; Arrow 1932: 226; Balthasar 1938: 220; Blackwelder 1944: 203; Martínez
1959: 62; Martínez 1959: 63; Martínez 1987: 60; Ferreira & Galileo 1993: 8, 24–26; Monteresino et al. 1996: 107;
Génier & Vaz-de-Mello 2002: 188; Vaz-de-Mello & Génier 2005: 46
Trichillum elongatum Balthasar, 1939: Balthasar 1939: 24
Pedaridium elongatum: Martínez 1969: 119
Pedarium [sic] argentinum: Bacchus 1978: 99
Type series: Pedaridium argentinum: Lectotype
%
(designated by Génier & Vaz-de-Mello, 2002): ARGEN-
TINA: Chacho: Río Salado (BMNH), paralectotypes 4
%
3
&&
, same data as lectotype (3
%%
BMNH), same
data except: Icano (1
%
, 2
&&
BMNH, 1
&
MNHN).
Trichillum elongatum: Lectotype
%
(designated by Vaz-de-Mello & Génier, 2005), paralectotypes 2
&&
:
ARGENTINA: Cordoba (NMP).
Non-type material examined: ARGENTINA: Chaco: Gancedo, XII-1939, Biraben-Bezzi (1 CMNC);
Córdoba: San Vincente, J. Franzel S. (3 MNHU); 4 km NE Cruz del Eje, 20-II-1982, H&A Howden (7
CMNC); Cruz del Eje, I-1977, Martínez (4 CMNC); Do. Santa Maria, Diquecito, XII-1965, Martínez (2
CMNC); La Falda, I-1945, Martínez (2 CMNC); Formosa: Ing. Juarez, XII-1953, Martínez (1 CMNC);
Mendoza (1 MZSP); Salta: Do. San Martín, Hickmann, II-1949, Martínez (1 CMNC); San Luis: Do. Capi-
tal, San Gerónimo, II-1980, A Martínez (1 BDGC, 1 CMNC, 1 FVMC); Do. Ayacucho, El Milagro, XI-1966,
Martínez (1 CMNC); San Geronimo, XII-1972, GJ Williner (4 CMNC); Bruch (1 CMNC); 18 km S Arizona,
250 m, 18-23-I-1982, H&A Howden (1 FVMC); Santa Fé?: Carcarana (1 UNSM); Santiago del Estero: Ojo
de Agua, II-1974, A Martínez (1 BDGC, 4 CMNC); no data (9 MNHU, 5 FVMC); Ciudad, XI-1947, A
Amigo (1 CMNC); Frías, I-1949, Martínez (1 CMNC); Ojo de Agua, XI-1944, Maldonado (1 CMNC); Río
Salado, Wagner (2 CMNC); Santiago del Estero, Wagner (3 CMNC); 11-XII-1939, Biraben-Bezzi (1 CMNC);
Chaco de Santiago, Wagner (1 MZSP); no locality, 1936, Wagner (2 NHMB); Tuc um á n: Do. Capital, Rio
Sali, XI-1950, Martínez (1 CMNC); Province?: Between Santa Fe and Reconquista, 23-XII-1965, Hungarian
Soil-Zool. Exp (1 HMNH).
10. Onoreidium, new genus
Diagnosis: Body oval (Figs. 27–28). Head with clypeal teeth upturned and widely separated, emargination
shallow and very widely U-shaped; clypeal margin with sides rounded (separately from gena) or straight, giv-
VAZ-DE-MELLO
38 · Zootaxa 1955 © 2008
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ing clypeus a trapezoidal shape. Head disc strongly convex with clypeofrontal suture at least weakly elevated
and indicated laterally, frequently strongly carinated. Clypeal teeth separated by distance at least as wide as
half interocular width, generally much more (Figs. 79, 82–83). Mesotibiae, in ventral view, with strong single
tooth near middle of lateral border, with very strong short seta inserted apically; tibial ventral apical transverse
margin interrupted by similar structures and without longer setae (Fig. 81). Pseudoepipleura gradually nar-
rowed posteriorly (Fig. 6), elytral discal interstriae with two rows of setose punctures.
Description: Length 3.5–5.5 mm. Body oval (Figs. 27–28). Color grey to black, sometimes with strong
metallic sheen. Clypeus with two very widely separated triangular, upturned teeth, emargination shallow.
Clypeus rounded laterally (separately from gena) or straight, general clypeal form trapezoidal. Head disc
strongly convex, clypeofrontal suture well indicated, carinate at least laterally. Clypeogenal suture indicated at
least in front of each eye. Eyes, in dorsal view, elongated to oval, width half to fifth of eye length, interocular
width seven to 14 times eye width (Figs. 79, 82–83). Pronotum covered by elongated setose punctures; lack-
ing anterior and posterior beads; separated from hypomeron by very strong longitudinal carina. Hypomeron
with strong lateral longitudinal carina, mesoepimera with strong anterior transverse carina; metasternal disc
with setose punctures around middle. Elytra with flat interstriae, discal interstriae with biseriate punctures;
striae deeper and wider apically. Pseudoepipleura with uniseriate setose punctures and additional scattered
setose punctures on basal half. Protibiae with three teeth distributed along apical half or slightly less, without
ventral scale-like setae. Protarsi with apical tarsomere modified to receive claws. All claws evenly curved, but
with very sharp basal tooth. Mesotibiae (and metatibiae to lesser degree), in ventral view, with strong single
tooth approximately at midlength of lateral border, with very strong short seta inserted apically, this tooth
being similar to others interrupting ventral transverse apical carina of same tibia (Fig. 81). Mesotibiae and
metatibiae very strongly expanded laterally very near apex, about 3.5 to four times as long as wide at apex.
Parameres flattened apically (Fig. 80), shorter than half of phallobase length. Internal sac with very thick heli-
coidal pseudoflagellum. Coxites symmetrical, laminar, strongly toothed mesally. Spermatheca thin, C-shaped,
gradually narrowed apically, rounded basally, with very long partially sclerotized duct.
Sexual dimorphism: Males possess an apically directed small apicomesal tooth in protibiae; concave
metasternal disc (flat in females), flat shorter abdominal last sternite (longer and convex in females), and
longer pygidium.
Type species: Trichillum cristatum Arrow, 1931 (present designation) = Onoreidium cristatum (Arrow,
1931), new combination.
Etymology: After Giovanni Onore, who kindly sent me specimens for study and is a great activist on the
study of Latin American scarabaeoids. Gender neutral.
Distribution: Ecuador, northern Peru, and Panama, comprising the Arid Ecuador and Tumbes-Piura in
the Northwerstern South American dominion, and Eastern Central America, in the Mesoamerican dominion,
both in the Caribbean subregion. Cited from coastal Venezuela by Ferreira & Galileo (1993).
Remarks: Onoreidium has several apomorphies, such as, the unusual clypeal form, uniquely developed
lateral mesotibial and metatibial tooth, thick pseudoflagellum, and convex head. It appears to be the sister
genus of Pereiraidium, sharing with that genus the modified anterior tarsi in both males and females, strong
tooth in claws, and same general head shape. This clade appears to be related to the Trichillidium-Bradypodid-
ium clade (absence of ventral protibial scale-like setae, widely emarginated clypeus, strong lateral pronotal
carina, and relative apical position of protibial tooth); however, it is not clear if that is a sister-group relation-
ship, or if they form different grades of a paraphyletic group in relation to the remaining genera of Scatimina
with fused abdominal sternites.
Composition: Besides the type species, the new genus contains O. ohausi (Arrow, 1931), new combina-
tion (described as Trichillum), Onoreidium howdeni (Ferreira & Galileo, 1993), new combination, Onore-
idium bottimeri (Howden & Young, 1981), new combination (described as Pedaridium); and at least two
new species are under study.
Zootaxa 1955 © 2008 Magnolia Press · 39
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Material examined:
10.1. Onoreidium bottimeri (Howden & Young, 1981), new combination
Pedaridium bottimeri: Howden & Young 1981: 45; Ferreira & Galileo 1993: 6, 11–12; Ratcliffe 2004: 14; Harvey et al.
2006: 578.
Type series: Holotype
%
: PANAMA: Canal Zone: Barro Colorado Island, 13-VI-1963, LJ Bottimer (CNCI).
Non-type material examined: PANAMA: Panamá: Aeropuerto de Panamá, VII-1975, Martínez
(unknown
&
? CMNC, doubtful) (Fig. 83).
10.2. Onoreidium cristatum (Arrow, 1931)
Trichillum cristatum: Arrow 1931: 610; Paulian 1936: 206; Balthasar 1939b: 22; Blackwelder 1944: 204; Vulcano &
Pereira 1967: 578; Martínez 1969: 119; Bacchus 1978: 101; Génier & Vaz-de-Mello 2002: 191
Pedaridium equatoriensis Ferreira & Galileo 1993: Ferreira & Galileo 1993: 5–7, 14–15.
Type series: Trichillum cristatum: Lectotype
&
(Designated by Génier & Vaz-de-Mello, 2002): ECUADOR:
Loja: Piscobamba, M. Witt (BMNH); Paralectotype: ECUADOR: Loja, Ohaus S. 2200m (1 BMNH).
Pedaridium equatoriensis Holotype
&
: ECUADOR: Loja: Maracá/Catacocha, 1100 m, 14-VIII-1977, L Peña
(CMNC).
Non-type material examined: ECUADOR: Loja: Maracá/Catacocha, 1100 m, 14-VIII-1977, L Peña (1
BDGC, 4 CMNC); Río Catamayo, 29-VIII-1997, C Carpio (1 FVMC, 1 PUCE); PERU: Lambayeque: 28 mi.
E Olmos, 2000 m, 19-I-1955 (1 WDEC, 1 FVMC).
10.3. Onoreidium howdeni (Ferreira & Galileo, 1993)
Pedaridium howdeni: Ferreira & Galileo 1993: 8, 26–27
Type series: Holotype
&
: ECUADOR: Guayas: 40 km SW Guayaquil, 50 m, 21-22-II-1981, HF Howden
(CMNC).
Paratypes: ECUADOR: Guayas: 40 km SW Guayaquil, 50 m, 21-22-II-1981, HF Howden (1 MZSP); 45
km W Guayaque, 22-II-1981, HF Howden (3 CMNC).
Non-type material examined: ECUADOR: Guayas: 40 km SW Guayaquil, 50 m, 21-22-II-1981, B Gill (8
BDGC); HF Howden (3 CMNC); 22-II-1981 (42 CMNC, 1 FVMC, 4 LEMQ); Guayaquil, 50 m, 21-22-II-
1981, HF Howden (9 CMNC, 2 FVMC).
10.4. Onoreidium ohausi (Arrow, 1931)
Trichillum ohausi: Arrow 1931: 610; Paulian 1936: 206; Balthasar 1939b: 22; Blackwelder 1944: 204; Vulcano &
Pereira 1967: 578; Bacchus 1978: 106
Pedaridium ohausi: Martínez 1969: 119; Ferreira & Galileo 1993: 12; Génier & Vaz-de-Mello 2002: 190–191
Type series: Lectotype
%
(designated by Génier & Vaz-de-Mello, 2002): Ecuador: Loja: Punzara (BMNH);
Paralectotypes: Same data as lectotype except: Loja Calvario, 4.8.05 (1 BMNH), Loja (1 BMNH), Ecuador,
no locality (2 BMNH).
Non-type material examined: ECUADOR: Loja: Loja, Ohaus (2 NHMB, 1 FVMC); III-1965, L Peña (1
BDGC, 6 CMNC, 1 NHMB); XII-1984, P. Ponce (1 FVMC); Abé Gaujon (1 NHMB, 1 FVMC).
VAZ-DE-MELLO
40 · Zootaxa 1955 © 2008
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11. Pedaridium Harold, 1868
Diagnosis: Body very large (5.2–8.6 mm) and elongated (Fig. 29). Pronotum clearly flat to concave posteri-
orly (Fig. 87). Elytra tectiform (suture elevated in relation to disc, each elytron separately flattened), disc
forming a weak fold at apex; interstriae with biseriate setose punctures (Fig. 29). Males with proclaws
strongly bent (Fig. 86). Scattered longer setae present in sides and anterior part of pronotum and elytral apex
(Fig. 29).
Description: Length 5.2–8.6 mm, body very elongated and parallel-sided (Fig. 29), color brown to dark
grey, completely opaque and lacking metallic sheen, with yellow to orange setae. Clypeus with two teeth, sep-
arated by wide shallow emargination rounded at bottom; laterally unequally rounded, mesally strongly so and
laterally straight, continuous to genal margin. Head flat, feebly depressed near eyes. Clypeofrontal and cly-
peogenal sutures indistinct. Eyes, in dorsal view, as wide as half their length; interocular width ten to 15 times
eye width (Fig. 84). Pronotum lacking anterior and posterior beads, separated from hypomeron by distinct
longitudinal carina; disc covered by setose puntures, setae much longer anteriorly and laterally; disc surface
with posterior shallow to deep concavity, projected anteriorly along midline; and slightly expanded (shallowly
concave) on anterior angles. Pronotal lateral outline sinuate, wider at anterior angles than at middle (Fig. 87).
Hypomeron with distinct longitudinal carina; mesoepimeron with distinct transverse anterior carina. Elytral
discal interstriae with two rows of setiferous punctures, flat. Elytra tectiform for entire length, with sutural
interstria clearly elevated and each elytron flat on disc; apically with lateral fold between declivity and pseu-
doepipleuron. Striae clearly deeper and wider at apex. Pseudoepipleura gradually narrowed to apex (Fig. 8),
with row of large setiferous punctures along entire length and smaller irregular setae in the anterior half.
Protibiae with three teeth distributed along apical half of lateral margin, basally denticulate and with small
scale-like setae on the ventral face of teeth only. Mesofemora and metafemora elongate, mesotibiae and metat-
ibiae four times as long as wide apically. Pygidium transverse, vertical, last abdominal sternite long. Phallo-
base very thin, parameres as long as third of phallobase length, apically flattened (Fig. 85). Internal sac with
very thin canaliculated straight pseudoflagellum, with basal curved auxiliar elongated lamella. Spermatheca
C-shaped on apical half, gradually narrowed apically, basally strongly narrowed, pointed (Fig. 12), duct not
sclerotized. Coxites minute, symmetrical and multidentate apically.
Sexual dimorphism: Males have proclaws strongly bent (Fig. 86), a very strong mesoapical protibial
tooth; small mesotibial and metatibial mesoapical teeth (Figs. 1–2); metasternal disc strongly concave (flat in
females), abdominal disc flat (convex in females), and less transverse pygidium than in females.
Type species: Pedaria hirsuta Harold, 1859 (implicit monotypy) = Pedaridium hirsutum (Harold, 1859).
Distribution: Brazil (Minas Gerais, São Paulo, Rio de Janeiro, and Paraná states). Present only in the
Eastern Parana subregion, in the Parana Forest and Araucaria angustifolia Forest provinces.
Remarks: Pedaridium species are the only representatives of the group that really have a body shape sim-
ilar to that of species in the African genus Pedaria. Synapomorphies shared by species of this genus are the
unique form of the pronotum and elytra; dual-length dorsal pilosity; reduction of number of internal sac scler-
ites; reduction of coxite size; and elongation of phallobase (see also under Genieridium).
Composition: In the present restricted sense, Pedaridium is monotypic. However, it contains at least one
undescribed species currently under study, in addition to the type species.
Material examined:
11.1. Pedaridium hirsutum (Harold, 1859)
Pedaria hirsuta: Harold 1859: 194–195
Pedaridium hirsutum: Harold 1869a: 1001; Gillet 1911: 48; Arrow 1913: 458; Arrow 1932: 224, 226; Balthasar 1938:
219; Blackwelder 1944: 203; Ferreira & Galileo 1993: 7, 16–18, 48–51; Vaz-de-Mello & Canhedo 1998: 100; Vaz-
de-Mello 2000: 194
Zootaxa 1955 © 2008 Magnolia Press · 41
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
Type series: Lectotype is here designated in order to maintain nomenclatoral stability (see remarks). LECTO-
TYPE here designated:
%
, pinned, in MNHU. Labels: [1.] 26438 / [2.] São João del Rey, Sellow / [3. green
label] hirsutum Harold / [4.] Zool. Mus. Berlin / [5. red label] LECTOTYPE / [6.] Pedaria hirsuta Har. LEC-
TOTYPE, Vaz-de-Mello det. 2001
PARALECTOTYPE:
%
, pinned, in MNHU. Labels: [1. green label] St Joao d R, Sello, Nr 26438 / [2.]
Zool. Mus. Berlin / [3. yellow label] PARALECTOTYPE / [4.] Pedaria hirsuta Har. PARALECTOTYPE,
Vaz-de-Mello det. 2001.
Non-type material examined: BRAZIL: Minas Gerais: Vila Monte Verde, 20-IV-1966, J. Halik (4
MZSP); Paraná: Campo Largo da Roseira, VII-1942, Hatsbach (1 CMNC); Curitiba, V-1942, Hatsbach (1
CMNC); no date (1 MNRJ); IV-1942 (1 IBSP, 3 MZSP); Deodoro, 1943, Hatsbach (2 CMNC); VI-1942 (1
CMNC, 3 MZSP); Florestal (Deodoro), X-1943, Hatsbach (1 CMNC); Florestal (Piraquara), XII-1942, B.
Pohl (2 MZSP); Guaraúna, VI-1937, Pinheiro Machado (1 MZSP); VI-1947 (1 MNRJ); Penha, VI-1940,
Hatsbach (1 MZSP); Purunã, V-1942, Hatsbach (1 MZSP); Rincão, IV-1942, Hatsbach (1 CMNC); Xaxim, V-
1942 (1 IBSP, 1 MZSP); Rio de Janeiro: Itatiaia, 19-IV-1992, CL Godinho Jr (2 FVMC); I-1992 (1 FVMC);
IV-1995 (2 FVMC); Nova Friburgo, Três Picos, Salinas, X-2000, P Grossi (4 FVMC); Nova Friburgo, VI-
2000, P Grossi (1 FVMC); Santa Catarina: São Bento, II-1989, Pereira (1 CMNC); São Paulo: São Ber-
nardo, VIII-1934, Guérin (2 IBSP); No data: (2 IRSN, 1 NMHB); Candèze (2 IRSN).
Remarks: The designation of a lectotype for P. hirsutum is necessary because of the sympatry and great
resemblance of that species to another yet undescribed one. Although the two examined specimens that could
be confirmed to belong to the type series are conspecific, Harold (1859) does not mention the number of spec-
imens seen by him for the description, and the original description fits both species. So, given the possibility
of existence of other syntypes of P. hirsutum, which may not be conspecific to the ones studied, I decide to
here fix the name-bearing type in order to fix the application of the species name in its current sense.
12. Pereiraidium, new genus
Diagnosis: Large species (5.5–6.1 mm), sides of head sinuate, clypeogenal margin not incised (Figs. 88, 90).
Male with two long frontal horns (Figs. 30, 88), females with two poorly defined tubercles (Figs. 31, 90).
Pronotum with anterior bead distinct. Claws strongly toothed at base; protarsi with apical tarsomere modified
in both sexes.
Description: Length 5.5–6.1 mm, body oval, elongated (Figs. 30–31), color dark brown, opaque, lacking
metallic sheen. Clypeus with two large, weakly defined teeth, separated by wide V-shaped emargination.
Clypeus laterally sinuated, continuous with genal margin. Clypeofrontal and clypeogenal sutures indistinct;
frons mesally with longitudinal concavity separating frontal structures (each side with either long horn or con-
vexity, mesad to eyes), and transverse concavities anterior to eyes. Eyes, in dorsal view, wider than half of
their length; interocular width approximately six times eye width (Figs. 88, 90). Pronotum with distinct ante-
rior bead (in some cases indistinct mesally), but lacking posterior one; lateral callosity indistinct; disc with
setiferous punctures. Pronotum separated from hypomeron by distinct longitudinal carina. Hypomeron later-
ally with distinct longitudinal carina, mesoepimeron with distinct transverse carina. Elytral interstriae convex,
each with two widely separated rows of setiferous punctures. Striae much deeper and wider on apical decliv-
ity. Pseudoepipleura gradually narrowed posteriorly, with uniseriate setiferous punctures along length and
sparse setiferous punctures ventrally on the anterior half. Protibiae with three strong teeth, located on apical
half, ventrally lacking scale-like setae. Mesotibiae and metatibiae longer than three times their apical width.
Pygidium vertical. Parameres apically flattened (Fig. 89), as long as third of phallobase length. Internal sac
with strong sinuated and bent pseudoflagellum and two auxiliary lamellae. Coxites small, symmetrical and tri-
angular. Spermatheca very elongated, slightly spiralled apically, gradually narrowed at both ends.
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Sexual dimorphism: Males with strong frontal, upturned horns (Figs. 30, 88) that can be as long as, or
slightly longer than prothorax; horns with hairs similar to those on remainder of head at least on basal half
(Fig. 88); females with only two distinct convexities in place of horns (Fig. 90). Pronotum of large males with
flat anterior declivity (Fig. 30), declivity absent or very slightly indicated in females (Fig. 31). Metasternum
concave in males and flat in females; and pygidium slightly longer in males.
Type species: Pedaridium almeidai Pereira, 1946 = Pereiraidium almeidai (Pereira, 1946), new combi-
nation.
Etymology: The name is after Padre Francisco Silvério Pereira, CMF (1912–1992), Brazilian scara-
beidologist who described the single species here included in this genus. Gender neutral.
Distribution: South and southeastern Brazil (Rio Grande do Sul, Paraná and São Paulo). Endemic to the
southernmost part of the Brazilian Atlantic Forest province (Fig. 108).
Remarks: The main apomorphies of this genus are the presence of horns in males and two frontal con-
cavities in females, an anterior pronotal bead, and convex discal interstriae. It may be diagnosed also by the
absence of the synapomorphies supporting Onoreidium, its sister-genus. See more remarks under Onoreidium.
Composition: Only the type species is known.
Material examined:
12.1. Pereiraidium almeidai (Pereira, 1946), new combination
Pedaridium almeidai: Pereira 1946: 289; Ferreira & Galileo 1993: 6, 10–11; Vaz-de-Mello 2000: 194; Verdú & Galante
2001: 597–599
Type series: Holotype
%
and allotype
&
: BRAZIL: Paraná: Deodoro, V-1942, Hatsbach (MZSP).
Paratypes: BRAZIL: Paraná: Guaraúna, V-1937, J. Pinheiro Machado (1 CMNC); XII-1937 (1 CMNC);
Deodoro, V-1942, Hatsbach (2 MZSP); Rio Grande do Sul: Glória, 1927, P. Buck (1 IRSN).
Non-type material examined: BRAZIL: Rio Grande do Sul: Glória, 06-VI-1927, P. Buck (2 CMNC); 20-
IX-1927 (1 CMNC); X-1927 (1 NMHB); 26-VII-1928, P Buck (1 FVMC); São Paulo: Est Biol. Boracéia, 24-
VIII-1996, JR Verdú (2 FVMC).
13. Scatimus Erichson, 1847
Diagnosis: Head with transverse carina or conical tubercle (Fig. 32). Dorsum lacking setiferous punctures.
Abdomen with intersternal sutures clearly visible. Trochanterofemoral pit of anterior leg rounded. Mesotibiae
and metatibiae each with two transverse lateral carinae.
Type species: Scatimus cucullatus Erichson, 1847 (monotypy).
Distribution: From Sonora province in the Continental Nearctic dominion of the North American Pacific
subregion of the Nearctic region to Napo, Guyana and Roraima provinces of the Amazonian subregion of the
Neotropical region.
Composition: This genus contains twelve species and has been recently revised by Génier and Kohlmann
(2003). Valid species and their synonyms are:
13.1. Scatimus cribrosus Génier & Kohlmann, 2003
13.2. Scatimus cucullatus Erichson, 1847
13.3. Scatimus erynnios Kohlmann & Solís, 1996
13.4. Scatimus fernandezi Martínez, 1988
13.5. Scatimus furcatus Balthasar, 1939
13.6. Scatimus monstrosus Balthasar, 1939
Zootaxa 1955 © 2008 Magnolia Press · 43
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
13.7. Scatimus onorei Génier & Kohlmann, 2003
13.8. Scatimus ovatus Harold, 1862
=Scatimus patruelis Preudhomme de Borre, 1886
=Scatimus quadridentatus Balthasar, 1939
13.9. Scatimus pacificus Génier & Kohlmann, 2003
13.10. Scatimus quadricuspis Génier & Kohlmann, 2003
13.11. Scatimus simulator Martínez, 1988
13.12. Scatimus strandi Balthasar, 1939
14. Scatrichus Génier & Kohlmann, 2003
Diagnosis: Head with two parallel transverse carinae (Fig. 33). Pronotum with sparse setiferous punctures lat-
erally; elytra with setiferous punctures on apex and pseudoepipleura. Abdomen with sternites clearly differen-
tiated on disc. Trochanterofemoral pit of anterior leg oval. Mesotibiae and metatibiae each with two transverse
lateral carinae ventrally.
Type species: Scatimus bicarinatus Harold, 1869 (original designation).
Distribution: Cerrado province of the Chacoan subregion, and Brazilian Atlantic Forest and Parana For-
est provinces of Parana subregion.
Composition: Scatrichus contains three species and has been recently revised by Génier & Kohlmann
(2003) and this work is here refered to. Recognized species are:
14.1. Scatrichus bicarinatus (Harold, 1869)
= Scatimus bicarinatus Harold, 1869
14.2. Scatrichus goiasensis Génier & Kohlmann, 2003
14.3. Scatrichus sulcifer Génier & Kohlmann, 2003
15. Silvinha, new genus
Diagnosis: Dark brown to black, shiny, with sparse setae only on sides and apices of elytra (Fig. 34); clypeal
teeth acute, emerging from below clypeal margin, which is sinuate mesally (Fig. 91). Pseudoepipleura
strongly narrowed posteriorly, forming sharp angle near metacoxal apex (Fig. 9). Aedeagus with lateroapical
region of parameres expanded and curved inwardly (Fig. 92).
Description: Length 3.5–3.8 mm, body form elongate, oval (Fig. 34), dark brown to black, shiny, without
traces of metallic sheen. Clypeus with two very acute teeth, separated from clypeus by transverse carina
delimiting clypeal margin; clypeal margin between teeth simply sinuate in the middle, with rounded to trun-
cate lobe over each clypeal tooth; sides straight to lateral genal angle. Clypeofrontal suture inconspicuous,
indicated only laterally, clypeogenal suture inconspicuous. Eyes, in dorsal view, as wide as half of their
length, not narrowed posteriorly; interocular width approximately eight times eye width (Fig. 91). Pronotum
lacking anterior and posterior beads, with lateral callus very feebly indicated; separated from hypomeron by
strong longitudinal carina. Hypomeron with strong lateral longitudinal carina, mesoepimeron lacking anterior
transverse carina; metasternal disc smooth. Elytral interstriae on disc lacking rows of punctures and setae, flat;
very few (at most three) setose uniseriate punctures present only apically. Mesal striae with enlarged punc-
tures only apically. Pseudoepipleura without distinct separation from disc anteriorly, glabrous, sinuated near
metasternal lateral lobe, strongly narrowed and strongly angulate near metacoxa (Fig. 9). Protibiae with three
lateral teeth, distributed along apical half, not denticulate basally, with ventral scale-like setae. Mesotibiae and
metatibiae gradually widened to apex, apex as wide as fourth of their length. Metatarsi with basal tarsomere as
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long as 1.3 times the second one. Abdomen with pygidium vertical, transverse. Parameres smaller than half of
phallobase length, laminate, divergent mesally and strongly convergent apically (Fig. 92). Internal sac with a
three-folded pseudoflagellum and two acessory lamellae. Coxites small, symmetrical, and triangular. Sper-
matheca C-shaped, bulbous at base and gradually narrowed apically, apex inwardly curved.
Sexual dimorphism: Males have protibiae with triangular apically and ventrally directed apicomesal
tooth; the apical tarsomere modified to receive claws, and protarsi with ventral scale-like setae; protibiae slen-
der and gently curved inwardly (almost straight and wider in females), metasternum concave (nearly flat in
females), last abdominal sternite shorter than in females, and pygidium less transverse.
Type species: Silvinha unica new species (monotypy).
Composition: Monobasic.
Etymology: Named after my wife, Silvia (Silvinha) Altoé Falqueto. The name is feminine in gender.
Distribution: Southeastern Brazil (São Paulo, Minas Gerais, and Rio de Janeiro states). Endemic to the
southeastern part of the Parana Forest province in the Parana subregion (Fig. 108).
Remarks: This genus is related to Trichillum, Besourenga, Eutrichillum, Degallieridium, and Feeridium,
although its exact position in that group is not resolved yet. Its main autapomorphies are the absence of orga-
nized punctation in elytral discal interstriae, uniquely shaped pseudoepipleura and clypeal teeth. See more
remarks under Besourenga.
15.1. Silvinha unica new species
Type series: Holotype
%
: BRAZIL: Rio de Janeiro: Nova Friburgo, VII-1994, P Grossi (IBSP ex-FVMC).
Paratypes: BRAZIL: Minas Gerais: Viçosa, X-1998, FZ Vaz-de-Mello (2 FVMC); II-1995, FZ Vaz-de-
Mello (1 FVMC); Rio de Janeiro: Nova Friburgo, VII-1994, P Grossi (1 FVMC); VI-1994, FZ Vaz-de-Mello
(1 CMNC); 1000 m, XII-1996, P&E Grossi (2 FVMC); São Paulo: Serra do Japi, 1050 m, floresta, 1998,
MIM Hernández, armadilha pitfall com fezes (5 FVMC).
Description: Holotype
%
3.6 mm. Body color very dark brown. Head covered by simple punctures sepa-
rated by one to three diameters, clypeus and gena with few intermixed larger setose punctures, clypeus anteri-
orly with also minute punctures intermixed (Fig. 91). Pronotum covered by simple round punctures, on disc
punctures separated by three to five diameters and minute in size, laterally punctures much larger and denser.
Hypomeron covered posteriorly by large round punctures separated by four to six diameters. Elytral inter-
striae covered by minute unorganized punctures; discal striae with large round punctures, about as wide as
twice the strial width, and separated by about four diameters on disc. Mesosternum covered by large trans-
verse ocellated punctures separated by less than one diameter, slightly larger than similar punctures present on
metasternal lateral lobes. Protibiae with anteriorly and ventrally directed triangular apicomesal tooth. Abdom-
inal disc with ocellate punctures separated by one to three diameters. Pygidium with scattered setose and
minute punctures intermixed. Parameres as in Fig. 92.
Variation: Paratypes vary only in color (brown to nearly black), size (3.4–3.8 mm) and sexual features
(see above).
Etymology: unica: unique, only one, referring to only one species known in the genus.
16. Trichillidium, new genus
Diagnosis: Clypeus with four variably-shaped teeth, inner pair separated by U-shaped emargination. Laterally
to outer teeth clypeal margin concave, continuous with that of gena (Figs. 93, 95, 97). Elytral striae very
strongly impressed apically. Pseudoepipleura gradually narrowed apically, with uniseriate setose punctures
and irregular ventral punctures in the anterior half. Protibiae with two to three apical teeth; if three then basal
Zootaxa 1955 © 2008 Magnolia Press · 45
SYNOPSIS OF THE NEW SUBTRIBE SCATIMINA
one very small; teeth distributed along apical half or less of tibial margin length. Mesotibiae and metatibiae
with dense setae apically and ventrally.
Description: Length 2.7–4.1 mm. Body short oval, with maximum width mesally on elytra (Figs. 35–36);
color grey to black, sometimes with slight metallic sheen. Clypeus with four teeth, inner two separated by
wide U-shaped emargination. Laterally to outer teeth, , clypeal margin continuous with genal margin, con-
cave; genal lateral angle projected. Head uniformly convex, with shallow concavities anterior to eyes. Clypeo-
frontal suture distinct only laterally, clypeogenal sutures indistinct. Eyes, in dorsal view, as wide as third to
half of their length; interocular width eight to 12 times eye width (Figs. 93, 95, 97). Pronotum lacking anterior
or posterior beads, laterally with callosity only very feebly indicated; hypomeron separated from pronotum by
strong longitudinal carina. Hypomeron with strong lateral longitudinal carina; mesoepimera anteriorly with
transverse carina indistinct; metasternal disc covered by small setose punctures. Elytral interstriae flat, with
biseriate setose punctures. Striae moniliform, with large coalescent punctures, which are strongly widened and
deeper at apex. Pseudoepipleura gradually narrowed apically, with uniseriate setose punctures. Protibiae with
two to three teeth (if three then basal one very small), distributed along less than half of tibial length; scale-
like setae lacking ventrally. Mesotibiae and metatibiae slender, apical width less than third of tibial length,
with very dense setae apically and ventrally, surface difficult to see. Protarsomeres and claws simple. Pygid-
ium elongated, inclined, not exactly vertical. Internal sac with large, thick pseudoflagellum. Coxites large,
laminate, triangular and symmetrical. Spermatheca C-shaped, base bulbous, gradually narrowed to acute
apex.
Sexual dimorphism: Males with small apicomesal tooth in protibiae, scale-like setae ventrally in pro-
tarsi, metasternal disc convex (concave in females), and abdominal disc shorter than in females.
Type species: Pedaridium quadridens Arrow, 1913 = Trichillidium quadridens (Arrow, 1913), new com-
bination.
Etymology: Combination of Trichillum and Pedaridium, due to the fact that species included have been
originally described in these different genera. This name was also found in some of A. Martínez's identifica-
tion labels. Gender neutral.
Distribution: Southern Nicaragua to Central Argentina, absent in Western Amazonia, Central and North-
eastern Brazil. Exhibits disjunct distribution comprising southern Mesoamerican dominion (Eastern Central
America and Western Panamanian Isthmus provinces) and Choco and Western Ecuador provinces in North-
western South American dominion, in the Caribbean subregion; Amapa and southern Pantanal provinces in
the Amazonian subregion; Chaco and Pampa provinces in Chacoan subregion, and Parana Forest in Parana
subregion.
Remarks: This genus is undoubtedly related to Bradypodidium (q.v.), and is defined by the following
apomorphies: projection of lateral part of gena; convexity of head disc, and shallow concavities anterior to the
eyes. This genus has very variable parameral shapes (e.g., Figs. 94, 96).
Composition: Apart of the type species, this genus includes T. brevisetosum (Howden & Young, 1981),
new combination (described as Pedaridium), T. caingua (Martínez, 1974), new combination (described as
Pedaridium with a question mark), T. pilosum (Robinson, 1948), new combination (described as Trichillum),
and at least one or two species yet undescribed.
Material examined:
16.1. Trichillidium brevisetosum (Howden & Young, 1981), new combination
Pedaridium brevisetosum: Howden & Young 1981: 44; Ferreira & Galileo 1993: 8, 32; Ratcliffe 2004: 14
Type series: Holotype
%
: PANAMA: Canal Zone: Barro Colorado Island, 16-VI-1977, HA Hespenheide
(CMNC).
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16.2. Trichillidium caingua (Martínez, 1974)
Pedaridium (?) caingua: Martínez 1974: 65
Pedaridium caingua: Ferreira & Galileo 1993: 8, 31–32
Type series: Holotype
&
: ARGENTINA: Misiones: P.N. Iguazú, Pto. Iguazú, XII-1958, Martínez (BRBA).
Paratype: ARGENTINA: Misiones: Do. Frontera, San Antonio, IX-1957, Martínez (1 CMNC).
Non-type material examined: BRAZIL: Rio de Janeiro: Miguel Pereira, XI-1997, J Carlos (1 AMBC);
Santa Catarina: Nova Teutônia, XI-1975, F Plaumann