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The species of the Deltochilum subgenus Aganhyboma, endemic to Central and South America, are reviewed. The species have been divided into complexes, with a brief diagnosis presented for each. A key for identification of 26 currently recognized species is provided in English and Portuguese. Two major groups within the subgenus are recognized (trisignatum and valgum). The first is represented by species typically belonging to the subgenus Aganhyboma (Deltochilum (A.) trisignatum, D. (A.) kolbei, D. (A.) violaceum, D. (A.) cupreicolle and D. (A.) viridescens new status) and three new species described here (D. (A.) amandaarcanjoae, D. (A.) viridicatum and D. (A.) titovidaurrei). The second group (valgum) is represented by: D. (A.) valgum, D. (A.) longiceps new status, D. (A.) acropyge new status, D. (A.) acanthus, D. (A.) icaroides and D. (A.) icariforme), previously assigned to the subgenus Deltohyboma. Twelve new species are described for the valgum group (D. (A.) schefflerorum; D. (A.) streblopodum; D. (A.) feeri; D. (A.) larseni; D. (A.) arturoi; D. (A.) finestriatum; D. (A.) cangalha; D. (A.) alpercata; D. (A.) ritamourae; D. (A.) kolleri; D. (A.) paresi; D. (A.) subrubrum). A lectotype is designated for D. (A.) trisignatum and D. (A.) icariforme. A detailed literature review, synonymies, description, illustration of key morphological characters, data of the studied material and geographic distribution is provided for each species.
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Accepted by S. Tarasov: 30 Jan. 2015; published: 3 Mar. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 3925 (4): 451
504
www.mapress.com
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http://dx.doi.org/10.11646/zootaxa.3925.4.1
http://zoobank.org/urn:lsid:zoobank.org:pub:69DADAD6-84DA-4CBE-B653-6527293BA560
A revision of the Deltochilum subgenus Aganhyboma Kolbe, 1893
(Coleoptera: Scarabaeidae:
Scarabaeinae)
FERNANDO A. B. SILVA
1
, JÚLIO LOUZADA
2
& FERNANDO VAZ-DE-MELLO
3
1
Universidade Federal do Pará, Instituto de Ciências Biológicas, setor de Zoologia. Rua Augusto Corrêa, 01. Guamá. Belém PA
66075-110, Brazil. E-mail: fernandoabsilva@yahoo.com.br
2
Universidade Federal de Lavras, Departamento de Biologia, setor de Ecologia. Campus Universitário. Lavras MG 37200-000, Bra-
zil. E-mail: jlouzada@gmail.com
3
Muséum National d’Histoire Naturelle, Département Systématique et Évolution, Entomologie, 57 rue Cuvier, F-75231 Paris cedex
05, France. Permanent address: Universidade Federal de Mato Grosso, Instituto de Biociências, Departamento de Biologia e Zoolo-
gia. Av. Fernando Correa da Costa, 2367. Boa Esperança. Cuiabá MT 78060-900, Brazil.
E-mail: vazdemello@gmail.com. CNPq fellow.
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452
Resumo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452
Material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 453
Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 454
Key to adults of species of Deltochilum subgenus Aganhyboma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455
Chave para adultos das espécies de Deltochilum do subgênero Aganhyboma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456
Subgenus AGANHYBOMA Kolbe 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 468
trisignatum group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 470
trisignatum complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471
Deltochilum (Aganhyboma) trisignatum Harold 1881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471
Deltochilum (Aganhyboma) kolbei Paulian 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476
Deltochilum (Aganhyboma) violaceum Paulian 1938. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
Deltochilum (Aganhyboma) amandaarcanjoae new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 478
cupreicolle complex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479
Deltochilum (Aganhyboma) cupreicolle (Blanchard 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479
Deltochilum (Aganhyboma) titovidaurrei new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .480
Deltochilum (Aganhyboma) viridescens Martínez 1948 new status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .481
Deltochilum (Aganhyboma) viridicatum new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 483
valgum group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 484
acropyge complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 485
Deltochilum (Aganhyboma) acropyge Bates 1887 new subgeneric placement and new status . . . . . . . . . . . . . . . . . . . . . . . .485
Deltochilum (Aganhyboma) feeri new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .486
Deltochilum (Aganhyboma) schefflerorum new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .487
Deltochilum (Aganhyboma) streblopodum new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487
Deltochilum (Aganhyboma) longiceps Paulian 1938 new subgeneric placement and new status . . . . . . . . . . . . . . . . . . . . . 488
Deltochilum (Aganhyboma) acanthus Kohlmann & Solís 2012 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .489
Deltochilum (Aganhyboma) arturoi new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 490
Deltochilum (Aganhyboma) larseni new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .491
finestriatum complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 492
Deltochilum (Aganhyboma) cangalha new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .492
Deltochilum (Aganhyboma) alpercata new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493
Deltochilum (Aganhyboma) finestriatum new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 494
valgum complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495
Deltochilum (Aganhyboma) icaroides Balthasar 1939 new subgeneric placement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495
Deltochilum (Aganhyboma) valgum Burmeister 1873 new subgeneric placement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .496
icariforme subcomplex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 497
Deltochilum (Aganhyboma) ritamourae new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 498
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Deltochilum (Aganhyboma) icariforme Paulian 1938 new subgeneric placement. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 498
Deltochilum (Aganhyboma) paresi new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 500
Deltochilum (Aganhyboma) kolleri new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 500
Deltochilum (Aganhyboma) subrubrum new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .502
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .502
Abstract
The species of the Deltochilum subgenus Aganhyboma, endemic to Central and South America, are reviewed. The species
have been divided into complexes, with a brief diagnosis presented for each. A key for identification of 26 currently rec-
ognized species is provided in English and Portuguese. Two major groups within the subgenus are recognized (trisignatum
and valgum). The first is represented by species typically belonging to the subgenus Aganhyboma (Deltochilum (A.) trisig-
natum, D. (A.) kolbei, D. (A.) violaceum, D. (A.) cupreicolle and D. (A.) viridescens new status) and three new species
described here (D. (A.) amandaarcanjoae, D. (A.) viridicatum and D. (A.) titovidaurrei). The second group (valgum) is
represented by: D. (A.) valgum, D. (A.) longiceps new status, D. (A.) acropyge new status, D. (A.) acanthus, D. (A.) ic-
aroides and D. (A.) icariforme), previously assigned to the subgenus Deltohyboma. Twelve new species are described for
the valgum group (D. (A.) schefflerorum; D. (A.) streblopodum; D. (A.) feeri; D. (A.) larseni; D. (A.) arturoi; D. (A.) fin-
estriatum; D. (A.) cangalha; D. (A.) alpercata; D. (A.) ritamourae; D. (A.) kolleri; D. (A.) paresi; D. (A.) subrubrum). A
lectotype is designated for D. (A.) trisignatum and D. (A.) icariforme. A detailed literature review, synonymies, descrip-
tion, illustration of key morphological characters, data of the studied material and geographic distribution is provided for
each species.
Key words: Systematics, dung beetles, millipede predators, Deltochilum, Aganhyboma
Resumo
As espécies de Deltochilum do subgênero Aganhyboma, endêmico das Américas do Sul e Central, são revisadas. As es-
pécies foram aqui organizadas em complexos, com uma breve diagnose apresentada. Uma chave para identificação de 26
espécies reconhecidas é apresentada em inglês e português. Dois grandes grupos dentro do gênero são reconhecidos
(trisignatum e valgum). O primeiro representado pelas espécies pertencentes tipicamente ao subgênero Aganhyboma (Del-
tochilum (A.) trisignatum, D. (A.) kolbei, D. (A.) violaceum, D. (A.) cupreicolle e D. (A.) viridescens novo status) e três
novas espécies descritas aqui (D. (A.) amandaarcanjoae, D. (A.) viridicatum e D. (A.) titovidaurrei). O segundo grupo
(valgum) é representado pelas espécies: D. (A.) valgum, D. (A.) longiceps novo status, D. (A.) acropyge novo status, D.
(A.) acanthus, D. (A.) icaroides e D. (A.) icariforme), pertencentes anteriormente ao subgênero Deltohyboma. Doze novas
espécies são descritas para o grupo valgum (D. (A.) schefflerorum; D. (A.) streblopodum; D. (A.) feeri; D. (A.) larseni; D.
(A.) arturoi; D. (A.) finestriatum; D. (A.) cangalha; D. (A.) alpercata; D. (A.) ritamourae; D. (A.) kolleri; D. (A.) paresi;
D. (A.) subrubrum). Um lectótipo é designado para D. (A.) trisignatum e D. (A.) icariforme. Uma revisão de literatura
detalhada, sinonímias, descrição, ilustração de importantes caracteres morfológicos, dados do material estudado e dis-
tribuição geográfica é apresentada para cada espécie.
Palavras-chave: Sistemática, rola-bosta, predadores de milípedes
, Deltochilum, Aganhyboma
Introduction
The subgenus Aganhyboma was proposed by Kolbe in 1893 for species of Deltochilum (D. trisignatum Harold, D.
cupricolle Blanchard, D. atrocoeruleum Kolbe [invalid name], D. signiferum Kolbe [invalid name]). However, he did
not designate a type species. The characteristics on which Kolbe defined this subgenus were: medium body size;
complete lateral elytral carina, extending the entire length of the elytra and separated from the epipleural carina;
absence of anterior tarsi.
Shipp (1897) treated Aganhyboma as a genus and designated D. trisignatum as the type species. In the most
recent revision of Aganhyboma by Paulian (1938), two new species (D. kolbei Paulian and D. violaceum Paulian)
Zootaxa 3925 (4) © 2015 Magnolia Press
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A REVISI ON OF AGANHYBOMA KOLBE, 1893
were described and a key to the species was provided. Paulian (1938) treated Aganhyboma as a subgenus of
Deltochilum. In the same paper Paulian erroneously designated D. cupreicolle Blanchard as the type species of the
subgenus, an action nullified by the earlier designation by Shipp (1897). Since Paulian (1938), only Martínez (1948)
has contributed to this subgenus with the description of a new subspecies, D. cupreicolle viridescens Martínez.
Paulian (1938) placed D. valgum Burmeister and D. acropyge Bates in the subgenus Deltochilum. In this same
work, Paulian described a variety, D. valgum var. longiceps, and considered valgum to consist of three subspecies (var.
valgum, var. acropyge, var. longiceps). D. icariforme Paulian 1938 was described and placed close to valgum. The
valgum group was established for this set of species. Balthasar (1939) described a new species (D. icaroides Balthasar
1939) which he placed in the valgum group.
Lane (1946) pointed out Paulian’s (1938) error in designating D. submetallicum (Castelnau 1840) as the type of
the subgenus Deltochilum, a monobasic genus including only the de facto type, D. dentipes Eschscholtz 1822. In the
same work, Lane established the subgenus Deltohyboma and designated D. submetallicum as type and in which he
placed all species of the valgum group. Kohlmann and Solís (2012) described a new species in this group, D.
acanthus.
Paulian (1938) recognized the morphological proximity of the valgum group and the subgenus Aganhyboma
(shape of the head and pygidium, and the flattening of the elytral disc, as well as the simple parameres shape) but did
not combine the two.
The purpose of this paper is to review the taxonomy of the subgenus Aganhyboma, including the species
comprising the valgum group formerly placed in the subgenus Deltohyboma.
Material
This study was based on the examination of 306 adult specimens of Aganhyboma and the valgum group. Material
was kindly provided by the curators of several institutions and private collections as listed below.
AFIC Adrian Forsyth Insect Collection, housed at National Museum of Natural History, Smithsonian
Institution, Washington, USA (Trond Larsen).
BMNH Natural History Museum, London, England (Max Barclay and Malcolm Kerley)
CEMT Seção de Entomologia da Coleção Zoológica, Departamento de Biologia e Zoologia,
Universidade Federal de Mato Grosso, Cuiabá, Brasil (Fernando Vaz-de-Mello).
CEUFPB Coleção do Laboratório de Entomologia, Universidade Federal da Paraíba, João Pessoa, Brasil
(Adelmar Gomes Bandeira).
CMN Canadian Museum of Nature, Ottawa, Canadá (François Génier).
CREN Coleção de Referência de Escarabeíneos Neotropicais, Departamento de Biologia, Setor de
Ecologia, Universidade Federal de Lavras, Lavras, Brasil (Júlio Louzada).
DZUP Coleção Entomológica Padre Jesus Santiago Moure, Departamento de Zoologia, Universidade
Federal do Paraná, Curitiba, Brasil (Lúcia Massutti de Almeida).
INBio collection National Biodiversity Institute, Santo Domingo de Heredia, Costa Rica.
INPA Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil (Márcio Luiz de Oliveira and
Thiago Mahlmann).
MACN Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina (Axel Bachmann).
MNHN Muséum National d'Histoire Naturelle, Paris, França (Olivier Montreuil e Antoine Mantilleri)
MNKM Museo Noel Kempff Mercado, Universidad Autonoma Gabriel Rene Moreno, Santa Cruz de la
Sierra, Bolívia (Julieta Ledezma).
MZUEFS Museu de Zoologia da Universidade Estadual de Feira de Santana, Feira de Santana, Brasil
(Freddy Bravo).
MZUSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brasil (Carlos Campaner).
NMPC National Museum of Natural History, Prague, Czech Republic (Jiří Hájek)
WDEC W. David Edmonds, private collection, Marfa, Texas.
MNHU Museum für Naturkunde, Humboldt-Universität, Berlin, Germany (Johannes Frisch and Joachim
Willers).
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Methods
Examination of the aedeagus, endophallic sclerites and genital segment was undertaken to clarify to the taxonomic
differentiation of species. In describing these structures here, we generally follow Tarasov and Solodovnikov
(2011), with some slight modifications. The terminology used for these sclerites is presented in Figures 1–3. The
endophallus was removed from the aedeagus through the basal foramen of the phallobase, and its sclerites were
illustrated according to the orientation inside the body (the symetric aedeagus rests on its left side, with its right
lateral to up); endophallic sclerites were drawn in dorsal view in the resting position (seen through the right lateral
of the aedeagus) as shown in Figure 1, except for the superior right peripheral sclerite (SRP), which was illustrated
in ventral view in resting position (seen from the left side of the aedeagus). The sclerites were found to be
taxonomically useful: lamella copulatrix (Fig. 1), superior right peripheral sclerite (Figs 1, 3) and fronto-lateral
peripheral sclerite (Fig. 1). Whereas, the complex of subaxial and axial sclerites (Fig. 59) did not provide
significant taxonomic value.
Dissection of genitalia utilized soapy, hot water hydration, followed by immersion in warm 5% KOH solution,
followed by washing in clean water and alcohol and stored in a vial with distilled glycerine.
The illustrations were made using a digitizing tablet and image editing software (Adobe Photoshop CS4).
Some illustrations were drawn with the aid of a camera system coupled to a Leica stereomicroscope M205A.
Images of specimens and key characters were taken with a Leica stereomicroscope M205A, using image stacking
software (Leica Application Suite, version 3.7.0).
FIGURES 1–3. Male genital sclerites in Deltochilum (Aganhyboma). 1. Position and nomenclature of endophallic sclerites
(modified from Medina et al. 2013). 2. Nomenclature of the main parts of the genital segment (D. amandaarcanjoae new
species). 3. Nomenclature of the main parts of the superior right peripheral sclerite (SRP) (D. paresi new species). Figures
scale = 1 mm.
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A REVISI ON OF AGANHYBOMA KOLBE, 1893
Key to adults of species of Deltochilum subgenus Aganhyboma
1. Dorsal interocular distance nine to ten times eye width (Fig. 60); ninth interstria completely longitudinally carinate; elytral lat-
eral edge bounded by the carina of ninth interstria (Fig. 62); length of sixth abdominal ventrite at midline equal to or less than
that of fifth ventrite; metafemur clearly narrowed at base (Fig. 64). (trisignatum group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Dorsal interocular distance three to four times eye width (Fig. 61); carina of ninth elytral interstria at most extending 3/4 length
of interstria; posterior part of elytral lateral edge formed by the pseudepipleural carina (Fig. 63); length of sixth abdominal ven-
trite at midline greater than that of fifth ventrite; metafemur slightly narrowed at base (Fig. 65). (valgum group) . . . . . . . . . .9
2. Apex of pygidium rounded; basal third of the protibia abruptly expanded along inner edge (weakly indicated in D. amandaar-
canjoae) (Fig. 66); medial lateral tooth of protibia closer to apical tooth than to basal tooth; apical one-third of metatibia
strongly curved inward (weakly indicated in D. amandaarcanjoae) (Fig. 67) (trisignatum complex) . . . . . . . . . . . . . . . . . . . .3
- Apex of pygidium acuminate (Fig. 73); protibia not abruptly widened; medial lateral tooth of protibia about equidistant from
basal and apical teeth; apical one-third of metatibia only slightly curved inward (Fig. 68) (cupreicolle complex) . . . . . . . . . .6
3. Body completely bluish-black or violaceous (Fig. 98); margin of pronotum (close to lateral angles) straight, without depres-
sion; punctuation of abdominal sternites concentrated anterolaterally; anterior edge of profemur, in ventral view, with a com-
plete margin. Male genitalia fig. 8. Brazil and Paraguay (Cerrado and Chaco) . . . . . . . . . . . . . .D. amandaarcanjoae (Fig. 98).
- Body not colored as above; margin of pronotum (close to lateral angles) curved inward; punctuation of abdominal sternites
concentrated on anterior margin and posterior margin; anterior margin of profemur effaced at middle portion (Fig. 69) . . . . . 4
4. Head and pronotum green; pronotum lacking distinct spots; basal carina of seventh interstria longer than metatibial spur . . . 5
- Head and pronotum differently coloured; anterior edge of pronotum with three black spots; elytra brown; basal carina of sev-
enth interstria short (Fig. 95), approximately the same length as metatibial spur. Male genitalia figs 9, 27, 35, 44, 52. Brazil
(southern Bahia, Espirito Santo and Minas Gerais) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..D. trisignatum (Fig. 95).
5. Elytra with ocellate punctures (microtubercles) (Fig. 70); elytra surface slightly rugose, with an opaque sheen, except in ocel-
late punctures, which are shiny; apex of second interstria with tubercle. Brazil (Minas Gerais) . . . . . . D. violaceum (Fig. 97).
- Elytra surface smooth, without ocellate punctures and with strong bright metallic color (Fig. 96); apex of second interstria
without tubercle or tubercle weakly indicated. Brazil (Atlantic Forest of northeast region) . . . . . . . . . . . . . .D. kolbei (Fig. 96).
6. Basal carina of seventh interstria elongate, length about four times the width of the seventh interestria (Fig. 71); foveiform
punctures on anterior portion of third interstria small, separated by three times or more their diameter; punctures at center of
the third elytral stria separated by 1.5 times or less their diameter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
- Basal carina of seventh interstria short, length about twice the width of the seventh interestria (Fig. 72); foveiform punctures
on anterior portion of third interstria separated by less than three times their diameter; punctures at center of the third elytral
stria separated by two or more times their diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
7. Head with strong green coloration and metallic sheen; posteromedial portion of pronotum with a semi-circular band which
extends up to half of pronotal disc (Fig. 101), or pronotum completely green (Figs 102, 103); elytra dark green; male with apex
of pygidium strongly acuminate (Fig. 73). Male genitalia figs 6, 34, 43, 51, 59. Brazil and Paraguay (Cerrado and Chaco) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. viridescens (Figs 101, 102, 103).
- Head with weak green or black-violaceous coloration, and an opaque sheen (Figs 104, 105); posteromedial portion of prono-
tum lacking greenish band or only faintly indicated; elytra with dark green or black-violaceous coloration; male with apex of
pygidium weakly acuminate or rounded. Male genitalia figs 7, 34. Argentina, southern Brazil, Uruguay [doubtful] . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. viridicatum (Figs 104, 105).
8. Elytra dark brown or black (Fig. 100); disc of pronotum with transverse oval spot (Fig. 100); aedeagus with short parameres
and phallobase (Fig. 4). Male genitalia figs 4, 42. Brazil (Goiás, Minas Gerais). . . . . . . . . . . . . . . . D. titovidaurrei (Fig. 100).
- Elytra reddish or light brown (Fig. 99); disc of pronotum lacking transverse oval spot. When pronotal disc presents this spot,
elytra never dark brown or black; aedeagus with parameres and phallobase more elongate (Fig. 5). Male genitalia figs 5, 25,
33, 50. Bolivia and Brazil (Cerrado and Chaco). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cupreicolle (Fig. 99).
9. Elytral striae narrow, with carinate margins close and almost touching (Fig. 74) (finestriatum complex) . . . . . . . . . . . . . . . .10
- Elytral striae wider, with carinate margins clearly separated (Fig. 75) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
10. Microtubercles of discal interstriae weakly indicated and united by depressed glossy areas making their delimitation difficult.
Elytra with dark brown sheen (Fig. 74); elytra oval-elongate with lateral edges slightly convex (Fig. 116). Male genitalia figs
11, 28, 36, 45, 53. Brazil (Atlantic Forest of south and southeast regions) . . . . . . . . . . . . . . . . . . . ..D. finestriatum (Fig. 116).
- Microtubercles of discal interstriae prominent, united or not by glossy spots. Elytral surface with or without red metallic sheen;
elytra rounded, sides distinctly convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Punctures of anteromedial portion of hypomera separated by at least one diameter (Fig. 78); pronotum with red metallic sheen
(Fig. 114); elytral microtubercles weakly delimited, with red-cupreous sheen (Fig. 76). Brazil (Minas Gerais) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cangalha (Fig. 114).
- Punctures of anteromedial portion of hypomera separated by less than one diameter (Fig. 79); pronotum black with an opaque
sheen or weak luster (Fig. 115); elytral microtubercles well delimited, with brown sheen (Fig. 77). Brazil (Alagoas) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..D. alpercata (Fig. 115).
12. Basal carina of ninth interstriae extending at most to midway along elytra (Fig. 80). If basal carina of ninth interstriae extend-
ing 3/4 along elytral length, pseudepipleural carina incomplete, effaced medially (Fig. 80) (acropyge complex) . . . . . . . . . . 13
- Basal carina of ninth interstria extending 3/4 or more along elytral length; pseudepipleural carina complete, visible entire
length of epipleuron (valgum complex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
13. Pseudepipleural carina complete, visible entire length of epipleuron; male genitalia figs 13, 46. Mexico to Panama . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. acropyge (Fig. 106).
SILVA ET AL.
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Zootaxa 3925 (4) © 2015 Magnolia Press
- Pseudepipleural carina incomplete, effaced medially (Fig. 80) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
14. Hypomera with foveiform punctures dense on anterior and posterior portions (Fig. 81) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
- Hypomera with dense foveiform punctures on anterior portion only (punctures not touching), and absent or dispersed punc-
tures in a diagonal band (Figs 82–83, indicated by arrow) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
15. Foveiform punctures of head almost touching, separated in dorsal interocular surface by about half diameter. Guyana, French
Guiana and Trinidad and Tobago . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. feeri (Fig. 107).
- Foveiform punctures of head touching. Brazil (South of the Amazon Basin in Mato Grosso and Pará) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. schefflerorum (Fig. 108).
16. Diagonal band of hypomera lacking punctures or with one or two isolated punctures (Fig. 83) . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Diagonal band of hypomera with dispersed but obvious punctures (Fig. 82). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
17. Punctures of elytral striae often elliptical, separated by irregular spacing. Some punctures close, while others are clearly dis-
persed (Fig. 84); length > 13 mm. Male genitalia figs 16, 37. Colombia . . . . . . . . . . . . . . . . . . . . . . . .D. longiceps (Fig. 110).
- Punctures of elytral striae round, or at most only weakly elliptical and almost always regularly spaced; length < 13 mm . . .18
18. Length of edge of pronotum between lateral and posterior angles less than that between anterior and lateral angles; elytral
striae with round punctures; apex of second interstria without carina. Brazil (Amazonas) . . . . . . . .D. streblopodum (Fig. 109).
- Length of anterior and posterior portions of lateral pronotal edge approximately equal; elytral striae with weakly elliptical
punctures; apex of second interstria with carina. Male genitalia fig. 15. Panama and Costa Rica . . . . . D. acanthus (Fig. 111).
19. Foveiform punctures of head separated in dorsal interocular surface by approximately half diameter; dorsal interocular dis-
tance approximately three times eye width; foveiform punctures of interstria surrounded by approximately four well-defined
shiny microtubercles; apex of second interstria with inconspicuous carina or tubercle; length > 13 mm. Ecuador . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. arturoi (Fig. 112).
- Foveiform punctures of head nearly touching; dorsal interocular distance approximately 2.5 times eye width; foveiform punc-
tures of interstriae surrounded by approximately five well-defined shiny microtubercles; apex of second interstria without
carina or tubercle; length < 13 mm. Male genitalia fig. 14. Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. larseni (Fig. 113).
20. Third interstria with foveiform punctures covering entire interstrial surface; elytral surface with opaque or silky sheen; black
coloration. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
- Foveiform punctures of third interstria denser laterally (punctures can be so scattered at center of interstria to produce smooth
midlongitudinal strip) (Fig. 86); elytra metallic; copper-red coloration (icariforme subcomplex) . . . . . . . . . . . . . . . . . . . . . .22
21. Apical carina of third interstria only somewhat more prominent than apical carinae of second and fourth interstriae (Fig. 87);
foveiform punctures of posteromedial portion of pronotum usually separated by less than one diameter; interstrial microtuber-
cles usually inconspicuous, sometimes blending in with remainder of interstria surface. Male genitalia figs 18, 31, 39, 48.
Argentina, Bolivia, Brazil and Paraguay. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. icaroides (Fig. 117).
- Apical carina of third interstria distinctly more prominent and round than apical carinae of second and fourth interstriae (Fig.
88); foveiform punctures of posteromedial portion of pronotum usually separated by at least one diameter; interstrial microtu-
bercles dense and prominent. Male genitalia figs 17, 30, 38, 47, 55. Argentina, Bolivia and Paraguay . . .D. valgum (Fig. 118).
22. Apical carina of seventh interstria elongate, length greater than the sum of lengths of apical carinae of fifth and sixth inter-
striae; basal carina of seventh interstria clearly shorter than apical carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23
- Apical carina of seventh interstria with length less than or equal to the sum of lengths of apical carinae of fifth and sixth inter-
striae; basal carina of seventh interstria equal to or longer than apical carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
23. Basal carina of ninth interstria surpassing anterior limit of apical carina of seventh interstria; elytral lateral edges distinctly
curved outward (Fig. 121). Male genitalia fig. 20. Brazil (Mato Grosso) . . . . . . . . . . . . . . . . . . . . . . . . . . D. paresi (Fig. 121).
- Basal carina of ninth interstria not surpassing anterior limit of apical carina of seventh interstria; elytral lateral edges slightly
curved outward (Fig. 120). Male genitalia figs 21, 32, 40, 57. Brazil (Goiás, Mato Grosso and São Paulo) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. icariforme (Fig. 120).
24. Lateral edges of elytra subparallel or slightly curved outward (Figs 122, 123). If curved outward, body never completely cop-
per with reddish metallic sheen; dark brown or black coloration, with greenish or slightly reddish metallic sheen . . . . . . . .25
- Elytral lateral edge distinctly curved outward; entirely copper and reddish metallic sheen. Male genitalia figs 22, 58. Brazil
(Bahia, Ceará, Minas Gerais and Piauí) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. ritamourae (Fig. 119).
25. Microtubercles of elytral interstriae absent or inconspicuous (Fig. 86); foveiform punctures near the center of posterior edge of
pronotum separated by at least one diameter. Male genitalia figs 23, 41. Argentina (Misiones). . . . . .D. subrubrum (Fig. 123).
- Elytral microtubercles more conspicuous on lateral interstriae; foveiform punctures near the center of posterior edge of prono-
tum separated by less than one diameter. Male genitalia figs 19, 49, 56. Brazil (Mato Grosso do Sul and São Paulo) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. kolleri (Fig. 122).
Chave para adultos das espécies de Deltochilum do subgênero Aganhyboma
1. Distância interocular dorsal nove a dez vezes a largura do olho (Fig. 60); nona interestria inteiramente carenada longitudinal-
mente; bordo lateral do élitro inteiramente formado pela carena da nona interestria (Fig. 62); sexto ventrito abdominal com
comprimento na linha do meio igual ou menor que aquele do quinto ventrito; metafêmur nitidamente mais estreito na base
(Fig. 64). (grupo trisignatum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Distância interocular dorsal três a quatro vezes a largura do olho (Fig. 61); carena da nona interestria elitral atinge no máximo
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A REVISI ON OF AGANHYBOMA KOLBE, 1893
3/4 do comprimento da interestria; parte posterior do bordo lateral do élitro formada pela carena pseudoepipleural (Fig. 63);
meio do sexto ventrito abdominal com comprimento maior que aquele do quinto ventrito; metafêmur apenas um pouco mais
estreito na base (Fig. 65). (grupo valgum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
2. Ápice do pigídio arredondado; terço basal da protíbia bruscamente alargado no bordo interno (fracamente indicado em D.
amandaarcanjoae) (Fig. 66); dente medial lateral da protíbia mais próximo do dente apical que do dente basal; metatíbia forte-
mente arqueada em direção ao corpo na sua terça parte apical (fracamente indicado em D. amandaarcanjoae) (Fig. 67) (com-
plexo trisignatum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
- Ápice do pigídio ponteagudo (Fig. 73); protíbia com largura subigual em toda sua extensão; dente medial lateral da protíbia
aproximadamente equidistante aos dentes apical e basal; metatíbia levemente arqueada em direção ao corpo (Fig. 68) (com-
plexo cupreicolle) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
3. Corpo inteiramente preto-azulado ou violáceo (Fig. 98); margem do pronoto (próximo aos ângulos laterais) aproximadamente
reta, sem reentrância; pontuação dos ventritos abdominais concentrada anterolateralmente; bordo anterior do profêmur, em
vista ventral, com emarginação completa. Genitália masculina fig. 8. Brasil e Paraguai (Cerrado e Chaco) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. amandaarcanjoae (Fig. 98).
- Corpo com coloração diferente; margem do pronoto (próximo aos ângulos laterais) com reentrância bem marcada; pontuação
dos ventritos abdominais concentrada nos bordos anterior e posterior da lateral dos ventritos; bordo anterior dos profêmures
com emarginação incompleta aproximadamente na metade do bordo (Fig. 69). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4. Cabeça e pronoto verde; pronoto unicolor, sem machas diferenciadas; comprimento da carena basal da sétima interestria maior
que aquele do cálcar da metatíbia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Coloração da cabeça diferente do pronoto; bordo anterior do pronoto com três distintas manchas pretas; élitro castanho; carena
basal da sétima interestria pequena (Fig. 95), com aproximadamente o mesmo comprimento do cálcar da metatíbia. Genitália
masculina figs 9, 27, 35, 44, 52. Brasil (sul da Bahia, Espírito Santo e Minas Gerais) . . . . . . . . . . . . . D. trisignatum (Fig. 95).
5. Élitros com microtubérculos ocelares (Fig. 70); superfície elitral levemente rugosa, com reflexo opaco e brilho apenas nos
microtubérculos da microescultura; tubérculo do ápice da segunda interestria presente. Brasil (Minas Gerais) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. violaceum (Fig. 97).
- Superfície elitral lisa, sem microtubérculos ocelares visíveis e com forte brilho metálico (Fig. 96); tubérculo do ápice da
segunda interestria ausente ou quase inconspícuo. Brasil (Mata Atlântica da região nordeste) . . . . . . . . . ..D. kolbei (Fig. 96).
6. Carena da base da sétima interestria alongada, com aproximadamente quatro vezes a largura da sétima interestria (Fig. 71);
pontos foveiformes da região anterior da terceira interestria pequenos, separados por três ou mais vezes o seu diâmetro; pontos
no centro da terceira estria elitral distanciados por 1,5 vezes ou menos o seu diâmetro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
- Carena da base da sétima interestria pequena, com aproximadamente duas vezes a largura da sétima interestria (Fig. 72); pon-
tos foveiformes da região anterior da terceira interestria separados por menos que três vezes o seu diâmetro; pontos no centro
da terceira estria elitral distanciados por duas ou mais vezes o seu diâmetro. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
7. Cabeça com coloração verde intensa e reflexos metálicos; região posterior do pronoto com forte mancha esverdeada semi-cir-
cular (Fig. 101), ou pronoto completamente verde (Figs 102, 103); élitros verde escuro; macho com o ápice do pigídio forte-
mente ponteagudo (Fig. 73). Genitália masculina figs 6, 34, 43, 51, 59. Brasil e Paraguai (Cerrado e Chaco) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. viridescens (Figs 101, 102, 103).
- Cabeça com coloração verde claro, ou preto-violáceo, sem brilho metálico (Figs 104, 105); região posterior do pronoto sem
mancha esverdeada semi-circular, ou com apenas uma indicação da mesma; élitros verde escuro, ou preto-violáceo; macho
com o ápice do pigídio suavemente acuminado ou arredondado. Genitália masculina figs 7, 34. Argentina, sul do Brasil, Uru-
guai [registro duvidoso] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. viridicatum (Figs 104, 105).
8. Élitros marrom escuro ou preto (Fig. 100); disco do pronoto com mancha oval transversa, (Fig. 100); edeago com parâmeros e
falobase curtos (Fig. 4). Genitália masculina figs 4, 42. Brasil (Goiás, Minas Gerais) . . . . . . . . . . . D. titovidaurrei (Fig. 100).
- Élitros avermelhados ou marrom claro (Fig. 99); disco do pronoto sem mancha oval transversa. Quando o disco do pronoto
apresenta essa mancha, élitro nunca marrom escuro ou preto; edeago com parâmeros e falobase mais alongados (Fig. 5). Geni-
tália masculina figs 5, 25, 33, 50. Bolívia e Brasil (Cerrado e Chaco) . . . . . . . . . . . . . . . . . . . . . . . . . . D. cupreicolle (Fig. 99).
9. Estrias elitrais delgadas, com margens carenadas que delimitam a estria próximas, quase tocando-se (Fig. 74) (complexo fines-
triatum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Estrias elitrais alargadas, com margens carenadas claramente separadas (Fig. 75) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
10. Microtubérculos das interestrias discais pouco destacados e unidos por áreas brilhantes de baixo relevo (Fig. 74). Isso torna
difícil a delimitação dos microtubérculos. Élitros com reflexo marrom escuro (Fig. 74); com formato oval alongado e bordos
laterais suavemente curvados para fora (Fig. 116). Genitália masculina figs 11, 28, 36, 45, 53. Brasil (Mata Atlântica da região
Sul e Sudeste) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..D. finestriatum (Fig. 116).
- Microtubérculos das interestrias discais proeminentes, unidos ou não por manchas brilhantes; élitros arredondados e bordos
laterais nitidamente curvados para fora. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Pontos da região antero-mediana do hipômero separados por no mínimo um diâmetro (Fig. 78); pronoto com reflexo vermelho
metálico (Fig. 114); microtubérculos elitrais pouco delimitados, com reflexo vermelho cúpreo (Fig. 76). Brasil (Minas Gerais)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. cangalha (Fig. 114).
- Pontos da região antero-mediana do hipômero separados por menos que um diâmetro (Fig. 79); pronoto preto, com reflexo
opaco ou fraco brilho (Fig. 115); microtubérculos elitrais bem delimitados, com reflexo marrom (Fig. 77). Brasil (Alagoas) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..D. alpercata (Fig. 115).
12. Carena da base da nona interestria geralmente não ultrapassa a metade do comprimento do élitro (Fig. 80). Se a carena da base
da nona interestria ultrapassar a metade do comprimento do élitro, carena pseudoepipleural incompleta, porção do meio apa-
gada (Fig. 80) (complexo acropyge) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
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- Carena da base da nona interestria se extende por 3/4 ou mais do comprimento do élitro; Carena pseudoepipleural completa,
visível em toda a extensão da epipleura (complexo valgum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
13. Carena pseudoepipleural completa, visível em toda a extensão da epipleura; genitália masculina figs 13, 46. México ao Pana-
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. acropyge (Fig. 106).
- Carena pseudoepipleural incompleta, porção do meio apagada (Fig. 80) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14. Hipômero com pontuação foveiforme densa nas partes anterior e posterior (Fig. 81). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- Hipômero com pontuação foveiforme densa apenas na parte anterior (pontos não se tocam), e ausente ou dispersa em uma
faixa diagonal (Figs 82-83, indicada por seta) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
15. Pontos foveiformes da cabeça quase tocando-se, separados na superfície interocular dorsal por aproximadamente metade do
diâmetro de um ponto. Guiana, Guiana Francesa e Trinidad e Tobago . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. feeri (Fig. 107).
- Pontos foveiformes da cabeça tocando-se. Brasil (sudeste do Rio Amazonas, no Mato Grosso e Pará) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..D. schefflerorum (Fig. 108).
16. Faixa diagonal do hipômero sem pontuação ou com um ou dois pontos isolados (Fig. 83) . . . . . . . . . . . . . . . . . . . . . . . . . . .17
- Faixa diagonal do hipômero com pontuação dispersa porém evidente (Fig. 82) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
17. Pontos das estrias elitrais frequentemente distintamente alongados (com formato elíptico), separados por espaçamento irregu-
lar. Alguns desses pontos são bem próximos enquanto outros são nitidamente dispersos (Fig. 84); comprimento > 13 mm. Gen-
itália masculina figs 16, 37. Colômbia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. longiceps (Fig. 110).
- Pontos das estrias elitrais arredondados, ou no máximo um pouco elípticos e sempre separados por espaçamento aproximada-
mente regular; comprimento < 13 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
18. Comprimento do bordo do pronoto, entre os ângulos lateral e posterior, menor que o comprimento do bordo entre os ângulos
anterior e lateral; estrias elitrais com pontos arredondados; ápice da segunda interestria sem carena. Brasil (Amazonas) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. streblopodum (Fig. 109).
- Comprimento das porções anterior e posterior do bordo lateral do pronoto aproximadamente iguais; estrias elitrais com pontos
fracamente elípticos; ápice da segunda interestria com carena. Genitália masculina fig. 15. Panamá e Costa Rica . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. acanthus (Fig. 111).
19. Pontos foveiformes da cabeça separados na superfície interocular dorsal por aproximadamente metade dos seus diâmetros; dis-
tância interocular dorsal com aproximadamente três vezes a largura do olho; pontos foveiformes das interestrias circundados
por aproximadamente quatro microtubérculos brilhantes bem definidos; ápice da segunda interestria com carena ou tubérculo
inconspícuo; comprimento > 13 mm. Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. arturoi (Fig. 112).
- Pontos foveiformes da cabeça praticamente tocam-se; distância interocular dorsal com aproximadamente 2,5 vezes a largura
do olho; pontos foveiformes das interestrias circundados por aproximadamente cinco microtubérculos brilhantes bem defini-
dos; ápice da segunda interestria sem carena ou tubérculo; comprimento < 13 mm. Genitália masculina fig. 14. Peru . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. larseni (Fig. 113).
20. Terceira interestria com pontos foveiformes cobrindo toda a superfície; superfície do élitro com reflexo opaco ou sedoso;
espécimes com coloração negra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- Pontos foveiformes da terceira interestria mais densos lateralmente (os pontos podem estar tão dispersos no centro da interes-
tria que produzem um aspecto liso em uma faixa longitudinal) (Fig. 86); élitros com brilho metálico; coloração vermelho
cúpreo (subcomplexo icariforme). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
21. Carena apical da terceira interestria apenas um pouco mais proeminente que as carenas do ápice da segunda e quarta interestria
(Fig. 87); pontos foveiformes próximos ao centro da região posterior do pronoto geralmente separados por menos que um
diâmetro; microtubérculos das interestrias geralmente inconspícuos, algumas vezes se misturam com a superfície ao redor.
Genitália masculina figs 18, 31, 39, 48. Argentina, Bolívia, Brasil e Paraguai . . . . . . . . . . . . . . . . . . . . D. icaroides (Fig. 117).
- Carena apical da terceira interestria distintamente mais proeminente e arredondada que as carenas do ápice da segunda e quarta
interestria (Fig. 88); pontos foveiformes próximos ao centro da região posterior do pronoto geralmente separados por no míni-
mo um diâmetro; microtubérculos das interestrias densos e conspícuos. Genitália masculina figs 17, 30, 38, 47, 55. Argentina,
Bolívia e Paraguai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. valgum (Fig. 118).
22. Carena apical da sétima interestria alongada, comprimento maior que a soma dos comprimentos das carenas do ápice da quinta
e sexta interestrias; carena da base da sétima interestria nitidamente menor que a carena apical . . . . . . . . . . . . . . . . . . . . . . . 23
- Carena apical da sétima interestria com comprimento menor ou igual a soma dos comprimentos das carenas do ápice da quinta
e sexta interestrias; comprimento da carena da base da sétima interestria mais longo ou aproximadamente igual ao da carena
apical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
23. Carena da base da nona interestria ultrapassa o limite anterior da carena apical da sétima interestria; bordo lateral do élitro niti-
damente curvado para fora (Fig. 121). Genitália masculina fig. 20. Brasil (Mato Grosso) . . . . . . . . . . . . . .D. paresi (Fig. 121).
- Carena da base da nona interestria não ultrapassa o limite anterior da carena do ápice da sétima interestria; bordo lateral do
élitro suavemente curvado para fora (Fig. 29). Genitália masculina figs 21, 32, 40, 57. Brasil (Goiás, Mato Grosso e São Paulo)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. icariforme (Fig. 120).
24. Bordos laterais dos élitros subparalelos ou levemente curvados para fora (Figs 122, 123). Se curvados, corpo nunca inteira-
mente cobre com reflexo metálico avermelhado; corpo marrom escuro, ou preto, com reflexo metálico esverdeado ou leve-
mente avermelhado . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
- Bordos laterais dos élitros distintamente curvados para fora; corpo inteiramente cobre e reflexo metálico avermelhado. Geni-
tália masculina figs 22, 58. Brasil (Bahia, Ceará, Minas Gerais e Piauí) . . . . . . . . . . . . . . . . . . . . . . D. ritamourae (Fig. 119).
25. Microtubérculos das interestrias elitrais ausentes ou inconspícuos (Fig. 86); pontos foveiformes próximos ao centro do bordo
posterior do pronoto separados por no mínimo o diâmetro de um ponto. Genitália masculina figs 23, 41. Argentina (Misiones)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D. subrubrum (Fig. 123).
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- Microtubérculos elitrais mais conspícuos nas interestrias laterais; pontos foveiformes próximos ao centro do bordo posterior
do pronoto bastante densos, separados por menos que o diâmetro de um ponto. Genitália masculina figs 19, 49, 56. Brasil
(Mato Grosso do Sul e São Paulo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. kolleri (Fig. 122).
Subgenus AGANHYBOMA Kolbe 1893
Deltochilum (Aganhyboma) Kolbe 1893: 192 (original description)
Shipp 1897: 195 (designation of type species; author treated Aganhyboma as genus)
Paulian 1938: 240, 243, 252 (identification keys, species descriptions)
Lane 1946: 172, 173, 174 (comment about history of names in Aganhyboma)
Pereira & Martínez 1956: 120, 121 (identification key)
Martínez 1959: 51 (catalogue)
Vulcano & Pereira 1964: 641 (catalogue)
Vulcano & Pereira 1967: 555, 556 (identification keys)
Vaz-de-Mello 2000: 192 (checklist)
Vaz-de-Mello et al. 2011: 5, 11, 18, 26, 33, 41, 44 (checklist and identification keys).
Deltochilum (Deltochilum) in part Eschscholtz 1822: 39 (original description)
Kolbe 1893: 191 (establishement of seven subgenera under Deltochilum)
Shipp 1897: 196 (taxonomic comments)
Paulian 1938: 240, 243, 268, 269 (description, identification key and comments)
Lane 1946: 172–175 (transfer of the valgum group to Deltohyboma).
Deltochilum (Deltohyboma) in part Lane 1946: 175 (designation of subgenus)
Martínez 1959: 53–56 (catalogue)
Pereira & Martínez 1956: 120, 121 (identification key)
Vulcano & Pereira 1964: 641 (catalogue)
Vulcano & Pereira 1967: 555–560 (identification keys)
Vaz-de-Mello 2000: 192 (checklist)
Vaz-de-Mello et al. 2011: 5, 11, 18, 26, 33, 41, 44 (checklist and identification keys)
Description. Body. Lateral margins as well as dorsal surface vary from weakly to strongly convex (Figs 95–123),
with some species quite round in shape. Dorsal surface glabrous, with at most minute setae in some species. Length
11 to 19 mm. Head. Surface with foveiform punctures, usually more conspicuous around eyes (Figs 60, 61).
Anterior portion (between clypeogenal sutures) prolonged, similar to a small ‘muzzle’ (Figs 60, 61), bidentate.
Margin of head, near clypeal teeth, straight and followed by a small tooth slightly pronounced (Fig. 61), or slightly
curved and without teeth (Fig. 60). Lateral margin of gena curved outward until eyes, with a strong convex shape in
the middle of this curvature, or with posterior half truncated, approximately straight. Clypeogenal suture well
defined and shallow. Eye margin not emarginate. Eye rounded or comma-shaped in dorsal view. Dorsal interocular
distance varies between three to ten times eye width (Figs 60, 61). Thorax. Pronotum with anterior angles acute,
directed anteriorly. Lateral margin slightly angled. Pronotum punctures variable, dense or sparse, consisting of
foveiform punctures. Sides of metasternum, anterior portion of hypomera, epimeron and episternum with a dense
foveiform punctures. Hypomera not excavated to receive head margin (Figs 78, 79). Posterior portion of
metasternum with a small, round, shallow fovea. Outer margin of mesocoxal cavity not reaching metaepimeron.
Elytra. Lateral margins ranging from semiparallel to outwardly curved. Elytral striae with foveiform punctures.
Interstriae with foveiform punctures of different sizes. Basal carina of sixth elytral interstria absent. Basal carina of
seventh interstria short or elongate, prominent or not (Figs 71, 72). Length of basal carina of ninth interstria reaches
from mid-interstria, to apex (Figs 62, 63, 80). Apices of interstriae 2, 3, 4, 5, 6 and 7 with tubercle or carina (that of
second interstria sometimes weak or absent). Length of apical carina of seventh interstria less than one-half that of
interstria. Pseudepipleural carina complete, visible entire length of epipleuron in most species (Figs 62, 63). In
some species this carina can be interrupted in the basal third, re-appearing halfway along elytral length (Fig. 80).
Abdomen. Abdominal ventrites shorter medially than laterally (Fig. 73). Punctures of ventrites most abundant and
conspicuous at sides. Pygidium with rounded or elliptical foveiform punctures. Apex of pygidium acuminate or
rounded. Legs. Protibia abruptly expanded (Fig. 66) or not in basal third. External apical margin of protibia bearing
three large prominent teeth. Medial lateral tooth can be equidistant between apical and basal tooth, or closer to
apical tooth. Metatibia short, truncated and only slightly curved inward (Fig. 68), or elongate and apical one-third
strongly curved inward (Fig. 67). Aedeagus. Parameres symmetric, short or elongate, simple, without
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ornamentation or setae (Fig. 24); endophallus: lobe in submedial portion; endophallic sclerites: lamella copulatrix
sclerites (LC), superior right peripheral sclerite (SRP), fronto-lateral peripheral sclerite (FLP) and complex of axial
and subaxial sclerites (A + SA) (Figs 1, 59).
Diagnosis. Anterior portion of head (between clypeogenal sutures) prolonged, similar to a small ‘muzzle’
(Figs 60, 61).
Remarks. The subgenus Aganhyboma, as defined here, includes 26 species distributed in South America, with
one species (D. acropyge Bates) reaching as far North as Mexico. The highest species richness occurs in Brazil.
This work recognizes two major species groups within the subgenus, the trisignatum and valgum groups. The first
comprises species historically grouped in Aganhyboma, including the genus type. The trisignatum group can be
further split into two species complexes, the trisignatum and the cupreicolle complexes. The valgum group is
comprised of three species complexes, the acropyge, finestriatum and the valgum complexes.
Genus Deltochilum Eschscholtz 1822
Subgenus Aganhyboma Kolbe 1893
trisignatum group
trisignatum complex
D. (A.) trisignatum Harold 1881
D. (A.) kolbei Paulian 1938
D. (A.) violaceum Paulian 1938
D. (A.) amandaarcanjoae new species
cupreicolle complex
D. (A.) cupreicolle (Blanchard 1841)
D. (A.) titovidaurrei new species
D. (A.) viridescens Martínez 1948 new status
D. (A.) viridicatum new species
valgum group
acropyge complex
D. (A.) acropyge Bates 1887 new subgeneric placement and new status
D. (A.) feeri new species
D. (A.) schefflerorum new species
D. (A.) streblopodum new species
D. (A.) longiceps Paulian 1938 new subgeneric placement and new status
D. (A.) acanthus Kohlmann & Solís 2012 new subgeneric placement
D. (A.) arturoi new species
D. (A.) larseni new species
finestriatum complex
D. (A.) cangalha new species
D. (A.) alpercata new species
D. (A.) finestriatum new species
valgum complex
D. (A.) icaroides Balthasar 1939 new subgeneric placement
D. (A.) valgum Burmeister 1873 new subgeneric placement
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icariforme subcomplex
D. (A.) ritamourae new species
D. (A.) icariforme Paulian 1938 new subgeneric placement
D. (A.) paresi new species
D. (A.) kolleri new species
D. (A.) subrubrum new species
trisignatum group
Description. Body. Small or medium (11–16.5 mm), weakly or strongly convex, with opaque or bright coloration
in shades of black, brown, coppery, green or violet. Head. Elongate. Dorsal interocular distance approximately
nine or ten times eye width (Fig. 60). Thorax. Pronotal margin, between anterior and lateral angles, oblique and
slightly curved outward. Margin between lateral and posterior angles approximately straight or curved outward.
Anterior and posterior portions of hypomera with dense foveiform punctures; anterior punctures rounded, posterior
punctures mostly elliptical. Elytra. With punctate striae. Striae conspicuous or not. Foveiform punctures
distributed evenly along the interstriae. Basal carina of seventh interstria prominent or not. Apices of interstriae 2,
3, 4, 5, 6 and 7 with tubercle or carina (that of second interstria sometimes weak or absent in D. kolbei). Apical
tubercles of 2nd, 3rd and 4th interstriae of approximately equal size and shape. Length of apical carinae of 5th and
6th interstriae approximately equal and slightly longer than apical carinae of 2nd, 3rd and 4th interstriae. Apical
carina of 7th interstria longer than carina of 6th interstria. Ninth interstria completely longitudinally carinate.
Elytral lateral margin completely bounded by the carina of ninth interstria (Fig. 62). Pseudepipleural carina
complete (Fig. 62). First interstria usually brighter than rest of elytra. Abdomen. Punctuation of abdominal
sternites concentrated anterolaterally (Fig. 73). Length of sixth ventrite, measured at midline, equal to or less than
that of fifth ventrite (Fig. 73). Pygidium often acuminate at apex (Fig. 73), or rounded, with large transverse
elliptical punctures. Legs. Most punctures on ventral surface of meso- and metafemora either transverse or oblique
elliptical in shape (Fig. 64). Protibia abruptly expanded (Fig. 66) or not along inner margin. Metafemur clearly
much wider in middle and apex than at base (Fig. 64). Apical one-third of metatibia slightly (Fig. 68) or strongly
(Fig. 67) curved inward, with five or six clearly visible longitudinal carinae. Lateral carina of metatibia with a row
of medium-size setae denser in apical half. Metatarsus much shorter or approximately two-thirds of metatibia
length. Secondary sexual characters. Head of male sometimes slightly more triangular than that of female. Male
abdomen usually slightly concave at lateral view, that of female usually convex. Male ventrites more flattened
antero-posteriorly than that of female. Abdominal suture separating 6th from 5th ventrite usually curved in male
(Fig. 73) and nearly straight in female. Apical margin of pygidium usually distended in male. Protibial spurs
generally vary in shape and size between male and female of some species. Aedeagus. Parameres symmetrical,
short, simple, without ornamentation or setae, and wider at base than at apex (Figs 4–9). Endophallus: with lobe in
submedial portion. Lamella copulatrix: left lobe "J" shaped, right lobe “baton” shaped (Figs 33–35). Superior right
peripheral sclerite (SRP): circular shaped, cable "n" shaped, and ring with margin "thread" shaped (Figs 42–44).
Fronto-lateral peripheral sclerite (FLP): two irregular sclerites, with a sclerotized cover that surrounds it (Figs
50–52). Complex of axial and subaxial sclerites (A+SA): several superposed and elongate sclerites (Fig. 59).
Genital segment: with lateral sclerotized arms in roughly a scalene triangle; central portion approximately "M"
shaped, showing portions with different degrees of sclerotization; superior margin well delimited, with
conspicuous sclerotization (Figs 25, 26, 27).
Diagnosis. Compared to the valgum group, the diagnostic characters of trisignatum group are: Dorsal
interocular distance approximately nine or ten times eye width (Fig. 60); apical tubercle of the 2nd, 3rd and 4th
interstriae with approximately equal size and shape (that of second interstria absent or weak in D. kolbei); length of
apical carinae of 5th and 6th interstriae approximately equal to or slightly longer than apical carinae of 2nd, 3rd and
4th; ninth interstria completely longitudinally carinate (Fig. 62); elytral lateral margin bounded by the carina of
ninth interstria (Fig. 62); ventral surface of meso and metafemora with transverse or oblique elliptical punctures
(Fig. 64); lamella copulatrix (LC): left lobe “J”-shaped and right lobe baton-shaped (Figs 33–35); cable of superior
right peripheral sclerite (SRP) "n"-shaped, and ring with margin “thread”-shaped (Figs 42–44); superior margin of
genital sclerite well-defined, with conspicuous sclerotization (Figs 25–27).
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trisignatum complex
Diagnosis. Apex of pygidium rounded; basal third of protibia sharply widened along its inner margin (Fig. 66);
middle protibial tooth closer to apical tooth than to basal tooth; apical one-third of metatibia strongly curved
inward (Fig. 67).
Deltochilum (Aganhyboma) trisignatum Harold 1881
(Figs 9, 27, 35, 44, 52, 66, 69, 89, 95)
Deltochilum trisignatum Harold 1881 [1880]: 150 (original description)
Heyne 1900: 61 (description)
Gillet 1911: 36 (catalogue)
Pessôa & Lane 1941: 428, 431 and 432 (description and identification key)
Blackwelder 1944: 203 (checklist)
Deltochilum (Aganhyboma) trisignatum Kolbe 1893: 192
Shipp 1897: 195
Paulian 1938: 240, 252, 253, 256, Figs 2, 18, map No 2 (description, taxonomic comments and identification key)
Lane 1946: 173, 174 (taxonomic comments)
Pereira & Martínez 1956: 121
Vulcano & Pereira 1964: 642 (catalogue)
Vaz-de-Mello 2000: 192 (checklist)
Description. Body. Head black with violet hue, pronotum yellowish brown with black spots, elytra reddish brown
(Fig. 95). Ventral surface black with green reflections. Length 17 to 19 mm. Head. Dorsal surface completely
covered by closely spaced to confluent foveiform punctures resulting in rugosity, anterior punctures smaller than
those at center. Punctures margin well-defined (impressed). Margin between clypeal teeth and genae curved
outward near clypeogenal suture. Thorax. Surface of pronotum with deep and dense foveiform punctures forming
rugosity. Anteromedial punctures much smaller than posteromedial punctures. Margin of pronotum (close to lateral
angles), curved inward, lacking emargination. Anterior margin with three black spots, one covering each angle and
one at center. Pronotum with small black spot close to lateral angles and dark spot in posterior part of disc. Margin
between lateral and posterior angles distinctly convex (curved outward). Elytra. Striae weakly marked, with
almost inconspicuous punctures. Lateral margins evenly curved outward (Fig. 95). Interstrial surface with weak,
medium-size foveiform punctures intermingled with dense microtubercles, some of which are confluent, shining
and forming a glossy zone. First interstria brighter than rest of elytra. Basal carina of seventh interstria weak and
short (Fig. 95), approximately the same length as metatibial spur. Length of apical carina of seventh interstria 1.5
times that of apex of sixth interstria. Abdomen. Punctures large and conspicuous, concentrated on anterior and
posterior margins of sternites and reduced to a narrow belt along midline of abdomen. Pygidium almost completely
covered by nearly confluent, large, foveiform punctures, leaving only a short, smooth medial line from base to
about midlength of pygidial surface. Apex of pygidium rounded. Legs. Protibia sharply widened along inner
margin from basal third to apex (Fig. 66). Middle protibial tooth closer to apical tooth than to basal tooth (Fig. 66).
Apical third of metatibia strongly curved inward (Fig. 67). Inner margin of metatibia with a longitudinal row of
small tubercles. Apex of metatibia with six clearly visible longitudinal carinae. Anterior margin of profemur
effaced at middle portion (Fig. 69). Most foveiform punctures of the profemur elliptical. Metatarsus approximately
one-third as long as metatibia. Secondary sexual characters. Male abdomen slightly more longitudinally flattened
than that of female. Apical margin of pygidium more dilated in male. Male protibial spur rounded at apex. Female
spur acuminate apically.
Material studied. Lectotype (MNHU) [here designated] : 1. [old white label, handwritten]. 8915; 2. [green
label, handwritten]. trisignatum Harold Brasilia; 3. [red label, bordered and printed in black, handwritten].
Lectotype Deltochilum trisignatum Harold des. F.Z. Vaz-de-Mello, 2014. Paralectotypes (MNHU) 2♀♀: ♀ 1. [old
white label, handwritten]. 8915; 2. [green label, handwritten]. Brasilia; 3. [yellow label, bordered and printed in
black, handwritten]. Paralectotype Deltochilum trisignatum Harold des. F.Z. Vaz-de-Mello, 2014./ ♀ 1. [old white
label, handwritten]. 8915; 2. [yellow label, bordered and printed in black, handwritten]. Paralectotype Deltochilum
trisignatum Harold des. F.Z. Vaz-de-Mello, 2014.
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Taxonomic notes. A lectotype is here designated in order to maintain nomenclatural stability by choosing the
only male (presenting characters that are easier to use for distinguishing from other species) among the available
syntypes as the single name-bearing type.
Non-type material [30]: ARGENTINA: TUCUMAN [probably mislabelled] (1 ♀ MNHN); BRAZIL:
BAHIA: Santa Teresinha, Pedra Branca, (12°51.33'S, 39°28.575'W), 2.II.2009, L.R.M. Oliveira (3 MZUEFS);
ESPÍRITO SANTO: P. Cachoeira, (2 ♀ MZUSP); Conceição da Barra, 22–28.X.1968, C. & C.T. Elias (1 ♀
DZUP); Linhares, II.1964 (1 ♂ DZUP); same, XII.1967 (1 ♀ DZUP); Linhares, Lagoa do Macuco, (19°03'50"S,
39°58'43"W), 26–27.I.2000, F. Génier & S. Ide (1 ♂ 3 ♀ CEMT, 2 ♀ CMN); Linhares, Povoação, (19°36'S,
39°48'W), 3.XII.2000, G. Schiffler (1♂ CEMT); MINAS GERAIS: Mar de Hespanha, 9.XII.1910, J.F. Zikán (1 ♀
CEMT, 1 ♂ MZUSP); Marliéria, Parque Estadual do Rio Doce, (19°42'09.22"S, 40°30'54.21"W), 11.I.2005, S.S.P.
Almeida & M.R. Pereira (10 ♂ 1 ♀ CEMT); RIO DE JANEIRO: Sto. Antonio dos Brotos, Districto de San Fidelis,
1876–1882, Auguste Vincent de Lyon (1 ♂ MNHN).
Sampling methods. Millipede carrion baited pitfall trap [3 specimens]; Flight interception trap [1]; un-baited
pitfall trap [11]. The un-baited pitfall traps contained formalin solution (as a preservative) may act as an attractant
for beetles predacious on millipedes, since millipedes often release defence secretions containing formaldehyde-
based compounds.
Habitat. Brazil (southern Bahia, Espirito Santo and Minas Gerais) (Fig. 89), from available data this species
inhabits lowland Atlantic forest, with sandy soil. A single specimen was collected from “cabruca” vegetation
(agricultural area where the cacao is planted in the midst of the natural vegetation of Atlantic forest). The known
elevation range is 10 to 130 m AMSL.
Diagnosis and remarks (within the trisignatum complex): head black with violet hue, pronotum yellowish-
brown with three distinct black spots, elytra reddishbrown; dorsal surface of head completely covered by closely
spaced to confluent foveiform punctures resulting in rugosity; margin of pronotum (close to lateral angles) curved
inward. The following aedeagus characters are also diagnostic: parameres shorter than phallobase, rounded at apex,
in lateral view forming an angle of approximately 30° with phallobase (Fig. 9). Lamella copulatrix: right lobe
"scaphopode" shaped (Fig. 35). Superior right peripheral sclerite (SRP) as in Fig. 44. Fronto-lateral peripheral
sclerite (FLP) as in Fig. 52. Center of genital segment with a well-sclerotized portion and "M"-shaped (Fig. 27).
There is some variation in the pronotum of four specimens examined, which display yellow coloration. The dark
spot on posterior portion of pronotal disc can be difficult to see in some individuals.
Deltochilum (Aganhyboma) kolbei Paulian 1938
(Figs 89, 96)
Deltochilum (Aganhyboma) kolbei Paulian 1938: 252, 254, Figs 4, 7, map n. 2 (original description, taxonomic comments and
identification key)
Vulcano & Pereira 1964: 642 (catalogue)
Vaz-de-Mello 2000: 192 (checklist)
Deltochilum kolbei: Blackwelder 1944: 203 (checklist)
Description. Body. Completely metallic green, with strong luster. Elytra a little darker than head and pronotum
(Fig. 96). Ventral surface dark green. Length 15 to 17 mm. Head. Dorsal surface with dense punctures close,
confluent on anterior and lateral portions. Punctures around clypeus larger and closer than those at center of dorsal
interocular surface. Anterior margin of each puncture not defined (impressed). Margin between clypeal teeth and
genae curved outward near clypeogenal suture. Thorax. Pronotum smooth, covered with small punctures,
apparently simple and uniformly distributed. Punctures on anterior angles larger, with a typical foveiform shape.
Margin of pronotum (close to lateral angles) curved inward. Margin between lateral and posterior angles distinctly
convex (curved outward). Posterior half of pronotum with rounded shape. Elytra. Smooth surface and strong
metallic bright (Fig. 96). Lateral margins evenly curved outward (Fig. 96). Elytral striae narrow, with carinate
margins close and almost touching. Punctures of elytral striae small and inconspicuous. Interstrial surface with
plenty small and scattered punctures. Surface of second interstria more irregular than the others. Basal carina of
seventh interstria prominent and short (Fig. 96). Length of apical carina of seventh interstria approximately two
times that of apical carina of sixth interstria. Apical tubercle of second interstria absent or weak. Abdomen.
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Punctures large and conspicuous on lateral portions of ventrites, concentrated on anterior and posterior margins and
reduced to a narrow belt along midline of abdomen. Pygidium with dense foveiform punctures over entire surface;
punctures small, transverse, elliptical and almost confluent. Apex of pygidium rounded. Legs. Protibia sharply
widened along inner margin from basal third to apex (as in Fig. 66). Middle protibial tooth closer to apical tooth
than to basal tooth (as in Fig. 66). Apical third of metatibia strongly curved inward (as in Fig. 67). Inner margin of
metatibia with a longitudinal row of small tubercles. Apex of metatibia with six clearly visible longitudinal carinae.
Anterior margin of profemur effaced at middle portion. Foveiform punctures of profemur with transverse elliptical
shape. Metatarsus with approximately one-half of length of metatibia. Secondary sexual characters. Since we
have been unable to examine a male specimen, it is not possible to verify which characters exhibit sexual
dimorphism. However, as this species is closely related to D. trisignatum, we believe the characters cited for D.
trisignatum will be similar.
Material studied. Holotype (MNHN) [examined]: BRAZIL: Bahia, Villa Victoria [Vitória da Conquista]
(CH. PUJOL, 1890; Col. R. OBERTHÜR).
Non-type material [5]: BRAZIL: PARAÍBA: Mataraca, (6°29'S, 34°56'W), IV.2004, V.H. Costa & B. Bellini
(1 ♀ CEMT); same, V.2004, V.H. Costa & B. Bellini (1 ♀ CEUFPB); Rio Tinto, Rebio Guaribas, (6°46'S,
35°04'W), III.2002, M.I.M. Hernández (2 ♀ CEUFPB); [RIO GRANDE DO NORTE] (1 ♀ CEMT).
Sampling methods. Flight Interception Trap [4 specimens].
Habitat. Brazil (Bahia, Paraíba and Rio Grande do Norte) (Fig. 89), from available data this species inhabits
lowland Atlantic forest with sandy soil.
Diagnosis and remarks (within the trisignatum complex): body completely metallic green, with strong luster
(Fig. 96); anterior margin of each head puncture not defined (impressed); margin of pronotum (close to lateral
angles) curved inward; elytra with smooth surface and strong metallic bright, without ocellate punctures
(microtubercles); surface of second interstria more irregular than the others, apical tubercle of second interstria
absent or weak.
The green coloration of this species is characteristic and unique among Aganhyboma. The strong bright
metallic color of elytra, among other characters, differentiates this species from D. violaceum.
Deltochilum (Aganhyboma) violaceum Paulian 1938
(Figs 70, 89, 97)
Deltochilum (Aganhyboma) violaceum Paulian 1938: 252, 253, Figs 7, 18 (original description, taxonomic comments and
identification key)
Vulcano & Pereira 1964: 642 (catalogue)
Vaz-de-Mello 2000: 192 (checklist)
Deltochilum violaceum: Blackwelder 1944: 203 (checklist)
Description. Body. Completely green or blue, with weak or opaque sheen. Elytra a little darker than head and
pronotum (Fig. 97). Ventral surface dark green. Length 15 mm. Head. Dorsal surface with dense punctures close,
confluent on anterior and lateral portions. Punctures around clypeus larger and closer than those at center of dorsal
interocular surface. Anterior margin of each puncture not defined (impressed). Margin between clypeal teeth and
genae curved outward near clypeogenal suture. Thorax. Pronotum smooth, covered with small punctures,
apparently simple and uniformly distributed. Punctures located on anterior angles larger, with a typical foveiform
shape. Margin of pronotum (close to lateral angles) curved inward. Margin between lateral and posterior angles
distinctly convex (curved outward). Posterior half of pronotum with rounded shape. Elytra. Slightly rugose due to
small and shiny microtubercles along interstriae (Fig. 70). Lateral margins evenly curved outward (Fig. 97). Elytral
striae narrow, with carinate margins close and almost touching. Punctures of elytral striae small and inconspicuous.
Interstrial surface with small and dispersed punctures intermingled with dense microtubercles, some of which are
confluent, shining and form an glossy zone (Fig. 70). Surface of second interstria more irregular than the others.
Basal carina of seventh interstria prominent and short (Fig. 97). Length of apical carina of seventh interstria
approximately two times that of apical carina of sixth interstria. Apical tubercle of second interstria present.
Abdomen. Punctures large and conspicuous on lateral portions of ventrites, concentrated on anterior and posterior
margins and reduced to a narrow belt along midline of abdomen. Pygidium with dense foveiform punctures over
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entire surface; punctures small, transverse, elliptical and almost confluent. Apex of pygidium rounded. Legs.
Protibia sharply widened along inner margin from basal third to apex (as in Fig. 66). Middle protibial tooth closer
to apical tooth than to basal tooth (as in Fig. 66). Apical third of metatibia strongly curved inward (as in Fig. 67).
Inner margin of metatibia with a longitudinal row of small tubercles. Apex of metatibia with six clearly visible
longitudinal carinae. Anterior margin of profemur effaced at middle portion. Foveiform punctures of profemur with
transverse elliptical shape. Metatarsus with approximately one-half of metatibia length.
Material studied. Holotype (MNHN) [examined]: BRAZIL (no locality given in original description). 1.
[old white label, handwritten]. Brésil; 2. [old white label, printed and bordered in black]. Ex Musaeo/ E. Allard/
1899; 3. [old white label, printed and bordered in black]. R. Paulian/ Vidit; 4. [old white label, handwritten]. D.
violaceum n. sp./ Type.; 5. [red label, printed in black]. Holotype.
Non-type material [1]: BRAZIL: MINAS GERAIS: Águas Vermelhas, XII.1997, A. Bello & F. Z. Vaz-de-
Mello (1 ♀ CEMT).
Sampling methods. millipede baited pitfall trap [1].
Habitat. Brazil (Minas Gerais) (Fig. 89), the single specimen with data was collected in “Carrasco”. This
habitat consists of dense shrubby xerophilous vegetation that occurs in semi-arid domain of Brazil.
Diagnosis and remarks (within the trisignatum complex): This species can be confused with D. kolbei.
However, D. violaceum can be easily distinguished by the following characters: elytra with dense and shiny
microtubercles (Fig. 70); elytra opaque or with weak luster; apical tubercle of second interstria present.
Deltochilum (Aganhyboma) amandaarcanjoae new species
(Figs 2, 8, 89, 98)
Etymology. Named in honour of Amanda Arcanjo, wife of the first author and scarabaeine cytogeneticist.
Description. Body. Completely bluish-black or violaceous, with weak sheen or opaque. Elytra a little darker
than head and pronotum (Fig. 98). Ventral surface black, with blue or green sheen. Length 13 mm. Head. Dorsal
surface with dense, but not confluent, foveiform punctures separated at center of dorsal interocular surface by
approximately twice their diameter. Punctures located near the clypeus larger and closer compared to those from
center of interocular surface. Anterior margin of each puncture not defined (impressed). Margin between clypeal
teeth and genae curved outward approximately at halfway its length. Thorax. Pronotum with smooth aspect,
covered by small punctures, apparently simple and uniformly distributed. Punctures on anterior angles larger, with
a typical foveiform shape. Margin of pronotum (close to lateral angles) straight, without depression. Margin
between lateral and posterior angles distinctly convex (curved outward). Posterior half of pronotum with rounded
shape (Fig. 98). Elytra. Lateral margins evenly curved outward (Fig. 98). Elytral striae narrow and shiny, with
carinate margins close and almost touching. Punctures of elytral striae small. Interstrial surface with small,
dispersed foveiform punctures amid small bright ocellate punctures, randomly distributed. First interstria brighter
than rest of elytra. Surface of second interstria with regular aspect. Basal carina of seventh interstria prominent and
short (Fig. 98). Length of apical carina of seventh interstria approximately two times that of apical carina of sixth
interstria. Apical tubercle of second interstria present. Abdomen. Punctures large and conspicuous, concentrated
anterolaterally on each sternite and reduced to a narrow belt along midline of abdomen. Pygidium with foveiform
punctures over entire surface; punctures transverse, elliptical and almost confluent. Apex of pygidium rounded.
Legs. Protibia widened along inner margin from basal third to apex; widened portion weakly indicated. Middle
protibial tooth closer to apical tooth than to basal tooth. Apical third of metatibia curved inward; curvature softer
than in other species of this complex (Fig. 98). Inner margin of metatibia with a longitudinal row of small tubercles.
Apex of metatibia with five clearly visible longitudinal carinae. Anterior edge of profemur completely emarginate,
with abundant setae for about two-thirds of its length. Foveiform punctures of profemur with transverse elliptical
shape. Metatarsus with approximately one-half of metatibia length.
Material studied. Holotype: BRAZIL: MATO GROSSO: Rosário Oeste, XII.1959 (1 ♂ DZUP).
Paratype [1]: PARAGUAY: CONCEPCIÓN: Horquetá, 1964, Martínez (1 ♂ CMN).
Diagnosis and remarks (within the trisignatum complex): bluish-black or violaceous coloration (Fig. 98);
margin of pronotum (close to lateral angles) straight, lacking depression; anterior edge of profemur completely
emarginate. The following aedeagus characters are also diagnostic: parameres short, rounded at apex, in lateral
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view forming an angle of approximately 20° with phallobase (Fig. 8). Lamella copulatrix: left lobe “J”-shaped (as
in Fig. 33); right lobe baton-shaped, slightly curved (as in Fig. 33). Fronto-lateral peripheral sclerite (FLP) as in
Fig. 50. Center of genital segment with a well-sclerotized portion and "T"-shaped (Fig. 2).
cupreicolle complex
Diagnosis. Apex of pygidium acuminate (Fig. 73); protibia not abruptly expanded along inner margin; middle
protibial tooth equidistant between apical and basal teeth; apical third of metatibia only slightly curved inward (Fig.
68).
Deltochilum (Aganhyboma) cupreicolle (Blanchard 1841)
(Figs 5, 25, 33, 50, 72, 90, 99)
Hyboma cupreicolle Blanchard 1841: plate 10, Fig. 4; Blanchard 1845: 157 (original description)
Canthon cupricolle Lucas 1859: 99 (original description, synonymy)
Deltochilum cupreicolle: Harold 1869a: 995 (catalogue)
Harold 1869b: 57 (synonymized Canthon cupricolle with Deltochilum cupreicolle)
Bruch 1911: 185 (catalogue)
Gillet 1911: 35 (catalogue)
Blackwelder 1944: 202 (checklist)
Deltochilum (Aganhyboma) cupricolle: Kolbe 1893: 192
Shipp 1897: 195
Vulcano & Pereira 1967: 556 (identification key)
Deltochilum (Aganhyboma) cupreicolle: Paulian 1938: 240, 252, 253, 255, Figs 7, 10, 18, map No 2 (description, taxonomic
comments and identification key)
Lane 1946: 173, 174 (taxonomic comments)
Martínez 1959: 51 (catalogue)
Vulcano & Pereira 1964: 641 (catalogue)
Vaz-de-Mello 2000: 192 (checklist)
Description. Body. Head dark green with weak sheen or opaque, pronotum in copper tone, with black or greenish
spots, elytra reddish-brown, with an opaque sheen (Figs 72, 99). Ventral surface black with a greenish sheen.
Length 11 to 13 mm. Head. Dorsal surface with dense foveiform punctures, not confluent, separated at center of
dorsal interocular surface by approximately twice their diameter. Punctures of variable size; largest around the
eyes, smallest closer to antero-lateral portion, near the clypeus. Anterior margin of each puncture not defined (as in
Fig. 60). Margin between clypeal teeth and genae curved outward approximately at halfway of its length. Thorax.
Surface of pronotum with small punctures, apparently simple and uniformly distributed. Punctures located on
anterior angles with a typical foveiform shape. Pronotum with a narrow greenish band extending the entire edge.
Posterior edge with band "I"-shaped (Fig. 99); this band can extend up to the disc of pronotum. Margin of
pronotum (close to lateral angles) straight and emarginated, without depression. Elytra. Lateral margins evenly
curved outward, with narrow dark spot, more visible on anterior portion. Striae faintly visible at a small
magnification (Fig. 72). Interstrial surface with small, dispersed foveiform punctures amid small bright ocellate
punctures, randomly distributed. First interstria brighter than rest of elytra. Basal carina of seventh interstria short,
length about twice the width of the seventh interestria (Fig. 72). Length of apical carina of seventh interstria
approximately 2.5 times that of apical carina of sixth interstria. Abdomen. Punctures large and conspicuous,
concentrated anterolaterally on each sternite, reduced to a narrow belt along midline of abdomen (as in Fig. 73).
Pygidium with dense foveiform punctures, transverse, elliptical and almost confluent at base. Apex of pygidium
acuminate (as in Fig. 73). Legs. Protibia not abruptly widened (Fig. 99). Middle protibial tooth equidistant between
apical and basal teeth. Protibial spur acuminate at apex. Apical third of metatibia only slightly curved inward (as in
Fig. 68), with five clearly visible longitudinal carinae. Inner margin of metatibia with a longitudinal row of small
tubercles. Anterior edge of profemur completely emarginate, with abundant setae in two-thirds of its length.
Foveiform punctures of profemur with transverse elliptical shape. Metatarsus approximately two-thirds as long as
metatibia. Secondary sexual characters. Male abdomen more longitudinally flattened than that of female. Males
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may possess a small tubercle medially between first and second ventrites. Apical margin of pygidium longer in
male. Protibial spur more acuminate in female.
Material studied. Holotype of Hyboma cupreicolle Blanchard (MNHN) [examined]: Andes (no locality
given in original description). 1. [red label, printed in black]. Type; 2. [green label, handwritten]. Deltochilum
cupreicolle Blanch; 3. [old white label, handwritten]. Deltochilum cupreicolle Blanch; 4. [old green label, round].
Holotype of Canthon cupricolle Lucas (MNHN) [examined]: (no locality given in original description). 1.
[old white label, handwritten]. Canthon cupricolle Luc.; 2. [old green label, round].
Non-type material [18]: BOLÍVIA: SANTA CRUZ: Andrés Ibañez, (17º43'S, 63°14'W), 15.II.1997, Paolo
Bettella (1 ♂ CEMT); Chiquitos (1 MNHN); Chiquitos, Santiago de Chiquitos, (18º19´S, 59º34´W), 11.XI.2008, T.
Vidaurre (1 ♀ CEMT, 1 ♀ CREN, 2 ♀ MNKM); same data, W.D. Edmonds & T. Vidaurre (2 ♂ CEMT); San
Ignacio, (16°24'S, 61°10'W), 19.I.2010, T. Vidaurre (1 ♂ MNKM, 1 ♂ CREN). BRAZIL: [no other data] (1
MNHN); GOIÁS: Colinas do Sul, Serra da Mesa, (14°01'S, 48°12'W), 15.XII.1995, C. Campaner (1 ♂ MZUSP);
MATO GROSSO: Cáceres, 18.XI.1984, C. Elias (3 ♂ DZUP); Cuiabá, Faz. Santhidi, 14.III.2009, (15°23'06"S,
56°06'53"W), L. R. Silva (1 ♀ CEMT); Barra dos Bugres, R. E. Serra das Araras, 16.II.1987, Y.O. Willis (1 ♀
CEMT); SÃO PAULO: Campinas, III.1943, M. Autuori (1 ♀ DZUP).
Sampling methods. baited pitfall trap with injured diplopods [6 specimens].
Habitat. Bolivia and Brazil (Goiás and Mato Grosso) (Fig. 90), from available data this species inhabits
Cerrado and Chaco. Six individuals were collected in "Chiquitano Cerrado".
Diagnosis and remarks (within the cupreicolle complex): basal carina of seventh interstria short, length about
twice the width of the seventh interestria (Fig. 72); elytra reddish-brown or dark brown, with an opaque sheen (Fig.
99). The following aedeagus characters are also diagnostic: parameres shorter than phallobase, slightly rounded at
apex, in lateral view forming an angle of approximately 45° with phallobase (Fig. 5). Lamella copulatrix (LC): left
lobe “J”-shaped (Fig. 33); right lobe baton-shaped (Fig. 33). Fronto-lateral peripheral sclerite (FLP) as in Fig. 50.
Center of genital segment with a well-sclerotized portion and "T"-shaped (Fig. 25).
The coloration of pronotum can vary between dark brown and copper. The color of elytra can also vary from
reddish-brown to dark brown. Most specimens do not present a round spot on disc of pronotum. When this spot is
present, elytra are never dark brown.
Based on examination of the types, we can find no significant difference between Hyboma cupreicolle
Blanchard 1841 and Canthon cupricolle Lucas 1859, we therefore confirm that C. cupricolle Lucas is a junior
synonym of Deltochilum cupreicolle (Blanchard 1841).
Deltochilum (Aganhyboma) titovidaurrei new species
(Figs 4, 42, 90, 100)
Etymology. Named in honour of Tito Vidaurre Sánchez, a friend and collaborator in scarabaeine studies.
Description. Body. Head dark green with a weak sheen or opaque, pronotum in copper tone, with black or
greenish spots, elytra dark brown, with an opaque sheen (Fig. 100). Ventral surface black. Length 11 to 12 mm.
Head. Dorsal surface with dense foveiform punctures, not confluent, separated at center of dorsal interocular
surface by approximately twice their diameter. Punctures of variable size; largest around the eyes, and smallest on
the anterolateral portion of the head, near the clypeus. Anterior margin of each puncture not defined (as in Fig. 60).
Margin between clypeal teeth and genae curved outward approximately at halfway of its length. Thorax. Surface
of pronotum with small punctures, apparently simple and uniformly distributed. Punctures on anterior angles
larger, with a typical foveiform shape. Pronotum with a narrow greenish band extending the entire edge; Disc with
transverse oval spot (Fig. 100). Margin of pronotum (close to lateral angles) straight and emarginated, without
depression. Margin between lateral and posterior angles approximately straight. Elytra. Lateral margins evenly
curved outward (Fig. 100). Striae punctate, however faintly visible at a small magnification. Interstrial surface with
small, dispersed foveiform punctures, amid small bright ocellate punctures, randomly distributed. First interstria
slightly brighter than rest of elytra. Basal carina of seventh interstria short, length about twice the width of the
seventh interestria (as in Fig. 72). Length of apical carina of seventh interstria with approximately 2.5 times that of
apical carina of sixth interstria. Abdomen. Punctures large and conspicuous, concentrated anterolaterally on each
sternite, reduced to a narrow belt along midline of abdomen (as in Fig. 73). Pygidium with dense foveiform
punctures, transverse, elliptical and almost confluent at base. Apex of pygidium acuminate (as in Fig. 73). Legs.
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Protibia not abruptly widened. Middle protibial tooth equidistant between apical and basal teeth. Protibial spur
acuminate at apex. Apical third of metatibia only slightly curved inward (as in Fig. 68), with five clearly visible
longitudinal carinae. Inner margin of metatibia without longitudinal row of small tubercles. Anterior edge of
profemur completely emarginate, with abundant setae in two-thirds of its length. Foveiform punctures of profemur
with transverse elliptical shape. Metatarsus approximately two-thirds as long as metatibia. Secondary sexual
characters. Male abdomen more longitudinally flattened than that of female. Males may possess medially between
first and second ventrites, a small tubercle. Apical margin of pygidium longer in male. Protibial spur more
acuminate in female.
Material studied. Holotype: BRAZIL: MINAS GERAIS: Lavras, near Parque Ecológico Quedas do Rio
Bonito, (21°20'21.3"S, 44°58'38.8"W), 10.II.2009, Fernando Silva & Tito Vidaurrei (1 ♂ CEMT).
Paratypes [5]: BRAZIL: GOIÁS: Brazlândia, I.1994, E. Grossi (1 ♂ CEMT); Jatahy, 1895–96, Ch. Pujol (2
MNHN); MINAS GERAIS: Lavras, 15.II.1999, Leila A.S. (1 ♂ CEMT); Lavras, near Parque Ecológico Quedas do
Rio Bonito, (21°20'21.3"S, 44°58'38.8"W), 10.II.2009, Fernando Silva & Tito Vidaurrei (1 ♀ CREN).
Sampling methods. baited pitfall trap with injured diplopods [2 specimens].
Habitat. Brazil (Goiás and Minas Gerais) (Fig. 90), from available data this species inhabits Cerrado. Two
individuals were collected in "Campo Limpo", Cerrado, 1,170 m AMSL.
Diagnosis and remarks (within the cupreicolle complex): basal carina of seventh interstria short, length about
twice the width of the seventh interestria (as in Fig. 72); elytral coloration can vary from dark brown to black; disc
of pronotum with transverse oval spot (Fig. 100). The following aedeagus characters are also diagnostic: parameres
shorter than phallobase, slightly rounded at apex, in lateral view forming an angle of approximately 30° with
phallobase (Fig. 4). Lamella copulatrix: left lobe “J”-shaped (as in Fig. 33); right lobe baton-shaped (as in Fig. 33).
Superior right peripheral sclerite (SRP) as in Fig. 42. Center of genital segment with a sclerotized portion and "T"-
shaped.
The specimens present the parameres and phallobase shorter or truncated than D. cupreicolle.
Deltochilum (Aganhyboma) viridescens Martínez 1948 new status
(Figs 6, 26, 34, 43, 51, 60, 64, 68, 71, 73, 90, 101, 102, 103)
Deltochilum cupreicolle ssp. viridescens Martínez 1948, 171 (original description)
Deltochilum (Aganhyboma) cupreicolle viridescens Martínez 1959: 52 (catalogue)
Martínez 1987: 55 (checklist)
Deltochilum cupreicolle viridescens Silva et al. 2012: 1359–1367 (predatory behaviour study)
Description. Body. Head and elytra dark green, with a metallic sheen. Pronotum light brown, with green spots
(Fig. 101). Ventral surface with metallic dark green. Length 12 to 14 mm. Head. Dorsal surface with dense
foveiform punctures not confluent, separated at center of dorsal interocular surface by approximately twice their
diameter (Fig. 60). Punctures of variable size; largest around the eyes and smallest on the anterolateral portion, next
to the clypeus. Anterior margin of each puncture not defined (Fig. 60). Margin between clypeal teeth and genae
curved outward approximately at halfway of its length. Thorax. Surface of pronotum with small punctures,
apparently simple and uniformly distributed. Punctures located on anterior angles larger, with a typical foveiform
shape. Pronotum with a narrow greenish band extending the entire edge; posteromedial portion with a semi-circular
band which extends up to half of pronotal disc (Fig. 101). Margin of pronotum (close to lateral angles) straight and
emarginated, without depression. Margin between lateral and posterior angles approximately straight. Elytra.
Slightly elongate (Figs 101–103). Striae well impressed and punctate (Fig. 71). Interstrial surface with small,
dispersed foveiform punctures, amid small bright ocellate punctures, randomly distributed. First interstria brighter
than rest of elytra. Basal carina of seventh interstria prominent and elongate, extending a little further than humeral
callus portion (Fig. 71). Length of apical carina of seventh interstria three times that of apical carina of sixth
interstria. Abdomen. Punctures large and conspicuous, concentrated anterolaterally on each sternite, reduced to a
narrow belt along midline of abdomen (Fig. 73). Foveiform punctures of pygidium denser on medial portion, some
confluent. Apex of pygidium strongly acuminate (Fig. 73). Legs. Protibia not abruptly widened (Fig. 101). Middle
protibial tooth equidistant between apical and basal teeth. Protibial spur acuminate at apex. Apical third of
metatibia only slightly curved inward (Figs 68, 101), with five clearly visible longitudinal carinae. Inner margin of
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metatibia without longitudinal row of small tubercles. Anterior edge of profemur completely emarginate, with
abundant setae in two-thirds of its length. Foveiform punctures of metafemur with transverse or oblique elliptical
shape (Fig. 64). Metatarsus with two-thirds of metatibia length. Secondary sexual characters. Male abdomen
more longitudinally flattened than that of female. Males may possess medially between first and second ventrites, a
small tubercle. Apical margin of pygidium longer in male. Protibial spur more acuminate in female.
Material studied. Holotype (MACN) [not examined]: ARGENTINA: Prov. Salta, Depto. Rosario de la
Frontera, El Naranjo. The original description, as well as the examination of a specimen of CMN which was
compared to the holotype, are sufficient to confirm the identification of this species.
Non-type material [55]: BRAZIL: BAHIA [doubtful]: Barro Preto, Ch. Pujol (8 MNHN); DISTRITO
FEDERAL: Universidade de Brasília (1 ♂ CEMT); GOIÁS: Jataí, 1955 (1 ♀ MZUSP); Jatahy, 1895–96, Ch. Pujol
(1 MNHN); Mineiros, Parque Nacional das Emas, (18°04'S, 52°55'W), 15–19.XII.1987, expedição MZUSP (1 ♂ 1
♀ MZUSP); MATO GROSSO: Cuyaba (1 MNHN); Cuiabá, XII.1945 (1 ♀ MZUSP); Chapada dos Guimarães,
X.1937, Pinto (1 ♀ MZUSP); Chapada dos Guimarães, 27.X.1961, F.M. Oliveira (4 7 ♀ DZUP); Chapada dos
Guimarães, XI.1963, Alvarenga & Werner (1 ♀ MZUSP); Rosário Oeste, XII.1959, (1 ♂ DZUP); Uirapuru,
Chapada dos Parecis, XII.2002, A. Foucart (1 ♀ CEMT); 30 Km norte de Uirapuru, (14°17'S, 59°15'W),
15.XII.2001, A. Foucart (3 ♂ CEMT); SÃO PAULO, Itirapina, 15.III.2000, (1 ♂ CEMT); Itirapina, II.2000, G.
Machado (2 ♀ CEMT); Itirapina, Estação Ecológica, (22°13'S, 47°54'W), 30.I.2009, Fernando Silva (6 ♂ 9 ♀
CEMT). PARAGUAY: AMAMBAY: Cerro Corá, XII. 1993, Drechsel (1 ♂ CEMT); CAAGUAZÚ: Caaguazú,
XII.1977, Martínez/Fritz (1 ♀ CMN); CONCEPCIÓN: Horqueta, 1934, Schulze (1 ♀ CMN); GUAIRA: Villarrica,
II.1922 (1 ♂ CEMT).
Sampling methods. baited pitfall trap with injured diplopods [15 specimens]; flight interception trap [3].
Habitat. Brazil (Bahia, Distrito Federal, Goiás, Mato Grosso and São Paulo) and Paraguay (Fig. 90), from
available data this species inhabits Cerrado and Chaco. 15 individuals were collected in "Campo Sujo", Cerrado,
731 m AMSL.
Diagnosis and remarks (within the cupreicolle complex): head dark green, with a metallic sheen; basal carina
of seventh interstria elongate, extending a little further than humeral callus portion (Fig. 71); apex of pygidium
strongly acuminate in male (Fig. 73). The following aedeagus characters are also diagnostic: parameres shorter
than phallobase, rounded at apex, in lateral view forming an angle of approximately 30° with phallobase (Fig. 6).
Lamella copulatrix (LC): left lobe “J”-shaped (Fig. 34); right lobe with shape similar to a human leg (Fig. 34).
Superior right peripheral sclerite (SRP) as in Fig. 43. Fronto-lateral peripheral sclerite (FLP) as in Fig. 51. Center
of genital segment with two well-sclerotized portions with oval shape (Fig. 26).
Pronotum can be yellow, red or brown. Posteromedial portion of pronotum can show only a small and
inconspicuous greenish band.
Body coloration can be completely green (Fig. 102). Through the study of original description, as well as the
examination of a specimen of CMN, which had been compared with the holotype, we concluded that the type
specimen (entirely green) is a variant of the most common coloration pattern for species.
Deltochilum (Aganhyboma) viridicatum new species
(Figs 7, 34, 90, 104, 105)
Pereira & Martínez 1956: 93 (comment on predation of Diplopoda; referred as D. kolbei “misidentification”)
Martínez 1959: 52 (catalogue; cites Pereira & Martínez 1956)
Halffter 1959: 166 (comment on predation of Diplopoda; cites Pereira & Martínez 1956)
Halffter & Matthews 1966: 42 (comment on predation of Diplopoda; cites Pereira & Martínez 1956)
Cano 1998: 176 (cites Pereira & Martínez 1956)
Etymology. The green body color inspired the species name.
Description. Body. Head dark green or black-violaceous, with weak sheen or opaque, pronotum dark brown,
with dark green spots, elytra dark green (Fig. 104). Ventral surface dark green, with a smooth metallic sheen.
Length 13 to 14 mm. Head. Surface with dense foveiform punctures, not confluent, separated at center of dorsal
interocular surface by approximately one or two times their diameters. Punctures of variable size; largest around
eyes and smallest closer to clypeus. Anterior margin of each puncture not defined (as in Fig. 60). Margin between
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clypeal teeth and genae curved outward approximately at halfway of its length. Thorax. Surface of pronotum with
small punctures, apparently simple and uniformly distributed. Punctures located on anterior angles larger, with a
typical foveiform shape. Pronotum with a narrow greenish band extending the entire edge (Fig. 104). Margin of
pronotum (close to lateral angles) straight and emarginated, without depression. Margin between lateral and
posterior angles approximately straight. Elytra. Slightly elongate (Figs 104, 105). Striae well impressed and
punctate. Interstrial surface with small, dispersed foveiform punctures, amid small bright ocellate punctures,
randomly distributed. First interstria brighter than rest of elytra. Basal carina of seventh interstria prominent and
elongate, extending a little further than humeral callus portion (as in Fig. 71). Length of apical carina of seventh
interstria not surpassing three times that of apical carina of sixth interstria. Abdomen. Punctures large and
conspicuous, concentrated anterolaterally on each sternite, reduced to a narrow belt along midline of abdomen (as
in Fig. 73). Pygidium with foveiform punctures denser on medial portion, near the base. Apex of pygidium weakly
acuminate or rounded. Legs. Protibia not abruptly widened. Middle protibial tooth equidistant between apical and
basal teeth. Protibial spur acuminate at apex. Apical third of metatibia only slightly curved inward (as in Fig. 68),
with five clearly visible longitudinal carinae. Inner margin of metatibia without longitudinal row of small tubercles.
Anterior edge of profemur completely emarginate, with abundant setae in two-thirds of its length. Foveiform
punctures of profemur with transverse elliptical shape. Metatarsus longer than two-thirds of metatibia length.
Secondary sexual characters. Male abdomen more longitudinally flattened than that of female. Males may
possess medially between first and second ventrites, a small tubercle. Apical margin of pygidium longer in male.
Protibial spur more acuminate in female.
Material studied. Holotype: ARGENTINA: MISIONES: San Pedro, I. 1956 (1 ♂ CEMT).
Paratypes [7]: ARGENTINA: MISIONES: Loreto, 1955, F. H. Walz (1 ♀ MZUSP); Loreto, XI.1958,
Martínez (1 ♀ CEMT); Loreto, A. Ogloblin (1 ♂ MZUSP); [no indication of city], II.1954, F. H. Walz (1 ♂ CMN).
BRAZIL: RIO GRANDE DO SUL: Neu Wurttemberg [currently Panambi], 1915, E. Garbe (1 ♂ MZUSP);
Guaíba, 1976, M.H. Galileo (1 ♀ CEMT). URUGUAY [doubtful]: Puntas, Arenal (1 ♂ MZUSP).
Habitat. Argentina, southern Brazil and Uruguay [doubtful] (Fig. 90). Only one specimen with label data
"Mata de galeria" [riparian forest].
Diagnosis and remarks (within the cupreicolle complex): head dark green or black-violaceous, with a weak
sheen or opaque; apex of pygidium weakly acuminate or rounded; basal carina of seventh interstria prominent and
elongate, extending a little further than humeral callus portion (as in Fig. 71), though somewhat shorter and
inconspicuous on the basis than in D. viridescens. The following aedeagus characters are also diagnostic:
parameres shorter than phallobase, rounded at apex, in lateral view forming an angle of approximately 30° with
phallobase (Fig. 7); parameres more enlarged at the base in lateral view (Fig. 7) than in D. viridescens. Lamella
copulatrix (LC): left lobe “J”-shaped (Fig. 34); right lobe with shape similar to a human leg (Fig. 34). Fronto-
lateral peripheral sclerite (FLP) darkly sclerotized (as in Fig. 51). Center of genital segment with two well-
sclerotized portions with oval shape.
We examined a specimen with label data "Puntas, Arenal" and we believe this location is Puntas del Arenal,
Uruguay. This specimen is black (Fig. 105), and the foveiform punctures on central portion of head are extremely
close.
The species can show other variations in pattern of colors. Pronotum of some individuals can vary between
dark and light brown or reddish. Elytra can vary between dark green and black. Ventral surface may also vary from
dark green and black.
valgum group
Description. Body. Dorsal surface with foveiform punctures of variable size and density. Head, pronotum, elytra
and ventral surface with a black, copper or red coloration, with or without reddish, brownish, greenish sheen or
copper tones. Length 12 to 16.5 mm. Head. Surface with strong, deep and dense punctures (Fig. 61), almost
confluent in some species. Foveiform punctures of head with well-defined margin (Fig. 61). Margin of head, near
clypeal teeth, straight and followed by a small tooth slightly pronounced (Fig. 61). After this tooth, margin slightly
curved inward until the junction of clypeus with gena. Dorsal interocular distance approximately three or four
times eye width (Fig. 61). Thorax. Margin between anterior and lateral angles elongate and oblique (Fig. 85),
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approximately straight, with visible row of setae in dorsal view. Margin between lateral and posterior angles
straight or slightly rounded (curved outward). Posterior margin, after the posterior angle, slightly curved outward.
Pronotum surface with deep and dense foveiform punctures. Punctures smaller and more dispersed near the
anteromedial portion of pronotum (Fig. 85). Hypomera with foveiform punctures denser on anterior portion and
somewhat more dispersed on posterior portion (Figs 81–83). Elytra. Oval shape, irregular surface (slightly rugose)
and lateral margins ranging from semi-parallel to convex. Striae thick and well delimited (Fig. 75) or narrow and
almost inconspicuous (Fig. 74). Interstriae ranging from distinctly convex to flattened, with uniform punctures
along the width (Fig. 75), or more punctate on the sides, close to striae (Fig. 86). Interstriae of most species with
ocellate punctures (microtubercles), or shiny and glossy zones poorly defined amid foveiform punctures (Figs
74–77). Basal carina of seventh interstria short or elongate. Basal carina of ninth interstria, when present in three-
quarters of elytra, can reach the anterior limit of apical carina of seventh interstria. Basal carina of ninth interstria
never traverses the entire length of elytra (Fig. 63). In some species, basal carina of ninth interstria reaches no
farther than mid length of elytra (Fig. 80). Posterior part of elytral lateral margin formed by the pseudepipleural
carina (Fig. 80). Apices of interstriae 2, 3, 4, 5, 6 and 7 with tubercle or carina. Apical carina of second interstria
absent in some species. Apical carina of third interstria can be more evident than apical carinae of second and
fourth interstriae (Fig. 88); species which this carina is more evident, it presents a rounded form, resembling a
tubercle. Length of apical carina of sixth interstria longer than or equal to that of apical carina of fifth interstria.
Pseudepipleural carina complete, visible entire length of epipleuron in most species (Fig. 63); in some species this
carina may be interrupted at basal third, reappearing next to one-half of elytral length (Fig. 80). Abdomen.
Punctures large and conspicuous on lateral portion of ventrites; sparser or absent along midline. Punctures reduced
on medial portion to one row on anterior margin. Length of sixth ventrite, measured at midline, longer than that of
fifth ventrite. Pygidium with dense rounded punctures over entire surface. Apex of pygidium acuminate. Legs.
Most foveiform punctures of the metafemur rounded and dense (Fig. 65). Anterior edge of profemur completely
emarginate. Protibia not abruptly widened, with dense foveiform punctures from base to first tooth. Medial lateral
tooth of protibia closer to apical tooth than to basal tooth. Metafemora just slightly narrowed at base (Fig. 65).
Apical third of metatibia strongly curved inward (as in Fig. 67). Inner margin of metatibia without longitudinal row
of small tubercles, or tubercles inconspicuous. Metatarsus with approximately one-half of metatibia length.
Secondary sexual characters. Male abdomen more longitudinally flattened than that of female. Males with a
small tubercle at medial portion of abdomen between first and second ventrites. Protibial spur broader and more
curved in male. Inner margin of metatibia usually with a longitudinal row of small tubercles more conspicuous in
male. Aedeagus. Parameres symmetrical, simple, without ornamentation or setae, elongate or short (Figs 10–23).
Endophallus: lobe in submedial portion. Lamella copulatrix (LC): subrectangular or subtrapezoidal sclerites (Figs
36–41). Superior right peripheral sclerite (SRP): circular shaped, sub-straight cable and a small sclerotized
appendix. Ring with margin usually narrow, but never "thread"-shaped (Figs 45–49). Fronto-lateral peripheral
sclerite (FLP): two irregular sclerites with a sclerotized cover that surrounds it (Figs 53–58). Complex of axial and
subaxial sclerites (A+SA): several superposed and elongate sclerites. Genital segment: lateral arms sclerotized in a
scalene triangle shape; central portion “M”-shaped; superior margin weakly bounded, with weak sclerotization
(Figs 28–32).
Diagnosis. Compared to the trisignatum group, the diagnostic characters of valgum group are: Presence of
small clypeal teeth external to central teeth (Fig. 61); foveiform punctures of pronotal disc smaller than that of
lateral portion; posterior part of elytral lateral margin formed by the pseudepipleural carina (Figs 63, 80); lamella
copulatrix: subrectangular or subtrapezoidal sclerites (Figs 36–41); ring of superior right peripheral sclerite (SRP)
with thick and regular margin, never similar to a “thread” (Figs 45–49); SRP with a small appendix (Figs 45–49).
acropyge complex
Diagnosis. Posteromedial punctures of pronotum not touch the posterior margin of pronotum. Basal carina of ninth
interstriae extending at most to midway along elytra (Fig. 80)
Deltochilum (Aganhyboma) acropyge Bates 1887 new subgeneric placement and new status
(Figs 13, 46, 91, 106)
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Deltochilum acropyge Bates 1887: 36, plate 2, Fig. 15 (original description)
Gillet 1911: 35 (catalogue)
Blackwelder 1944: 202 (checklist)
Blackwelder 1973: 5 (checklist)
Deltochilum (Deltochilum) valgum var. acropyge: Paulian 1938: 269, 273 (taxonomic comments and identification key)
Deltochilum (Deltohyboma) valgum acropyge: Howden 1966: 733, 735 (identification key, taxonomic comments)
Deltochilum valgum acropyge: Howden & Young 1981: 36, 39 (identification key, description)
Cano 1998: 174, 175, 176, 177, Figs 2 and 3 (comment on habitat, distribution and behavior of predation)
Halffter 2003: 27, 28, Fig. 2.5 (description)
Description. Body. Head, pronotum, elytra and ventral surface black, with a silky sheen (Fig. 106). Length > 14
mm. Head. Punctures dense, almost touching; foveiform punctures separated at dorsal interocular surface by
approximately half diameter. Punctures around clypeus shallower, smaller and more dispersed than those on
interocular surface. Lateral margin of gena, nearby eyes, without row of setae or with almost inconspicuous setae.
Dorsal interocular distance approximately three times eye width (as in Fig. 61). Thorax. Surface of pronotum with
dense foveiform punctures, not touching each other. Punctures on the sides and anterior angles larger and
somewhat closer than that of pronotum disc. Discal punctures separated approximately by 1.5 or two times their
diameters. Anteromedial punctures smaller and more dispersed than discal punctures. Posteromedial punctures
usually separated by at least one diameter. Margin between anterior and lateral angles with row of small setae.
Margin between lateral and posterior angles straight, with approximately the same length of margin between
anterior and lateral angles. Hypomera with foveiform punctures much more dense on anterior portion;
anteromedial punctures separated by distance smaller or equal to one diameter; median portion with sparser
punctures. Diagonal band of hypomera from posteromedial portion to middle of lateral margin, with dispersed but
obvious punctures. Elytra. Lateral margins curved outward and elytral surface slightly irregular (rugose). Elytral
striae with carinate margins clearly separated. Interstriae flattened or faintly convex with foveiform punctures
surrounded by approximately four shiny microtubercles. Microtubercles with sheen more evident than surrounding
surface; some microtubercles indistinct. Foveiform punctures separate on the elytral disc by more than one
diameter. Basal carina of seventh interstria elongate, length about four times the width of the seventh interestria.
Basal carina of ninth interstria with approximately one-half of elytral length. Apical carina or tubercle of second
elytral interstria absent. Apical carina of third interstria weaker than apical carina of fourth interstria. Length of
apical carina of sixth interstria longer than that of apical carina of fifth interstria. Length of apical carina of seventh
interstria shorter than the sum of apical carinae of fifth and sixth interstriae. Pseudepipleural carina complete,
visible entire length of epipleuron (as in Fig. 63).
Material studied. Holotype (BMNH) [examined]: 1. [old white label, printed in black, bordered in red].
Type; 2. [old white label, printed in black]. B. Honduras./ Blancaneau.; 3. [old white label, printed in black]. Sp.
figured.; 4. [old white label, handwritten]. Deltochilum acropyge Bates.
Non-type material [2]: GUATEMALA: PETÉN: La Libertad, Comunidad Bethel, 14.VIII.1996, E. Cano (1
♂ 1 ♀ CEMT).
Sampling methods. baited pitfall trap with live diplopods [2 specimens].
Habitat. Belize, Costa Rica, Guatemala, Mexico and Panama (inhabits the Caribbean slope) [see too Halffter
2003; Kohlmann & Solis, 2012] (Fig. 91), from available data this species inhabits tropical rainforest.
Diagnosis and remarks (within the acropyge complex): Length > 14 mm; posteromedial punctures of
pronotum usually separated by at least one diameter; anteromedial punctures of hypomera usually separated by less
than one diameter; apex of second interstria without carina or tubercle; pseudepipleural carina complete, visible
entire length of epipleuron (as in Fig. 63). Elytra are slightly more convex than in other species of this complex,
with clearly impressed striae. The following aedeagus characters are also diagnostic: parameres symmetric and
quite elongate, in lateral view forming an angle of approximately 45° with phallobase; apex rounded (Fig. 13).
Lamella copulatrix (LC): subrectangular sclerites. Superior right peripheral sclerite (SRP) as in Fig. 46.
Deltochilum (Aganhyboma) feeri new species
(Figs 91, 107)
Etymology. Named in honour of François Feer, an excellent scarab ecologist who was responsible for collecting a
large number of the type specimens.
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Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with a silky sheen (Fig.
107). Length < 14 mm. Head. Punctures dense and almost touching; around clypeus shallower, smaller and more
dispersed than those on the dorsal interocular surface. Lateral margin of gena, around eyes, without row of setae or
with almost inconspicuous setae; margin with central portion clearly convex. Dorsal interocular distance
approximately three times eye width (as in Fig. 61). Thorax. Surface of pronotum with dense foveiform punctures,
nearly touching on the sides, anterior angles and posterior portion. Punctures of pronotal disc more dispersed than
those of lateral portion. Density, size and proximity of anteromedial punctures similar to those on disk.
Posteromedial punctures usually separated by less than one diameter. Margin between anterior and lateral angles
with row of small setae. Margin between lateral and posterior angles with approximately the same length of margin
between anterior and lateral angles. Hypomera with foveiform punctures dense at anterior, median and posterior
portions; punctures of anteromedial portion nearly touching. Diagonal band of hypomera from posteromedial
portion to middle of lateral margin with obvious punctures. Elytra. Lateral margins slightly curved outward, and
elytral surface slightly irregular (rugose). Elytral striae with carinate margins clearly separated. Interstriae
flattened, with foveiform punctures surrounded by approximately four well-defined shiny microtubercles.
Microtubercles with sheen more apparent than surrounding surface. Foveiform punctures separated on elytral disc
by one diameter or less. Basal carina of seventh interstria elongate, length about four times the width of the seventh
interestria. Basal carina of ninth interstria with approximately one-half of elytral length (as in Fig. 80). Apical
carina or tubercle of second elytral interstria absent. Apical carina of third interstria weaker than apical carina of
fourth interstria. Length of apical carina of sixth interstria longer than that of apical carina of fifth interstria. Length
of apical carina of seventh interstria less than the sum of lengths of apical carinae of fifth and sixth interstriae.
Pseudepipleural carina incomplete, effaced medially (as in Fig. 80).
Material studied. Holotype: FRENCH GUIANA: Nouragues, II.2001, F. Feer (1 ♂ CEMT).
Paratypes [10]: GUIANA: Isherton, 14.XI.1931 (2 ♀ MZUSP). TRINIDAD AND TOBAGO: Tunapuna, Mt.
St. Benedict, 21.VI.1993, S. & J. Peck (1 ♀ CMN). FRENCH GUIANA: Cayenne, (4°48'18"N, 28°52'41"W),
9.VI.1997, J. Ashe & R. Brooks (1 ♀ CMN); Nouragues, II.2001, F. Feer (1 ♂ CEMT); same, III.2002, F. Feer (1
♂ CEMT); same, III.2006, F. Feer (1 ♂ CEMT). SURINAM: SIPALIWINI DIST.: Kwamalasamutu region,
Sipaliwini River (Werehpai), 02°21'N, 56°41'W, CI RAP, 3–7.IX.2010, T. Larsen, 250m (1 ♂ CEMT); same but
Kutari River, 02°10'N, 56°47'W, ~250masl. CI RAP Survey, 18–24.VIII.2010 (1 ♂ AFIC); same but CI RAP 3
sites between 02°21'N, 56°41'W and 02°10'N, 56°47'W. ~250masl, VIII–IX.2010 (1 ♀ AFIC).
Sampling methods. flight interception trap [2 specimens].
Habitat. Guyana, French Guiana and Trinidad and Tobago (Fig. 91). One specimen with label data "Summit
Rain Forest".
Diagnosis and remarks (within the acropyge complex): length < 14 mm; foveiform punctures of head almost
touching, separated in dorsal interocular surface by about half diameter; posteromedial punctures of pronotum
usually separated by less than one diameter; anteromedial punctures of hypomera nearly touching; apical carina or
tubercle of second elytral interstria absent; pseudepipleural carina incomplete, effaced medially (as in Fig. 80)..
Hypomera punctures in individuals of this species, as well as in D. acropyge and D. schefflerorum new species, are
the most dense of acropyge complex.
Deltochilum (Aganhyboma) schefflerorum new species
(Figs 12, 29, 54, 81, 91, 108)
Etymology. Named in honour of Pamela Scheffler and Timothy Scheffler who collected a part of the type
specimens.
Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with a silky sheen.
Length 12 to 13.5 mm. Head. Punctures extremely dense and touching; around clypeus shallower, smaller and
more dispersed than those on the dorsal interocular surface. Lateral margin of gena, around eyes, without row of
setae or with almost inconspicuous setae; margin with central portion clearly convex. Dorsal interocular distance
approximately 2.5 times eye width. Thorax. Surface of pronotum with dense foveiform punctures, nearly touching
on the sides, anterior angles and posterior portion. Discal punctures only somewhat smaller and more dispersed
than those of lateral portion. Density, size and proximity of anteromedial punctures similar to those on disk.
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Posteromedial punctures usually separated by less than one diameter. Margin between anterior and lateral angles
with row of small setae. Margin between lateral and posterior angles with length somewhat smaller than that of
margin between anterior and lateral angles. Hypomera with foveiform punctures dense at anterior, median and
posterior portions (Fig. 81); anteromedial punctures nearly touching. Diagonal band of hypomera from
posteromedial portion to middle of lateral margin with obvious punctures (Fig. 81). Elytra. Lateral margins
slightly curved outward (Fig. 108), and surface slightly irregular (rugose). Elytral striae with carinate margins
clearly separated. Interstriae flattened, with foveiform punctures surrounded by approximately four well-defined
shiny microtubercles. Microtubercles with sheen more evident than surrounding surface. Foveiform punctures
separated on elytral disc by one diameter or less. Basal carina of seventh interstria elongate, length about four times
the width of the seventh interestria. Basal carina of ninth interstria with approximately one-half of elytral length (as
in Fig. 80). Apical carina or tubercle of second elytral interstria absent. Apical carina of third interstria weaker than
apical carina of fourth interstria. Length of apical carina of sixth interstria longer than that of apical carina of fifth
interstria. Length of apical carina of seventh interstria less than the sum of lengths of apical carinae of fifth and
sixth interstriae. Pseudepipleural carina incomplete, effaced medially (as in Fig. 80).
Material studied. Holotype: BRAZIL: MATO GROSSO: Cotriguaçu, Faz. S. Nicolau, (09°50'53"S, 58°
14'36"W), 12.X.2009, F. Vaz-de-Mello (1 ♂ CEMT).
Paratypes [9]: BRAZIL: MATO GROSSO: Alta Floresta, (09°35'55"S, 55°56'10"W), VI.2008, E. Berenguer
(1 ♂ CEMT); Cotriguaçu, Faz. São Nicolau, (09°50'24"S, 58°15'10"W), 9.X.2009, F. Vaz-de-Mello (1 ♂ CEMT);
same, (09°49'22"S, 58°15'48"W), 08.XII.2010, F. Vaz-de-Mello (1 ♂ 1 ♀ CEMT); Cotriguaçu, V.2011, R. E.
Vicente (1 ♂ CEMT); PARÁ, Redenção, (07°46'S, 51°58'W), XI.1998, P & T. Scheffler (2 ♂ 2 ♀ CEMT).
Sampling methods. flight interception trap [4 specimens]; at light-trap [1]; collected manually on millipedes
[2].
Habitat. Brazil (Mato Grosso and Pará) (Fig. 91), from available data this species inhabits Amazon rainforest.
Diagnosis and remarks (within the acropyge complex): foveiform punctures of dorsal interocular surface
touching; anteromedial punctures of pronotum separated by less than one diameter; diagonal band of hypomera
with obvious punctures (Fig. 81); apical carina or tubercle of second elytral interstria absent; pseudepipleural
carina incomplete, effaced medially (as in Fig. 80). The following aedeagus characters are also diagnostic:
parameres symmetric and elongate, in lateral view forming an angle of approximately 45° with phallobase; apex
acuminate (Fig. 12). Genital segment with superior margin weakly sclerotized (Fig. 29). Lamella copulatrix (LC):
subrectangular sclerites. Fronto-lateral peripheral sclerite (FLP) as in Fig. 54.
Deltochilum (Aganhyboma) streblopodum new species
(Figs 91, 109)
Etymology. Streblopodum (crooked + legs), Latin adjective, refers to the posterior tibiae of individuals, which are
curved.
Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with an evident sheen.
Length < 13 mm. Head. Punctures extremely dense, almost touching; foveiform punctures separated at dorsal
interocular surface by half diameter or less. Punctures around clypeus shallower, smaller and more dispersed than
those on the interocular surface. Lateral margin of gena, around eyes, without row of setae or with almost
inconspicuous setae; margin with central portion clearly convex. Dorsal interocular distance approximately three
times eye width (as in Fig. 61). Thorax. Surface of pronotum with dense foveiform punctures, nearly touching on
the sides, anterior angles and posterior portion. Discal punctures somewhat smaller and more dispersed than those
of lateral portion. Density, size and proximity of anteromedial punctures similar to those on disk. Posteromedial
punctures separated by one diameter or less. Margin between anterior and lateral angles with row of small setae.
Margin between lateral and posterior angles with length somewhat smaller than that of margin between anterior
and lateral angles. Hypomera with dense foveiform punctures at anterior portion and absent at posteromedial
portion. Diagonal band of hypomera from posteromedial portion to middle of lateral margin lacking punctures or
with one or two isolated punctures on otherwise smooth surface (as in Fig. 83). Elytra. Lateral margins slightly
curved outward (Fig. 109) and surface slightly irregular (rugose). Elytral striae with carinate margins clearly
separated. Interstriae flattened, with foveiform punctures surrounded by approximately four well-defined shiny
microtubercles. Microtubercles with sheen more evident than surrounding surface. Foveiform punctures separated
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on elytral disc by one diameter or less. Basal carina of seventh interstria elongate, length about four times the width
of the seventh interestria. Basal carina of ninth interstria with approximately one-half of elytral length. Apical
carina or tubercle of second elytral interstria absent. Apical carina of third interstria weaker than apical carina of
fourth interstria. Length of apical carina of sixth interstria longer than that of apical carina of fifth interstria. Length
of apical carina of seventh interstria less than the sum of lengths of apical carinae of fifth and sixth interstriae.
Pseudepipleural carina incomplete, effaced medially (as in Fig. 80). Aedeagus. The specimen examined presented
the aedeagus weakly sclerotized, without defined morphology.
Material studied. Holotype: BRAZIL: AMAZONAS: Barcelos, Igarapé Erere/Coruja, (00°06'16"N,
63°51'01"W), 18–25.VI.2008, F.F. Xavier (1 ♂ INPA).
Sampling methods. at light-trap [1 specimen].
Habitat. Brazil (Amazonas) (Fig. 91), from available data this species inhabits Amazon rainforest.
Diagnosis and remarks (within the acropyge complex): margin of pronotum between lateral and posterior
angles narrower than that between anterior and lateral angles; diagonal band of hypomera lacking punctures or with
one or two isolated punctures on otherwise smooth surface (as in Fig. 83); apical carina or tubercle of second
elytral interstria absent; elytral striae with round punctures; pseudepipleural carina incomplete, effaced medially
(as in Fig. 80).
Deltochilum (Aganhyboma) longiceps Paulian 1938 new subgeneric placement and new status
(Figs 16, 37, 61, 80, 83, 84, 91, 110)
Deltochilum (Deltochilum) valgum var. longiceps Paulian 1938: 269, 273 (original description, identification key)
Deltochilum longiceps: Blackwelder 1944: 203 (checklist)
Contreras 1951: 222 (catalogue)
Deltochilum (Deltohyboma) valgum longiceps: Vulcano & Pereira 1967: 557 (identification key)
Deltochilum valgum longiceps: Cano 1998: 177 (comment on feeding behavior)
Escobar 2003: 131 (observations of feeding behavior, referred as Deltochilum acropyge, "misidentification")
Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with an evident sheen. Length
> 13 mm. Head. Punctures dense, but not touching each other; foveiform punctures separated at the dorsal
interocular surface by approximately one diameter (Fig. 61). Punctures around clypeus shallower, smaller and
more dispersed than those on the interocular surface. Lateral margin of gena, around eyes, without row of setae or
with almost inconspicuous setae. Dorsal interocular distance approximately three times eye width (Fig. 61).
Thorax. Surface of pronotum with dense foveiform punctures, not touching each other. Punctures on the sides and
posterior angles as close as those on anterior angles. Discal punctures smaller and more dispersed than those of
lateral portion, separated by approximately two or three times their diameter. Anteromedial punctures smaller and
more dispersed than discal punctures. Posteromedial punctures usually separated by at least one diameter. Margin
between anterior and lateral angles with row of small setae. Margin between lateral and posterior angles straight,
with approximately the same length of margin between anterior and lateral angles. Hypomera with dense foveiform
punctures at anterior portion and absent at posteromedial portion (Fig. 83); anteromedial punctures usually
separated by less than one diameter. Diagonal band of hypomera from posteromedial portion to middle of lateral
margin lacking punctures or with one or two isolated punctures on otherwise smooth surface (Fig. 83). Elytra.
Lateral margins slightly curved outward (Fig. 110) and surface slightly irregular (rugose). Elytral striae with
carinate margins clearly separated; strial punctures often elliptical, separated by irregular spacing (Fig. 84).
Interstriae flattened, with foveiform punctures surrounded by approximately five well-defined shiny
microtubercles. Foveiform punctures separated on elytral disc by approximately half diameter. Basal carina of
seventh interstria elongate, length about four times the width of the seventh interestria. Basal carina of ninth
interstria with approximately one-half of elytral length (Fig. 80). Apical carina or tubercle of second interstria
inconspicuous. Apical carina of third interstria weaker than apical carina of fourth interstria. Length of apical
carina of sixth interstria longer than that of apical carina of fifth interstria. Length of apical carina of seventh
interstria noticeably smaller than the sum of lengths of apical carinae of fifth and sixth interstriae. Pseudepipleural
carina incomplete, effaced medially (Fig. 80). Parameres symmetric and elongate, length equal to or longer than
phallobase; in lateral view forming an angle greater than 60° with phallobase; apex rounded (Fig. 16). Lamella
copulatrix (LC): subrectangular sclerites (Fig. 37). Fronto-lateral peripheral sclerite (FLP) as in Fig. 54.
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Material studied. Holotype (BMNH) [examined] : COLOMBIA (no locality given in original
description). 1. [white label, handwritten]. 416; 2. [white label, printed in black]. 6745; 3. [white label, bordered in
red, round]. Type; 4. [white label, handwritten]. longiceps/ Reiche/ Colombie; 5. [white label, handwritten].
Deltochilum/ valgum var. longiceps/ Paulian/ Type.
Non-type material [1]: COLOMBIA: TOLIMA: Mariquita, XI.1995, F. Escobar (1 ♂ CEMT).
Sampling methods. un-baited pitfall trap [1].
Habitat. Colombia (Fig. 91), one specimen with label data "Bosque Seco".
Diagnosis and remarks (within the acropyge complex): foveiform punctures separated at the dorsal
interocular surface by approximately one diameter (Fig. 61); foveiform punctures of pronotal disc separated by
approximately two or three times their diameter; punctures of elytral striae often elliptical, separated by irregular
spacing (Fig. 84); apical carina or tubercle of second elytral interstria inconspicuous; pseudepipleural carina
incomplete, effaced medially (Fig. 80). The pattern of hypomera punctures on individuals of this species is also
characteristic (Fig. 83). The following aedeagus characters are also diagnostic: parameres symmetric and elongate,
length equal to or longer than phallobase; in lateral view forming an angle greater than 60° with phallobase; apex
rounded (Fig. 16). Lamella copulatrix (LC): subrectangular sclerites (Fig. 37). Fronto-lateral peripheral sclerite
(FLP) as in Fig. 54.
Escobar (2003) reported the feeding behavior of seven specimens from Colombia (Tolima), supposedly
identified as D. valgum acropyge Bates. As far as we know, D. acropyge Bates inhabits only the Caribbean slope of
Central America to Mexico. From Colombia we know only specimens of D. longiceps. Therefore, we belive that
those specimens reported from Colombia are D. longiceps.
Deltochilum (Aganhyboma) acanthus Kohlmann & Solís 2012
(Figs 15, 91, 111)
Deltochilum (Deltohyboma) acanthus Kohlmann & Solís 2012: 35, 36, 37, 38, Figures 6, 7 and 15 (original description)
Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with an evident sheen. Length
< 13 mm. Head. Punctures dense, nearly touching at central portion; around clypeus shallower, smaller and more
dispersed than those on the interocular surface. Lateral margin of gena, around eyes, without row of setae or with
almost inconspicuous setae. Dorsal interocular distance approximately 2.5 times eye width. Thorax. Surface of
pronotum with dense foveiform punctures, not touching each other. Punctures on the sides, anterior and posterior
angles separated by half diameter or less. Discal punctures smaller and more dispersed than those of lateral portion,
separated by approximately two times their diameter. Anteromedial punctures smaller and more dispersed than
discal punctures. Posteromedial punctures separated by less than one diameter. Margin between anterior and lateral
angles with row of small setae. Margin between lateral and posterior angles straight, with the same length of
margin between anterior and lateral angles. Hypomera with dense foveiform punctures at anterior portion and
absent at posteromedial portion; anteromedial punctures usually separated by less than one diameter. Diagonal
band of hypomera from posteromedial portion to middle of lateral margin lacking punctures or with one or two
isolated punctures on otherwise smooth surface (as in Fig. 83). Elytra. Lateral margins semiparallel (Fig. 111) and
surface slightly irregular (rugose). Elytral striae with carinate margins clearly separated; several punctures clearly
elliptical, separated by regular spacing. Interstriae flattened, with foveiform punctures surrounded by
approximately four well-defined shiny microtubercles. Foveiform punctures separated on elytral disc by
approximately one diameter. Basal carina of seventh interstria elongate, length about four times the width of the
seventh interestria. Basal carina of ninth interstria with approximately one-half of elytral length (as in Fig. 80).
Apical carina or tubercle of second interstria present. Apical carina of third interstria weaker than apical carina of
fourth interstria. Length of apical carina of sixth interstria longer than that of apical carina of fifth interstria. Length
of apical carina of seventh interstria noticeably less than that of sum of apical carinae of fifth and sixth interstriae.
Pseudepipleural carina incomplete, effaced medially (as in Fig 80).
Material studied. Holotype (INBio) [not examined]: COSTA RICA: Prov. Puntarenas, Res. Biol. Carara,
Est. Quebrada Bonita, VI. 1993, 50 m, J.C. Saborío, L-N-194500, 469850, CRI001185073 (1 ♂ INBio collection).
The original description is sufficient to confirm the identification of species.
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Non-type material [2]: PANAMA: PANAMA: Gatún Lake, Barro Colorado Island, 5.V.1981, B. Gill (1 ♂
CMN). COSTA RICA: PUNTARENAS: Osa Peninsula. Río Piro, la Joya 350m, 8°23'56"N, 83°19'51"W,
V–IX.1997, N. Gibson (A. Forsyth) (1 ♀ AFIC).
Sampling methods. flight interception trap [1 specimen].
Habitat. Panama and Costa Rica (Fig. 91). According to Kohlmann & Solis (2012), this species inhabits
Pacific tropical rain forest of Costa Rica and the Canal Zone of Panama, ranging from 0 to 100 m AMSL.
Diagnosis and remarks (within the acropyge complex): foveiform punctures of dorsal interocular surface
nearly touching; foveiform punctures of pronotal disc separated by approximately twice their diameter;
posteromedial punctures of pronotum usually separated by less than one diameter; pronotal margin, between lateral
and posterior angles, with approximately the same length of margin between anterior and lateral angles; elytra with
lateral margins semiparallel (Fig. 111); punctures of elytral striae often elliptical, separated by regular spacing;
apical carina or tubercle of second elytral interstria present; pseudepipleural carina incomplete, effaced medially
(as in Fig 80). The following aedeagus characters are also diagnostic: parameres symmetric and elongate, length
approximately equal to phallobase; in lateral view forming an angle of approximately 45° with phallobase; apex
rounded (Fig. 15). Lamella copulatrix (LC): subrectangular sclerites. Fronto-lateral peripheral sclerite (FLP) as in
Fig. 54.
Deltochilum (Aganhyboma) arturoi new species
(Figs 82, 91, 112)
Etymology. Named in honour of Fabio Arturo González Alvarado, friend of both authors and good dung beetle
taxonomist.
Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with an evident sheen.
Length > 14 mm. Head. Punctures dense but not touching each other; foveiform punctures separated at the dorsal
interocular surface by approximately half diameter. Punctures around clypeus shallower, smaller and more
dispersed than those on the interocular surface. Lateral margin of gena, around eyes, without row of setae, or with
almost inconspicuous setae. Dorsal interocular distance approximately three times eye width (as in Fig. 61).
Thorax. Surface of pronotum with dense foveiform punctures. Punctures on the sides, anterior and posterior angles
nearly touching. Discal punctures smaller and more dispersed than those of lateral portion, separated
approximately by one diameter. Anteromedial punctures smaller and more dispersed than discal punctures.
Posteromedial punctures usually separated by at least one diameter. Margin between anterior and lateral angles
with row of small setae. Margin between lateral and posterior angles straight, with approximately the same length
of margin between anterior and lateral angles. Hypomera with dense foveiform punctures on anterior portion and
dispersed on posteromedial portion (Fig. 82); anteromedial punctures separated by approximately half diameter.
Diagonal band of hypomera from posteromedial portion to middle of lateral margin with few isolated punctures on
otherwise smooth surface (Fig. 82). Elytra. Lateral margins slightly curved outward (Fig. 112) and surface slightly
irregular (rugose). Elytral striae with carinate margins clearly separated; some punctures slightly elliptical,
separated by regular spacing. Interstriae flattened, with foveiform punctures surrounded by approximately four
well-defined shiny microtubercles. Foveiform punctures separated on elytral disc by at least one diameter. Basal
carina of seventh interstria elongate, length about four times the width of the seventh interestria. Basal carina of
ninth interstria with approximately one-half of elytral length (as in Fig. 80). Apex of second interstria with carina
or tubercle inconspicuous. Apical carina of third interstria weaker than apical carina of fourth interstria. Length of
apical carina of sixth interstria longer than that of apical carina of fifth interstria. Length of apical carina of seventh
interstria approximately equal to the sum of lengths of apical carinae of fifth and sixth interstriae. Pseudepipleural
carina incomplete, effaced medially (as in Fig. 80).
Material studied. Holotype: ECUADOR: PICHINCHA: B.P. Milpe, 1200m, IV.2003, Fumigación Dosel,
P.Araujo & William Chamorro (1 ♀ CEMT).
Paratype [1]: ECUADOR: SANTO DOMINGO: Santo Domingo (16 Km E), Tinalandia, 25.VII.1985, S. & J.
Peck (1 ♀ CMN).
Sampling methods. flight interception trap [1 specimen].
Habitat. Ecuador (Fig. 91), one specimen with label data “tropical rain forest”.
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Diagnosis and remarks (within the acropyge complex): length > 14 mm; foveiform punctures of dorsal
interocular surface separated by approximately half diameter; foveiform punctures of pronotal disc separated
approximately by one diameter; posteromedial punctures of pronotum usually separated by at least one diameter;
pronotal margin, between lateral and posterior angles, with approximately the same length of margin between
anterior and lateral angles; some punctures of elytral striae slightly elliptical, separated by regular spacing; apical
carina or tubercle of second elytral interstria inconspicuous; pseudepipleural carina incomplete, effaced medially
(as in Fig. 80). The pattern of hypomera punctures on individuals of this species is also characteristic (Fig. 82).
Deltochilum (Aganhyboma) larseni new species
(Figs 14, 91, 113)
Etymology. A patronym honoring Trond Larsen, great and enthousiatic scarab ecologist who collected most of the
type specimens.
Description. Body. Black or dark brown coloration, with obvious sheen (Fig. 113). Length < 13 mm. Head.
Punctures dense, nearly touching; around clypeus shallower, smaller and more dispersed than those on the
interocular surface. Lateral margin of gena, around eyes, without row of setae or with almost inconspicuous setae.
Dorsal interocular distance approximately 2.5 times eye width. Thorax. Surface of pronotum with dense foveiform
punctures. Punctures on the sides, anterior and posterior angles nearly touching. Discal punctures smaller and more
dispersed than those of lateral portion, separated approximately by one diameter. Anteromedial punctures smaller
and more dispersed than discal punctures. Posteromedial punctures usually separated by at least one diameter.
Margin between anterior and lateral angles with row of small setae. Margin between lateral and posterior angles
with length somewhat shorter than that of margin between anterior and lateral angles. Hypomera with dense
foveiform punctures on anterior portion and dispersed on posteromedial portion; anteromedial punctures almost
touching. Diagonal band of hypomera from posteromedial portion to middle of lateral margin with dispersed
punctures on otherwise smooth surface. Elytra. Lateral margins slightly curved outward (Fig. 113) and surface
slightly irregular (rugose). Elytral striae with carinate margins clearly separated; strial punctures separated by
regular spacing; most of punctures round intermingled with some slightly elliptical. Interstriae flattened, with
foveiform punctures surrounded by approximately five obvious shiny microtubercles. Foveiform punctures
separated on elytral disc by approximately one diameter. Basal carina of seventh interstria elongate, length about
four times the width of the seventh interestria. Basal carina of ninth interstria with approximately one-half of elytral
length (as in Fig. 80). Apical carina or tubercle of second elytral interstria absent. Apical carina of third interstria
weaker than apical carina of fourth interstria. Length of apical carina of sixth interstria longer than that of apical
carina of fifth interstria. Length of apical carina of seventh interstria longer than or equal to the sum of lengths of
apical carinae of fifth and sixth interstriae. Pseudepipleural carina incomplete, effaced medially (as in Fig. 80).
Material studied. Holotype: ECUADOR: SUCUMBIOS: R.P.F. Cuyabeno, Trocha Zábalo-Güepí, km 10,
9.VIII.2010, Colección Manual nocturna, Bosque de tierra firme colinado, Pablo Araujo, LOTE 557 (1 ♂ CEMT).
Paratypes [15]: ECUADOR: SUCUMBIOS: R.P.F. Cuyabeno, Trocha Zábalo-Güepí, km 10, 9.VIII.2010,
Colección Manual nocturna, Bosque de tierra firme colinado, Pablo Araujo, LOTE 557 (1 ♂ CEMT); PASTAZA:
B.P. Oglán, T4, Bsvpm. 554 msnm, 201146E 9853376N, 23.VIII.2008, Col. W. Chamorro & V. Guasumba (1 ♀
CEMT). PERU: valle cerca Abancay, 2.VII.1948, C.S. Carbonell (1 ♂ MZUSP); Upper Rio Marañón, 8.X.1928 (1
♂ MZUSP); MADRE DE DIOS: Río Tambopata, 300 m, Ccolpa de Guacamayos, 13°08.5'S, 69°36.4'W, A
Forsyth, X.1995 (1 ♂ AFIC); Rio Madre de Dios, Rio Los Amigos, base camp, 1° terra firme, 290 m, pitfall
crushed but live spirobolid millipede, 12°34'10.0"S, 70°06'01.4"W, T. Larsen, IV.2000 (1 ♂ CEMT, 1♂ 1♀ AFIC);
same but 12–13.IV.2000 (1♂ CEMT); same but Rio Amiguillos, small rever flood plain, 260 m, pitfall millipede,
12°22'25.4"S, 70°22'13.2"W, T. Larsen, V.2000 (1♂ AFIC, 1♀ CEMT); same but Rio Palma Real Grande, Limon
Camp, 12°32'20"S, 68°51'40"W, Flight intercept trap, 220 m, T. Larsen, 11–12.X.1999 (1♀ CEMT); same but pitf.
crished millipede, 12–13.X.1999 (1♀ AFIC); same but perching night, 11.X.1999 (1♀ AFIC); same but
12°32'20"S, 68°51'41"W, Flight interception trap, 400 m, 2–3.IV.1999 (1♀ AFIC).
Habitat. Ecuador and Peru (Fig. 91), from available data this species inhabits Amazon rainforest.
Diagnosis and remarks (within the acropyge complex): length < 13 mm; punctures of pronotal disc separated
by approximately one diameter; posteromedial punctures of pronotum usually separated by at least one diameter;
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pronotal margin, between lateral and posterior angles, with length somewhat shorter than margin between anterior
and lateral angles; elytral striae with carinate margins clearly separated; strial punctures separated by regular
spacing; most of strial punctures round intermingled with some slightly elliptical; apical carina or tubercle of
second interstria absent; pseudepipleural carina incomplete, effaced medially (as in Fig. 80). The following
aedeagus characters are also diagnostic: parameres symmetric and short, length shorter than phallobase; in lateral
view forming an angle of approximately 45° with phallobase (Fig. 14). Lamella copulatrix (LC): subrectangular
sclerites. Fronto-lateral peripheral sclerite (FLP) as in Fig. 54.
finestriatum complex
Diagnosis. Elytral striae narrow, carinate margins close and almost touching (Fig. 74).
Deltochilum (Aganhyboma) cangalha new species
(Figs 10, 76, 78, 92, 114)
Etymology. The new species name, "cangalha", is a typical term of the northeastern region of Brazil, which means
"bowlegged", a characteristic usually found in individuals of Aganhyboma. The epithet is a name in apposition.
Description. Body. Head, pronotum, elytra, and ventral surface black, with a copper-red sheen. Length > 14
mm. Head. Punctures extremely dense and touching; around clypeus shallower, smaller and more dispersed than
those on the interocular surface. Lateral margin of gena, around eyes, without row of setae, or with almost
inconspicuous setae. Dorsal interocular distance approximately three times eye width (as in Fig 61). Thorax.
Pronotum with a red metallic sheen (Fig. 114). Surface of pronotum with dense foveiform punctures, nearly
touching on the sides, anterior angles and posterior portion. Discal punctures smaller and more dispersed than those
of lateral portion. Anteromedial punctures smaller and more dispersed than discal punctures. Posteromedial
punctures usually separated by less than one diameter. Margin between anterior and lateral angles with row of
small setae. Margin between lateral and posterior angles approximately straight. Hypomera with moderate
foveiform punctures on anterior and posterior portions (Fig. 78); anteromedial punctures separated by at least one
diameter (Fig. 78). Elytra. Elytra rounded, sides bulging outward (Fig. 114) and surface slightly irregular (rugose).
Elytral striae narrow, carinate margins close and almost touching (as in Figs 74, 76). Interstriae flattened, with
foveiform punctures surrounded by shiny areas poorly defined. Foveiform punctures of interstriae separated on
elytral disc by less than one diameter. Microtubercles of interstriae dense and weakly defined on most of surface,
united by glossy spots with a red-cupreous sheen (Fig. 76). Basal carina of seventh interstria short, length about
twice the width of the seventh interestria. Basal carina of ninth interstria present in three-quarters of elytra, but it
never reaches the anterior limit of apical carina of seventh interstria. Apical carina of third interstria somewhat
more prominent than apical carinae of second and fourth interstriae. Length of apical carina of sixth interstria
longer than that of apical carina of fifth interstria. Length of apical carina of seventh interstria less than the sum of
lengths of apical carinae of fifth and sixth interstriae. Pseudepipleural carina complete, visible entire length of
epipleuron (as in Fig. 63).
Material studied. Holotype: BRAZIL: MINAS GERAIS: Águas Vermelhas, XII.1998, A. Bello & F. Z. Vaz-
de-Mello (1 ♂ CEMT).
Habitat. Brazil (Minas Gerais) (Fig. 92), from available data this species inhabits “Carrasco”. This habitat
consists of dense shrubby xerophilous vegetation that occurs in semi-arid domain of Brazil.
Diagnosis and remarks (within the finestriatum complex): pronotal margin, between lateral and posterior
angles, straight; Elytra rounded, sides bulging outward (Fig. 114); anteromedial punctures of hypomera separated
by at least one diameter (Fig. 78); pronotum with a red metallic sheen; elytral microtubercles weakly defined, with
a red-cupreous sheen (Fig. 76). The following aedeagus characters are also diagnostic: parameres symmetric and
elongate, in lateral view forming an angle of approximately 45° with phallobase; apex rounded (Fig. 10). Lamella
copulatrix (LC): left lobe subtrapezoidal; right lobe subrectangular. Fronto-lateral peripheral sclerite (FLP) as in
Fig. 53.
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Although the aedeagus of D. cangalha new species and D. alpercata new species do not present significant
difference, the elytral punctures in these species are distinct.
Deltochilum (Aganhyboma) alpercata new species
(Figs 10, 77, 79, 92, 115)
Etymology. “alpercata" is a typical footwear of the northeastern region of Brazil. The epithet is a name in
apposition.
Description. Body. Head, pronotum, elytra, and ventral surface black, with a silky sheen. Length > 14 mm.
Head. Punctures extremely dense and touching; around clypeus shallower, smaller and more dispersed than those
on the interocular surface. Lateral margin of gena, around eyes, without row of setae, or with almost inconspicuous
setae. Dorsal interocular distance approximately three times eye width (as in Fig. 61). Thorax. Pronotum black
with an opaque sheen or a weak luster (Fig. 115). Surface of pronotum with dense foveiform punctures, nearly
touching on the sides, anterior angles and posterior portion. Discal punctures somewhat smaller and more dispersed
than those of lateral portion. Anteromedial punctures smaller and more dispersed than discal punctures.
Posteromedial punctures usually separated by less than one diameter. Margin between anterior and lateral angles
with row of small setae. Margin between lateral and posterior angles approximately straight. Hypomera with dense
foveiform punctures on anterior and posterior portions (Fig. 79); anteromedial punctures separated by less than one
diameter (Fig. 79). Elytra. Rounded, sides bulging outward (Fig. 115) and surface slightly irregular (rugose).
Elytral striae narrow, carinate margins close and almost touching (as in Figs 74, 77). Interstriae flattened, with
foveiform punctures surrounded by four or five shiny microtubercles. Foveiform punctures separated on elytral
disc by less than one diameter. Microtubercles of interstriae well delimited and conspicuous, with strong luster;
interestrial surface around microtubercles opaque (Fig. 77). Basal carina of seventh interstria short, length about
twice the width of the seventh interestria. Basal carina of ninth interstria present in three-quarters of elytra, but it
never reaches the anterior limit of apical carina of seventh interstria. Apical carina of third interstria more
prominent than apical carinae of second and fourth interstriae. Length of apical carina of sixth interstria longer than
that of apical carina of fifth interstria. Length of apical carina of seventh interstria less than or equal to the sum of
lengths of apical carinae of fifth and sixth interstriae. Pseudepipleural carina complete, visible entire length of
epipleuron (as in Fig. 63).
Material studied. Holotype: BRAZIL: ALAGOAS: Murici, S. Branca, V.1984, F.M. Oliveira (1 ♂ CEMT).
Habitat. Brazil (Alagoas) (Fig. 92), from available data this species inhabits lowland Atlantic forest.
Diagnosis and remarks (within the finestriatum complex): elytra rounded, sides bulging outward (Fig. 115);
punctures of anteromedial portion of hypomera separated by less than one diameter (Fig. 79); pronotum black with
an opaque sheen or a weak luster (Fig. 115); elytral microtubercles well delimited, with a brown sheen (Fig. 77);
interestrial surface around microtubercles opaque (Fig. 77). The following aedeagus characters are also diagnostic:
parameres symmetric and elongate, in lateral view forming an angle of approximately 45° with phallobase; apex
rounded (Fig. 10). Lamella copulatrix (LC): left lobe subtrapezoidal; right lobe subrectangular. Fronto-lateral
peripheral sclerite (FLP) as in Fig. 53.
Although the aedeagus of D. alpercata new species and D. cangalha new species do not present significant
difference, the elytral punctures in these species are distinct.
Deltochilum (Aganhyboma) finestriatum new species
(Figs 11, 28, 36, 45, 53, 63, 74, 92, 116)
Etymology. Named referring to the narrow elytral striae.
Description. Body. Head, pronotum, elytra, and ventral surface black, with a silky sheen. Length 13.3 to 15.8
mm. Head. Punctures extremely dense and touching. Foveiform punctures distributed along the entire dorsal
surface; around clypeus shallower, smaller and more dispersed than those on the interocular surface. Lateral margin
of gena, around eyes, without row of setae or with almost inconspicuous setae. Dorsal interocular distance
approximately three times eye width (as in Fig. 61). Thorax. Surface of pronotum with dense foveiform punctures,
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nearly touching on the sides, anterior angles and posterior portion. Discal punctures somewhat smaller and more
dispersed than those of lateral portion. Anteromedial punctures smaller and more dispersed than discal punctures.
Posteromedial punctures usually separated by less than one diameter. Margin between anterior and lateral angles
with row of small setae. Margin between lateral and posterior angles approximately straight. Hypomera with dense
foveiform punctures on anterior portion; anteromedial punctures separated by less than one diameter. Elytra. Oval-
elongate with lateral margins slightly curved outward (Fig. 116) and surface slightly irregular (rugose). Elytral
striae narrow, carinate margins close and almost touching (Fig. 74). Interstriae flattened, with foveiform punctures
surrounded by glossy areas weakly defined (Fig. 74). Foveiform punctures separated on elytral disc for less than
one diameter. Microtubercles dense and weakly defined on most of surface, united by depressed glossy areas
making their delimitation difficult (Fig. 74). Basal carina of seventh interstria elongate, length about four times the
width of the seventh interestria. Basal carina of ninth interstria present in three-quarters of elytra, but it never
reaches the anterior limit of apical carina of seventh interstria (Fig. 63). Apical carina of third interstria elongate,
more prominent than apical carinae of second and fourth interstriae. Length of apical carina of sixth interstria
longer than that of apical carina of fifth interstria. Length of apical carina of seventh interstria less than the sum of
lengths of apical carinae of fifth and sixth interstriae. Pseudepipleural carina complete, visible entire length of
epipleuron (Fig. 63).
Material studied. Holotype: BRAZIL: MINAS GERAIS: Viçosa, Mata do Paraíso, (20º48.135'S,
42º51.525'W), 14.II.2009, Fernando Silva (1 ♂ CEMT).
Paratypes [27]: BRAZIL: [no data] (1 ♀ MNHN); ESPÍRITO SANTO: Rio Bonito, II.1964 (1 ♀ DZUP);
MINAS GERAIS: Guanhães, 25.III.1994, J.C. Zamuncio (1 ♀ CEMT); Marliéria, Parque Estadual do Rio Doce,
(19°42'09.22"S, 42°30'54.21"W), 11.I.2005, S.S.P. Almeida & M.R. Pereira (1 ♂ CEMT); Ravena,
(19°49'46.52"S, 43°49'01.24"W), (1 ♂ CEMT); Viçosa, Mata do Paraíso, X.1998, F.Z. Vaz-de-Mello (1 ♀ CEMT);
same, XI.1998, F.Z. Vaz-de-Mello (1 ♀ CEMT); same, (20º48.135'S, 42º51.525'W), 14.II.2009, Fernando Silva (6
4 ♀ CREN); PARANÁ: Campina Grande do Sul, Estrada da Mandaçaia, I.2009, F.W.T. Freitas (1 ♀ CEMT);
Cornélio Procópio, Parque Estadual Mata São Franscisco, (23°09'20”S, 50°34'20”W), 14.XII.2009, N. Cipola (1 ♂
CEMT); Londrina, Parque Estadual Mata dos Godoy, (23°27'S, 51°14'W), 24.I.2000, J. Lopes (1 ♀ CEMT); RIO
DE JANEIRO: Rio de Janeiro, Jardim Botânico, XII.1992, F.Z. Vaz-de Mello (1 ♀ CEMT); SANTA CATARINA:
Corupá, XII.1953, Anton Maller (1 ♀ MZUSP); Corupá, 1961, Anton Maller (1 ♀ MZUSP); same, XII.1963, (1 ♀
DZUP); Florianópolis, Lagoa do Peri, (27°49'S, 48°32'W), X.2007, P. Condé & M. Hernández (1 ♀ CEMT);
Hansa, I.1941, (1 ♂ MZUSP); SÃO PAULO: Santo André, Rebio Alto da Serra do Paranapiacaba, (23°46'48"S,
46°18'36"W), 18.II.2007, M. Uehara-Prado (1 ♀ CEMT).
Sampling methods. baited pitfall trap with injured diplopods [11 specimens]; flight interception trap [2].
Habitat. Brazil (Espírito Santo, Minas Gerais, Paraná, Rio de Janeiro and Santa Catarina) (Fig. 92). From the
available data this species inhabits Atlantic forest, ranging from 0 to 800 m AMSL.
Diagnosis and remarks (within the finestriatum complex): elytra with oval-elongate shape and lateral margins
slightly curved outward (Fig. 116); elytral microtubercles dense and poorly defined on most of surface, united by
glossy spots with dark brown or black sheen, never copper-red (Fig. 74). The following aedeagus characters are
also diagnostic: parameres symmetric and elongate; in lateral view forming an angle of approximately 45° with
phallobase; apex rounded (Fig. 11). Genital segment as in Fig. 28. Lamella copulatrix (LC): left and right lobes as
in Fig. 36. Superior right peripheral sclerite (SRP) as in Fig. 45. Fronto-lateral peripheral sclerite (FLP) as in Fig.
53.
valgum complex
Diagnosis. Interstriae convex.
Deltochilum (Aganhyboma) icaroides Balthasar 1939 new subgeneric placement
(Figs 18, 31, 39, 48, 85, 87, 93, 117)
Deltochilum (s. str.) icaroides Balthasar 1939: 10 (original description)
Deltochilum icaroides: Blackwelder 1944: 203 (checklist)
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Roze 1955: 43 (checklist)
Génier 2001: 4 (comments about type locality)
Deltochilum (s. str.) icarioides: Lane 1947: 109 (taxonomic comments)
Deltochilum (Deltohyboma) icaroides: Martínez 1959: 54 (catalogue)
Vulcano & Pereira 1964: 655 (catalogue)
Vulcano & Pereira 1967: 557 (identification key)
Vaz-de-Mello 2000: 192 (checklist)
Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with an opaque or silky sheen.
Length 12.8 to 13.5 mm. Head. Foveiform punctures extremely dense, usually touching on the dorsal interocular
surface. Punctures around clypeus shallower, smaller and more dispersed than those on the interocular surface.
Lateral margin of gena, around eyes, without row of setae or with almost inconspicuous setae. Dorsal interocular
distance approximately four times eye width. Thorax. Surface of pronotum with dense foveiform punctures, nearly
touching on the sides, anterior and posterior angles (Fig. 85). Discal punctures somewhat smaller and more
dispersed than those of lateral portion. Anteromedial punctures smaller and more dispersed than discal punctures.
Posteromedial punctures usually separated by less than one diameter. Margin between anterior and lateral angles
with row of small setae. Hypomera with dense foveiform punctures on anterior and posterior portions; foveiform
punctures almost touching. Elytra. Lateral margins slightly curved outward (Fig. 117) and surface slightly
irregular (rugose). Elytral striae wider, carinate margins clearly separated (as in Fig. 75). Interstriae slightly
convex; however, always less convex than that of the icariforme subcomplex. Interstriae with foveiform punctures
over entire surface. Elytral microtubercles almost inconspicuous, sometimes blending in with remainder of
interstria surface. Basal carina of seventh interstria elongate, length about four times the width of the seventh
interestria. Basal carina of ninth interstria present in three-quarters of elytra, but it never reaches the anterior limit
of apical carina of seventh interstria. Apical carina of third interstria somewhat more prominent than apical carinae
of second and fourth interstriae (Fig. 87). Length of apical carina of sixth interstria longer than that of apical carina
of fifth interstria. Length of apical carina of seventh interstria less than or equal to the sum of lengths of apical
carinae of fifth and sixth interstriae. Pseudepipleural carina complete, visible entire length of epipleuron (as in Fig.
63).
Material studied. Holotype (NMPC) [examined] : 1. [white label, printed in black]. San Bernardino / A.
Fischer S. G. / 8. 1912; 2. [white label, handwritten and printed in black]. Deltochilum / icariformis / Ohs. in litt. /
dr. V. Balthasar det.; 3. [red label, handwritten]. Typus; 4. [red label, printed in black]. HOLOTYPE; 5. [white
label, printed in black]. Type; 6. [green label, bordered in black, handwritten]. icaroides / m. See also Bezděk &
Hájek (2011).
The holotype has the following locality data on the label: «San Bernardino, A. Fischer, S. G., 8.1912». In the
original description, the author supposes that this location is in Venezuela (Balthasar 1939). However, Génier
(2001) believes that the specimen is from San Bernardino, Paraguay, because he examined an individual, according
to him, identical to the type, from Villarica, Paraguay.
Non-type material [44]: ARGENTINA: MISIONES: Loreto (1 ♀ MZUSP); Loreto, A. Uglobin (1 ♀
CEMT); P. N. Iguazú, 6.I.1991, S. & J. Peck (1 ♀ CMN); Puerto Iguazú, X.1997 (1 ♀ CEMT). BOLIVIA: SANTA
CRUZ: Chiquito, Santiago de Chiquito, (18º19´S, 59º34´W), 11.XI.2008, T. Vidaurre (1 ♂ 2 ♀ MNKM, 3 ♂ 1 ♀
CEMT); same, (18°20.103'S, 59°35.007'W), XI.2008, W.D.E. Edmonds & T. Vidaurre (7 ♂ 2 ♀ WDEC); 5km SSE
Buena Vista, Hotel Flora e Fauna, (17°29.925'S, 63°39.128'W), 15.XII.2003, S. & J. Peck (1 ♂ CMN). BRAZIL:
GOIÁS: [anonym] (2 ♀ MZUSP); MINAS GERAIS: Rio Doce, 5.III.1940 (1 ♀ MZUSP); MATO GROSSO:
Chapada dos Guimarães, 14.XI.2008, S.P. Rosa, F. Fernandes, J. Almeida & R. Kawada (2 ♀ CEMT); Diamantino,
Alto Rio Arinos, X.1999, E. Furtado (1 ♂ CEMT); same, II.2001, E. Furtado (1 ♂ CEMT); Nossa Senhora do
Livramento, Distrito de Pirizal, 15.X.2009, D. D. Pinheiro (1 ♀ CEMT); Rosário Oeste, XI.1959 (2 ♀ DZUP);
same, XI.1963, Alvarenga & Werner (1 ♀ MZUSP); same, II.1969 (1 ♀ DZUP); MATO GROSSO DO SUL:
Bodoquena, Ass. Canaã, (20°46.267'S, 56°46.038'W), XI.2009, F. O. Roque (1 ♂ 1 ♀ CEMT); Corumbá, Serra do
Urucum, (19°13'S, 57°33'W), 23.XI.1960, K. Lenko (1 ♀ DZUP); SÃO PAULO: Batatais, XII.1940 (1 ♀
MZUSP); Pres. Epitácio, J. Lane (1 ♂ MZUSP). PAR AGUAY: CANINDEYÚ: Lagunita, Res. Nat. Bosque
Mbaracayú, 10.XI.1996, C. F. Costa (1 ♀ CMN); Salto del Guaira, Melgarejo, 10.X.1994, Drechsel (1 ♂ CEMT);
GUAIRA: Villarrica, XI.1928, F. Schade (2 ♀ CEMT); same, 23.XII.1946, (1 ♀ MZUSP); Cerro Acati, XI.1993,
Drechsel (1 ♀ CEMT).
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Sampling methods. baited pitfall trap with injured diplopods [16 specimens].
Habitat. Argentina, Bolivia, Brazil (Goiás, Minas Gerais, Mato Grosso, Mato Grosso do Sul and São Paulo)
and Paraguay (Fig. 93). From available data this species inhabits mainly the Cerrado domain. The elevation data
ranges from 622 to 750 m AMSL.
Diagnosis and remarks (within the valgum complex): length 12.8 to 13.5 mm; foveiform punctures near the
posteromedial portion of pronotum usually separated by less than one diameter; elytral interstriae with
homogeneous and dense foveiform punctures over entire surface; elytral surface with an opaque or silky sheen;
apical carina of third interstria somewhat more prominent than apical carinae of second and fourth interstriae (Fig.
87). The following aedeagus characters are also diagnostic: parameres symmetric and elongate; in lateral view
forming an angle of approximately 45° with phallobase; apex rounded (Fig. 18). Genital segment as in Fig. 31.
Lamella copulatrix (LC): left and right lobes as in Fig. 39. Superior right peripheral sclerite (SRP) as in Fig. 48.
Fronto-lateral peripheral sclerite (FLP) as in Fig. 55.
Some specimens exhibit apical carina of sixth interstria shorter than carina of fifth interstria.
Deltochilum (Aganhyboma) valgum Burmeister 1873 new subgeneric placement
(Figs 17, 30, 38, 47, 55, 75, 88, 93, 118)
Deltochilum valgum Burmeister 1873: 409 (original description)
Gillet 1911: 36 (catalogue)
Bruch 1911: 186 (catalogue)
Blackwelder 1944: 185 (checklist)
Larsen et al. 2009: 1–4 (feeding behavior, "misidentification" probably refers to the species described here as D. larseni)
Deltochilum (Deltochilum) var. valgum: Paulian 1938: 252, 269, 271, 276, map n. 5 (description, taxonomic comments and
identification key)
Deltochilum (Deltohyboma) valgum: Martínez 1959: 55 (catalogue)
Vulcano & Pereira 1964: 659 (catalogue)
Martínez 1987: 55 (checklist)
Deltochilum valgum valgum: Cano 1998: 177 (ecological comments)
Deltochilum (Deltohyboma) valgum valgum: Vaz-de-Mello 2000: 192 (checklist)
Description. Body. Head, pronotum, elytra and ventral surface black or dark brown, with an opaque or silky sheen.
Length 14 to 15 mm. Head. Foveiform punctures dense, but not touching each other as in D. icaroides; around
clypeus quite shallower and smaller than those on the dorsal interocular surface. Lateral margin of gena, around
eyes, without row of setae or with almost inconspicuous setae. Interocular distance approximately four times eye
width. Thorax. Surface of pronotum with deep and dense foveiform punctures, nearly touching on the sides and
posterior angles. Discal punctures somewhat smaller and more dispersed than those of lateral portion.
Anteromedial punctures smaller and more dispersed than discal punctures. Posteromedial punctures usually
separated by at least one diameter. Margin between anterior and lateral angles with row of small setae. Margin
between lateral and posterior angles straight. Hypomera with dense foveiform punctures on anterior and posterior
portions; foveiform punctures almost touching. Elytra. Lateral margins slightly curved outward (Fig. 118), or
almost parallel, and surface slightly irregular (rugose). Elytral striae wider, carinate margins clearly separated (Fig.
75). Interstriae slightly convex; however, always less convex than that of the icariforme subcomplex. Interstriae
with foveiform punctures over entire surface (Fig. 75). Microtubercles of interstriae dense and conspicuous (Fig.
75). Basal carina of seventh interstria elongate, length about four times the width of the seventh interestria. Basal
carina of ninth interstria present in three-quarters of elytra, but it never reaches the anterior limit of apical carina of
seventh interstria. Apical carina of third interstria round, better defined as tubercle, distinctly more prominent than
the apical carina of second and fourth interstriae (Fig. 88). Length of apical carina of sixth interstria longer than
that of apical carina of fifth interstria. Length of apical carina of seventh interstria less than or equal to the sum of
lengths of apical carinae of fifth and sixth interstriae. Pseudepipleural carina complete, visible entire length of
epipleuron (as in Fig. 63).
Material studied. Holotype (MACN) [image examined]: 1. [white label, bordered and printed in black].
MACN-En/ 1525; 2. [red label, bordered in black, handwritten]. Deltochilum/ valgum/ Burmeister/ 1873/ Typus; 3.
[white label, bordered in black, handwritten and printed]. Deltochilum/ valgum valgum/ Lectotipo Burm/ A.
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Martínez Det. 1950; 4. [old white label, handwritten]. valgum/ Burm./ Rep. Arg.; 5. [white label, bordered in red,
handwritten]. 52977; 6. [old white label, printed in black]. Col./ Antigua; 7. [green label, printed in black].
Tucuman.
The locality data in original description was given as “near Tucumán”. Fabio Arturo González supplied high
resolution images of a specimen from Tucumán, deposited at the MACN, with a lectotype label placed by Antonio
Martínez. However, Martínez did not publish this designation. After studying the original description of
Burmeister, we belive that the author described this species from a single specimen. Therefore, the specimen with
the lectotype designation by Martínez is the holotype.
Non-type material [10]: ARGENTINA: CORDOBA: El Sauce, Diquecito, Martínez (1 ♀ CEMT); same,
XII.1964, Martínez (1 ♂ CMN); FORMOSA: San José, 20.XI.1948, E. Budim (1 ♂ CEMT); JUJUY: Km 38/Ruta
52, 17.I.2008, K.S. Ramos (1 ♂ CEMT); P. N. Calilegua (3 km NW de campground), (23°44.149'S,
064°51.044'W), 15.I.2008, D.M. Takiya (1 ♀ CEMT); P. N. Calilegua (Estaca el Cero), 28.XII.1987, S. & J. Peck
(1 ♀ CMN). BOLIVIA: SANTA CRUZ: Caballero, ~9Km Sudoeste Comarapa, (17°57.847'S, 64°27.745'W),
XII.2008, W.D.E. Edmonds & T. Vidaurre (1 ♂ WDEC); Chiquito, Santiago de Chiquito, (18º19'S, 59º34'W),
11.XI.2008, T. Vidaurre (1 ♂ 1 ♀ CEMT). PARAGUAY: PRESIDENTE HAYES: Pozo Colorado, Trans-Chaco
[highway], IV.1995, J.C. D'Alessandro (1 ♂ CEMT).
Sampling methods. at light-trap [3 specimens]; baited pitfall trap with injured diplopods [2].
Habitat. Argentina, Bolivia and Paraguay (Fig. 93). From available data this species inhabits mainly the
Chaco domain. Two specimens with label data "Chiquitano" [a transition zone between the moist Amazonian
forests of the north and dryer forest of the southern Chaco regions].
Diagnosis and remarks (within the valgum complex): length 14 to 15 mm; elytral interstriae with
homogeneous and dense foveiform punctures over entire surface; elytral surface with an opaque or silky sheen;
apical carina of third interstria distinctly more prominent and rounded than apical carinae of second and fourth
interstriae (Fig. 88). The following aedeagus characters are also diagnostic: parameres symmetric and short; in
lateral view forming an angle of approximately 30° with phallobase; apex rounded (Fig. 17). Genital segment as in
Fig. 30. Lamella copulatrix (LC): left lobe subrectangular (Fig. 38); right lobe "b"-shaped (Fig. 38). Superior right
peripheral sclerite (SRP) as in Fig. 47. Fronto-lateral peripheral sclerite (FLP) as in Fig. 55.
The specimens present the foveiform punctures of head and pronotum more sparse than that in D. icaroides.
icariforme subcomplex
Diagnosis. Body coloration copper with a metallic sheen (Figs 119–123); punctures of pronotal disc clearly more
dispersed than lateral punctures; elytral interstriae clearly convex; foveiform punctures of third interstria denser
laterally (punctures can be so scattered at center of interstria to produce smooth midlongitudinal strip) (Fig. 86).
Deltochilum (Aganhyboma) ritamourae new species
(Figs 22, 58, 94, 119)
Etymology. Named in honour of Rita de Cássia de Moura, an enthusiast in the study of dung beetles and specialist
in Scarabaeinae cytogenetics.
Description. Body. Head, pronotum, elytra and ventral surface with copper coloration and a reddish metallic
sheen. Length 13.5 to 14.5 mm. Head. Foveiform punctures dense, touching on dorsal interocular surface.
Punctures around clypeus shallower, smaller and more dispersed than those on interocular surface. Dorsal
interocular distance approximately four times eye width. Lateral margin of gena strongly curved outward, with row
of medium sized setae around the eyes. Thorax. Surface of pronotum with dense foveiform punctures, touching on
the sides and anterior angles (however not as deep as in D. icariforme). Density, size and proximity of anteromedial
punctures similar to those on disk. Posteromedial punctures separated by at least one diameter. Margin between
lateral and posterior angles straight. Elytra. Entirely copper and with a reddish metallic sheen (Fig. 119). Lateral
margin distinctly curved outward (Fig. 119) and surface slightly irregular (rugose). Elytral striae wider, carinate
margins clearly separated (as in Fig. 75). Interstriae clearly convex, with its central portion nearly flat. Interstrial
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surface with foveiform punctures surrounded by approximately six microtubercles. Interstrial surface usually more
punctate on sides, close to striae. Foveiform punctures of third interstria denser laterally (punctures can be so
scattered at center of interstria to produce smooth midlongitudinal strip) (as in Fig. 86). Basal carina of seventh
interstria elongate, length about three or four times seventh interestria width. Basal carina of ninth interstria
extending 3/4 or more along elytral length. Apical carina of third interstria rounded, better defined as tubercle,
more prominent than apical carinae of second and fourth interstriae. Length of apical carina of sixth interstria
longer than that of apical carina of fifth interstria. Length of apical carina of seventh interstria less than or equal the
sum of lengths of apical carinae of fifth and sixth interstriae. Pseudepipleural carina complete, visible entire length
of epipleuron (as in Fig. 63).
Material studied. Holotype: BRAZIL: MINAS GERAIS: Águas Vermelhas, XII.1998, A. Bello & F. Z. Vaz-
de-Mello (1 ♂ CEMT).
Paratypes [26]: BRAZIL: BAHIA: [no data] (1 MNHN); Encruzilhada, XI.1972, M. Alvarenga (1 ♀
CEMT); same, XII.1980, A. Martínez & M. Alvarenga (1 ♀ CMN); same, XII.1995, P. Arnaud (12 3 ♀ CEMT);
CEARÁ: Carquejo, Dirings (1 ♂ MZUSP); MINAS GERAIS: Águas Vermelhas, XII.1997, F.Z. Vaz-de-Mello (1
♂ CEMT); same, Faz. Faceiro, 15°23'56"S, 41°23'57"W, 12.XII.2012, JARafael & EJGrossi, Ar. luz. 850 m (2
INPA); Montes Claros, I.2002, G.L.D. Leite (1 ♂ CEMT); Fronteira Minas/Bahia, 14.II.1977, Celso Jr. (1 ♂
CEMT); PIAUÍ: São Raimundo Nonato, Parque Nacional Serra da Capivara, 4.XII.1998, C.A.R. Matrangolo (1 ♀
CEMT); SERGIPE: Poço Redondo, MONA Grota do Angico, (9°41'S, 38°31'W), 28–30.IV.2013, Maual, Santos-
Júnior (1 CEMT).
Sampling methods. at light [8 specimens].
Habitat. Brazil (Bahia, Ceará, Minas Gerais and Piauí) (Fig. 94). From the available data this species occurs
mainly in semi-arid domain of Brazil.
Diagnosis and remarks (within the icariforme subcomplex): length 13.5 to 14.5 mm; body completely cooper
and with a reddish metallic sheen (Fig. 119); length of apical carina of seventh interstria less than or equal the sum
of lengths of apical carinae of fifth and sixth interstriae; elytral lateral margin distinctly curved outward. The
following aedeagus characters are also diagnostic: parameres symmetric and elongate, in lateral view forming an
angle of approximately 45° with phallobase; apex acuminate (Fig. 22). Fronto-lateral peripheral sclerite (FLP) as in
Fig. 58.
In some specimens, apical carina of third interstria weaker than apical carina of second and fourth interstriae.
Deltochilum (Aganhyboma) icariforme Paulian 1938 new subgeneric placement
(Figs 21, 32, 40, 57, 94, 120)
Deltochilum (Deltochilum) icariforme Paulian 1938: 269, 273, 276, Fig. 18, map n.5 (original description, taxonomic
comments and identification key)
Deltochilum icariforme: Blackwelder 1944: 203 (checklist)
Deltochilum (Deltochilum) icariniforme: Martínez 1947: 274 (comments about geographical distribution)
Deltochilum (Deltohyboma) icariforme: Martínez 1959: 53 (catalogue)
Vulcano & Pereira 1964: 655 (catalogue); Vaz-de-Mello 2000: 192 (checklist)
Description. Body. Head, pronotum, elytra and ventral surface with copper coloration and a reddish metallic
sheen. Length 14.5 to 15.8 mm. Head. Foveiform punctures dense and deep, touching on dorsal interocular
surface. Dorsal interocular distance approximately three times eye width. Lateral margin of gena strongly curved
outward, with row of setae around eyes. Thorax. Surface of pronotum with deep and dense foveiform punctures,
nearly touching on the sides and anterior angles. Anteromedial punctures more dispersed than those of surrounding
surface. Margin between lateral and posterior angles straight. Posteromedial punctures separated by less than one
diameter. Elytra. Lateral margins slightly curved outward, almost parallel (Fig. 120). Surface slightly irregular
(rugose). Elytral striae wider, carinate margins clearly separated (as in Fig. 75). Interstriae clearly convex, with its
central portion nearly flat. Interstrial surface with foveiform punctures surrounded by microtubercles weakly
defined. Interstriae surface usually more punctate on sides, close to striae. Foveiform punctures of third interstria
denser laterally (punctures can be so scattered at center of interstria to produce smooth midlongitudinal strip) (as in
Fig. 86). Basal carina of seventh interstria elongate, length about four or more times seventh interestria width.
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Basal carina of ninth interstria present in three-quarters of elytra (not surpassing anterior limit of apical carina of
seventh interstria). Apical carina of third interstria weaker than apical carinae of second and fourth interstriae.
Length of apical carina of sixth interstria longer than that of apical carina of fifth interstria. Length of apical carina
of seventh interstria longer than the sum of lengths of apical carina of fifth and sixth interstriae. Pseudepipleural
carina complete, visible entire length of epipleuron (as in Fig. 63).
Material studied. Lectotype (MNHN) [here designated]: 1. [old white label, bordered and printed in black].
Jatahy/ Etat de Goyaz/ Ch. Pujol 1895–96; 2. [old white label, handwritten]. icariforme n. sp. Type; 3. [old white
label, bordered and printed in black]. R. PAULIAN/ Vidit; 4. [red label, bordered and printed in black].
LECTOTYPE; 5. [white label, handwritten, bordered and printed in black]. Deltochilum icariforme Paul. LECTOT
FVaz-de-Mello des. 2013.
In the original description, Paulian (1938) stated that the “type” was deposited in the Oberthür collection. After
examining the list of the specimens used by the author to describe the species, we found that only one individual
from Jataí (Goiás) and one from Lacerda (Bahia) were deposited in this collection.
We examined three of the five specimens listed in the original description. These were labeled by Paulian with
the designation of "type" and deposited in the Muséum National d'Histoire Naturelle (MNHN), Paris, France. Two
of them correspond to the specimens from Jataí and Lacerda mentioned above, and the third is from Paraguay.
After careful analysis, we concluded that these individuals represent three different species.
Thus, with the purpose of stabilizing nomenclature by choosing the name-bearing specimen, we designate the
specimen from Jataí (Goiás) as the lectotype of D. icariforme. The specimen from Bahia represents a new species
described here as D. ritamourae new species, and the specimen from Paraguay is D. icaroides.
Non-type material [5]: BRAZIL: MATO GROSSO: Rio Verde, XI.1963 (1 ♂ DZUP); SÃO PAULO:
Itirapina, 27.II.2000, G. Machado (1 ♀ CEMT); Itirapina, Estação Ecológica, (22°13'S, 47°54'W), 30.I.2009,
Fernando Silva (2 ♂ CEMT); Pirassununga, E.E.C.P., 29.X.1945, Schubart (1 ♂ MZUSP).
Sampling methods. baited pitfall trap with injured diplopods [2 specimens].
Habitat. Brazil (Goiás, Mato Grosso and São Paulo) (Fig. 94). From available data this species inhabits
Cerrado. Two specimens with label data "Campo Sujo", Cerrado.
Diagnosis and remarks (within the icariforme subcomplex): length 14.5 to 15.8 mm; length of apical carina
of seventh interstria longer than the sum of lengths of apical carina of fifth and sixth interstriae; basal carina of
ninth interstria present in three-quarters of elytra (not surpassing anterior limit of apical carina of seventh
interstria). The following aedeagus characters are also diagnostic: parameres symmetric and elongate, in lateral
view forming an angle of approximately 45° with phallobase; apex acuminate (Fig. 21). Genital segment as in Fig.
32. Lamella copulatrix (LC): left lobe trapezoid (Fig. 40); right lobe subrectangular (Fig. 40). Fronto-lateral
peripheral sclerite (FLP) as in Fig. 57.
The body coloration of some specimens can vary between dark brown, copper or red, with reddish or greenish
sheen. In some specimens, apical carina of third interstria can be somewhat more prominent than apical carina of
second and fourth interstriae, with a rounded form, resembling a tubercle.
Deltochilum (Aganhyboma) paresi new species
(Figs 3, 20, 94, 121)
Etymology. Named in honour of the indigenous people of the “Chapada dos Parecis”, known as the “Paresi”, from
region where the specimens were collected.
Description. Body. Head, pronotum, elytra and ventral surface dark brown with green or copper metallic
sheen. Length 16 to 16.5 mm. Head. Foveiform punctures dense and deep, touching on dorsal interocular surface.
Dorsal interocular distance approximately three times eye width (as in Fig. 61). Lateral margin of gena strongly
curved outward, with row of setae around eyes. Thorax. Surface of pronotum with deep and dense foveiform
punctures, touching on the sides and anterior angles. Anteromedial punctures more dispersed than those of
surrounding surface. Posteromedial punctures usually separated by less than one diameter. Margin between lateral
and posterior angles straight. Elytra. Lateral margin distinctly curved outward (Fig. 121) and surface slightly
irregular (rugose). Elytral striae wider, carinate margins clearly separated (as in Fig. 75). Interstriae clearly convex,
with its central portion nearly flat. Interstrial surface with foveiform punctures surrounded by microtubercles
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weakly defined. Interstriae usually more punctate on sides, close to striae. Foveiform punctures of third interstria
denser laterally (punctures can be so scattered at center of interstria to produce smooth midlongitudinal strip) (as in
Fig. 86). Basal carina of seventh interstria elongate, length about four or more times seventh interestria width.
Basal carina of ninth interstria surpassing anterior limit of apical carina of seventh interstria. Apical carina of third
interstria more prominent than apical carinae of second and fourth interstriae. Apical carina of sixth interstria
longer than apical carina of fifth interstria. Length of apical carina of seventh interstria longer than the sum of
lengths of apical carina of fifth and sixth interstriae. Pseudepipleural carina complete, visible entire length of
epipleuron (as in Fig. 63).
Material studied. Holotype: BRAZIL: MATO GROSSO: Nova Marilândia, (14°20'56.27"S,
57°42'40.02"W), II.2008, R.J. Silva (1 ♂ CEMT).
Paratypes [4]: BRAZIL: MATO GROSSO: Nova Marilândia, (14°20'56.27"S 57°42'40.02"W), II.2008, R.J.
Silva (1 ♀ CEMT); same, (14°19'41.37"S, 57°45'1.52"W), II.2008, R.J. Silva (1 ♂ CEMT); same, (14°20'11.36"S,
57°44'44.17"W), R.J. Silva (1 ♂ 1 ♀ CEMT).
Sampling methods. un-baited pitfall traps [5 specimens].
Habitat. Brazil (Mato Grosso) (Fig. 94). Two specimens with label data "Campo Sujo" (Cerrado) and three
specimens with "Cerrado s. str." The elevation data ranges from 522 to 613 m AMSL.
Diagnosis and remarks (within the icariforme subcomplex): length 16 to 16.5 mm; length of apical carina of
seventh interstria longer than the sum of lengths of apical carina of fifth and sixth interstriae; basal carina of ninth
interstria surpassing anterior limit of apical carina of seventh interstria. The following aedeagus characters are also
diagnostic: parameres symmetric and elongate, directed downward, almost at an angle of 90° with phallobase; apex
acuminate (Fig. 20). Lamella copulatrix (LC): left lobe subtrapezoidal (as in Fig. 40); right lobe subrectangular (as
in Fig. 40). Superior right peripheral sclerite (SRP) as in Fig. 3.
The lateral elytral margin is the most curved between species of the icariforme subcomplex. In some
specimens, apical carina of third interstria can be somewhat more prominent than apical carina of second and
fourth interstriae, with a rounded form, resembling a tubercle. Apical carina of seventh interstria presents the
largest length among the species of the valgum group.
Deltochilum (Aganhyboma) kolleri new species
(Figs 19, 49, 56, 94, 122)
Etymology. Named in honour of Wilson Werner Koller, the collecter of one of the specimens examined.
Description. Body. Head, pronotum, elytra and ventral surface dark brown or black, with a greenish metallic
sheen. Length > 13 mm. Head. Foveiform punctures dense, touching on dorsal interocular surface. Punctures
around clypeus shallower, smaller and more dispersed than those on the interocular surface. Dorsal interocular
distance approximately four times eye width. Lateral margin of gena strongly curved outward, with row of setae
around the eyes. Thorax. Surface of pronotum with dense and deep foveiform punctures, touching on the sides and
anterior angles. Discal punctures smaller and more dispersed than those of surrounding surface. Posteromedial
punctures usually separated by less than one diameter. Margin between lateral and posterior angles straight. Elytra.
Lateral margins almost pararell (Fig. 122) and surface slightly irregular (rugose). Elytral striae wider, carinate
margins clearly separated (as in Fig. 75). Interstriae clearly convex, with its central portion nearly flat. Interstrial
surface with foveiform punctures surrounded by microtubercles weakly defined, conspicuous on lateral of elytra.
Interstriae usually more punctate on the sides, close to striae. Foveiform punctures of third interstria denser
laterally (punctures can be so scattered at center of interstria to produce smooth midlongitudinal strip) (as in Fig.
86). Foveiform puncture surrounded by approximately six microtubercles. Basal carina of seventh interstria
elongate, length about four or more times seventh interestria width. Basal carina of ninth interstria present in three-
quarters of elytra (not reaching anterior limit of apical carina of seventh interstria). Apical carina of third interstria
rounded, better defined as tubercle, more prominent than apical carina of second and fourth interstriae. Length of
apical carina of sixth interstria longer than that of apical carina of fifth interstria. Length of apical carina of seventh
interstria less than or equal to the sum of lengths of apical carina of fifth and sixth interstriae. Pseudepipleural
carina complete, visible entire length of epipleuron (as in Fig. 63).
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Material studied. Holotype: BRAZIL: MATO GROSSO DO SUL: Coxim, 26.II.1986, Koller (1 ♂ CEMT).
Paratypes [5]: BRAZIL: MINAS GERAIS: São Roque de Minas, Parque Nacional Serra da Canastra,
(20°15'17"S, 46°25'11"W), 9.XI.2007, M. F. Souza (1 ♂ 2 ♀ CEMT); SÃO PAULO: Piracicaba, E.E.C.J.V.,
30.IV.1966 (1 ♂ CEMT); [no data] (1 ♂ CEMT).
Sampling methods. at light-trap [1 specimen].
Habitat. Brazil (Minas Gerais and Mato Grosso) (Fig. 94). From available data this species inhabits Cerrado.
Diagnosis and remarks (within the icariforme subcomplex): length > 13 mm; posteromedial punctures of
pronotum usually separated by less than one diameter; length of apical carina of seventh interstria less than or equal
to the sum of lengths of apical carinae of fifth and sixth interstriae; lateral margins of elytra almost parallel. The
following aedeagus characters are also diagnostic: parameres symmetric and elongate, directed downward, almost
at an angle of 60° with phallobase; apex weakly acuminate (Fig. 19). Lamella copulatrix (LC): left lobe
subrectangular (as in Fig. 41); right lobe “b”-shaped (as in Fig. 41). Superior right peripheral sclerite as in Fig. 49.
Fronto-lateral peripheral sclerite (FLP) as in Fig. 56.
The punctures at center of elytral interstriae, on anterior portion of elytra, are the most dense of icariforme
subcomplex. Apical carina of sixth interstria can present the same length of apical carina of fifth interstria.
Deltochilum (Aganhyboma) subrubrum new species
(Figs 23, 41, 86, 94, 123)
Etymology. Subrubrum (reddish), Latin adjective, refers to the color of individuals.
Description. Body. Head, pronotum, elytra and ventral surface dark brown or black, with greenish or reddish
metallic sheen. Length 14.8 to 15.8 mm. Head. Foveiform punctures dense, touching on dorsal interocular surface.
Punctures around clypeus shallower, smaller, and more dispersed than those on the interocular surface. Dorsal
interocular distance approximately four times eye width. Lateral margin of gena strongly curved outward, with row
of setae around eyes. Thorax. Surface of pronotum with dense foveiform punctures, touching on the sides and
anterior angles (however not as deep as in D. icariforme). Posteromedial punctures separated by at least one
diameter. Margin between lateral and posterior angles straight. Elytra. Lateral margins almost parallel (Fig. 123)
and surface slightly irregular (rugose). Elytral striae wider, carinate margins clearly separated (as in Fig. 75).
Interstriae clearly convex, with its central portion nearly flat. Interstriae usually more punctate on the sides, close to
striae. Foveiform punctures of third interstria denser laterally (punctures can be so scattered at center of interstria to
produce smooth midlongitudinal strip) (Fig. 86). Microtubercles of interstriae almost inconspicuous. Basal carina
of seventh interstria elongate, length about four or more times the width of the seventh interestria. Basal carina of
ninth interstria present in three-quarters of elytra (not reaching anterior limit of apical carina of seventh interstria).
Apical carina of third interstria rounded, better defined as tubercle, more prominent than apical carinae of second
and fourth interstriae. Length of apical carina of sixth interstria longer than that of apical carina of fifth interstria.
Length of apical carina of seventh interstria less than or equal to the sum of lengths of apical carinae of fifth and
sixth interstriae. Pseudepipleural carina complete, visible entire length of epipleuron (as in Fig. 63).
Material studied. Holotype: ARGENTINA: MISIONES: Loreto, 1955, F. H. Walz (1 ♂ DZUP).
Paratypes [3]: ARGENTINA: MISIONES: Loreto (1 ♀ CEMT); same, San Ignacio (1 ♂ 1 ♀ MNHN).
Diagnosis and remarks (within the icariforme subcomplex): length 14.8 to 15.8 mm; posteromedial punctures
of pronotum separated by at least one diameter; length of apical carina of seventh interstria less than or equal to the
sum of lengths of apical carinae of fifth and sixth interstriae; microtubercles of elytral interstriae absent or almost
inconspicuous. The following aedeagus characters are also diagnostic: parameres symmetric and elongate, directed
downward, almost at an angle of 60° with phallobase; apex weakly acuminate (Fig. 23). Lamella copulatrix (LC):
left lobe subrectangular (Fig. 41); right lobe "b"-shaped (Fig. 41). Fronto-lateral peripheral sclerite (FLP) as in Fig.
56.
We noted a single female with lateral margins of elytra distinctly curved outward.
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FIGURES 4–11. Aedeagus in Deltochilum (Aganhyboma). 4. D. titovidaurrei new species. 5. D. cupreicolle. 6. D. viridescens
new status. 7. D. viridicatum new species. 8. D. amandaarcanjoae new species. 9. D. trisignatum. 10. D. cangalha new
species. and D. alpercata new species. 11. D. finestriatum new species. Figures scale = 1 mm.
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FIGURES 12–19. Aedeagus in Deltochilum (Aganhyboma). 12. D. schefflerorum new species. 13. D. acropyge new status.
14. D. larseni new species. 15. D. acanthus. 16. D. longiceps new status. 17. D. valgum. 18. D. icaroides. 19. D. kolleri new
species. Figures scale = 1 mm.
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FIGURES 20–23. Aedeagus in Deltochilum (Aganhyboma). 20. D. paresi new species. 21. D. icariforme. 22. D. ritamourae
new species. 23. D. subrubrum new species. Figure 24. Aedeagus illustration showing the angle between the phallobase and
the parameres. Figures 25–27. Genital segment in Deltochilum (Aganhyboma). 25. D. cupreicolle. 26. D. viridescens new
status. 27. D. trisignatum. Figures scale = 1 mm.
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FIGURES 28–32. Genital segment in Deltochilum (Aganhyboma). 28. D. finestriatum new species. 29. D. schefflerorum new
species 30. D. valgum. 31. D. icaroides. 32. D. icariforme. Figures 33–36. Lamella copulatrix sclerites in D. (Aganhyboma).
33. D. cupreicolle. 34. D. viridescens new status and D. viridicatum new species 35. D. trisignatum. 36. D. finestriatum new
species. Figures scale = 1 mm.
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FIGURES 37–41. Lamella copulatrix sclerites in D. (Aganhyboma). 37. D. longiceps new status. 38. D. valgum. 39. D.
icaroides. 40. D. icariforme. 41. D. subrubrum new species. Figures 42–49. Superior right peripheral sclerite (SRP) in D.
(Aganhyboma). 42. D. titovidaurrei new species. 43. D. viridescens new status. 44. D. trisignatum. 45. D. finestriatum new
species. 46. D. acropyge new status. 47. D. valgum. 48. D. icaroides. 49. D. kolleri new species. Figures 50–54. Fronto-lateral
peripheral sclerite (FLP) in D. (Aganhyboma). 50. D. cupreicolle. 51. D. viridescens new status. 52. D. trisignatum. 53. D.
finestriatum new species. 54. D. schefflerorum new species. Figures scale = 1 mm.
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FIGURES 55–58. Fronto-lateral peripheral sclerite (FLP) in D. (Aganhyboma). 55. D. valgum. 56. D. kolleri new species. 57.
D. icariforme. 58. D. ritamourae new species. Figure 59. Complex of axial and subaxial sclerites (A+AS) in D. viridescens
new status. Figures 60–61. Head in D. (Aganhyboma). 60. D. viridescens new status. 61. D. longiceps new status. Figures
62–63. Lateral view of the elytra in D. (Aganhyboma). 62. D. viridescens new status (arrow points the basal carina of ninth
interstria). 63. D. finestriatum new species. (arrow points the basal carina of ninth interstria). Figures 64–69. Legs. 64.
Metafemur in D. viridescens new status. 65. Metafemur in D. longiceps new status. 66. Protibia in D. trisignatum. 67.
Metatibia in D. trisignatum. 68. Metatibia in D. viridescens new status. 69. Profemur in D. trisignatum (arrow indicates the
anterior margin discontinuous). Figures scale = 1 mm.
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FIGURE 70. Detail of the interstrial surface in D. violaceum. Figures 71–72. Detail of the basal carina of seventh interstria in
D. (Aganhyboma). 71. D. viridescens new status. 72. D. cupreicolle. Figure 73. Abdominal ventrites in D. viridescens new
status (male; arrow points to apex of pygidium). Figures 74–77. Detail of the third elytral interstria in D. (Aganhyboma). 74.
D. finestriatum new species. (arrow points to third elytral interstria). 75. D. valgum (arrow points to third elytral interstria). 76.
D. cangalha new species. 77. D. alpercata new species. Figures 78–79. hypomera in D. (Aganhyboma). 78. D. cangalha new
species. 79. D. alpercata new species. Figure 80. Lateral view of the elytra in D. longiceps new status.
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FIGURES 81–83. Illustrations of hypomera in D. (Aganhyboma). 81. D. schefflerorum new species. 82. D. arturoi new
species. 83. D. longiceps new status. Figure 84. Right elytron of D. longiceps new status (arrows indicate the irregular
spacing among the points of the elytral striae). Figure 85. Pronotum in D. icaroides. Figure 86. Apex of third interestria in D.
subrubrum new species. Figures 87–88. Detail of the apical tubercle of third interstria in D. (Aganhyboma). 87. D. icaroides.
88. D. valgum.
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FIGURES 89–92. Known distribution of D. (Aganhyboma). 89. trisignatum complex. 90. cupreicolle complex. 91. acropyge
complex. 92. finestriatum complex.
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FIGURES 93–94. Known distribution of D. (Aganhyboma). 93. D. icaroides and D. valgum. 94. icariforme subcomplex.
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FIGURES 95–102. Species of D. (Aganhyboma). 95. D. trisignatum. 96. D. kolbei. 97. D. violaceum. 98. D. amandaarcanjoae
new species. 99. D. cupreicolle. 100. D. titovidaurrei new species. 101. D. viridescens new status. 102. D. viridescens new
status. Figures scale = 1mm.
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FIGURES 103–110. Species of D. (Aganhyboma). 103. D. viridescens new status. 104. D. viridicatum new species. 105. D.
viridicatum new species. 106. D. acropyge new status. 107. D. feeri new species. 108. D. schefflerorum new species. 109. D.
streblopodum new species. 110. D. longiceps new status. Figures scale = 1 mm.
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FIGURES 111–118. Species of D. (Aganhyboma). 111. D. acanthus. 112. D. arturoi new species. 113 . D. larseni new species.
114 . D. cangalha new species. 115 . D. alpercata new species. 11 6. D. finestriatum new species. 117. D. icaroides. 118 . D.
valgum. Figures scale = 1 mm.
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FIGURES 119–123. Species of D. (Aganhyboma). 119. D. ritamourae new species. 120. D. icariforme. 121. D. paresi new
species. 122. D. kolleri new species. 123. D. subrubrum new species. Figures scale = 1 mm.
Acknowledgments
Fernando Silva was supported by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—CAPES during
his PhD. We also thank Conselho Nacional de Desenvolvimento Científico e Tecnológico—CNPq for providing
PQ2 fellowships and research grants for Fernando Vaz-de-Mello (304925/2010-1, 302997/2013-0, 405697/2013-9,
484035/2013-4, 202327/2013-2) and research grants for Fernando Silva (444020/2014-4). Thanks are due for all
curators or collections managers cited in the "material" section. We are especially grateful to the following
researchers for their assistance and important contribution to various aspects of this work: Arturo González,
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François Génier, Fredy Molano, Jorge Arias, Jiří Hájek and Sergei Golovatch. We are endlessly thankfull to W. D.
Edmonds, Jiri Zídek, Darren Mann, Sergei Tarasov for their valuable suggestions on the text of the manuscript and
to Andrey Frolov for reviewing the English.
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... The Neotropical genus Not long after of the works of Paulian [8,9] and Balthasar [30], some American scarabaeidologists recognised the need for a taxonomic revision of Deltochilum due to its heterogeneity, the amount of divisions (subgenera) and the numerous new species remaining to be described in collections [34][35][36][37][38]. The attempt towards a modern and comprehensive taxonomic revision of Deltochilum focused on subgenera resulted in the description of 24 new species [25,26,33,39,40] and the description of new genus [41]. ...
... It is possible to identify species belonging to Deltohyboma using the key for American genera of Scarabaeinae [59]. However, after Silva et al. [26] transferred some species of Deltohyboma to D. (Aganhyboma), the species now belonging to acropyge and valgum complexes of the subgenus Aganhyboma (see Silva et al., [26]) would still be identified as if they were Deltohyboma. However, these species can be correctly identified as belonging to D. (Aganhyboma) by the head, which is slightly wider than long (Fig 2D). ...
... It is possible to identify species belonging to Deltohyboma using the key for American genera of Scarabaeinae [59]. However, after Silva et al. [26] transferred some species of Deltohyboma to D. (Aganhyboma), the species now belonging to acropyge and valgum complexes of the subgenus Aganhyboma (see Silva et al., [26]) would still be identified as if they were Deltohyboma. However, these species can be correctly identified as belonging to D. (Aganhyboma) by the head, which is slightly wider than long (Fig 2D). ...
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Deltochilum Eschscholtz, 1822 is perhaps the most speciose genus of the tribe Deltochilini sensu Tarasov & Dimitrov (2016) (Scarabaeidae: Scarabaeinae) and has been traditionally divided into eight subgenera. Among them, the subgenus Deltohyboma Lane, 1946, is the most speciose with 47 species, five of which are described here ( D . genieri sp. nov. , D . gilli sp. nov. , D . susanae sp. nov. , D . bolivariensis sp. nov. and D . inesae sp. nov. ), and at least 165 species still undescribed. Due to the large number of species, and for practical purposes, the subgenus is here divided into 19 species-groups, with D . inesae sp. nov. left as incertae sedis . This division into species-groups will help in the reliable identification of species and will aid in the completion of the revision of subgenus. This report is the first part of the taxonomic revision of the subgenus Deltohyboma ; it is based on the examination of all type specimens and almost 9,800 specimens of which approximately 1,200, mostly males, had their genitalia studied. The 19 species-groups recognized here are based mainly on characters described for the first time for Deltohyboma , namely, the state of a) the anterior margin of the clypeus (between the clypeal teeth), b) the internal margin of hypomera, c) the ventral face of the protibia, d) the posterior margin of the metafemur, and e) several new characters resulting from the first detailed study of the aedeagus and the endophallus of the group. Diagnosis, description, geographic distribution, composition and identifications keys (for males as well as males and females) for the species-groups are provided.
... Fortunately, recent taxonomic reviews have allowed an accurate identification of several species belonging to the following genera addressed in the present work: Canthon Hoffmannsegg, 1817 (Nunes et al. 2018Vieira et al. 2019;Vaz-de-Mello et al. 2020), Copro phanaeus d' Olsoufieff, 1924 (Edmonds andZídek 2010), Deltochilum Eschscholtz, 1822 (Génier 2012;González-Alvarado and Vaz-de-Mello 2014;Silva et al. 2015), Dichotomius Hope, 1838 (Nunes and Vaz-de-Mello 2013Maldaner et al. 2015Maldaner et al. , 2018Valois et al. 2017;Arias-Buriticá and Vaz-de-Mello 2019;Rossini and Vaz-de-Mello 2020;Silva et al. 2020), Eurysternus Dalman, 1824 (Génier 2009), Phanaeus MacLeay, 1819 (Edmonds and Zídek 2012), and Oxysternon Castelnau, 1840 (Edmonds and Zídek 2004). ...
... For identification at genus level, the key to genera and subgenera of Vaz- was used. Some of the identifications at species level were accomplished with the aid of taxonomic reviews (Edmonds and Zídek 2004Génier 2009;Silva et al. 2015;Valois et al. 2017). ...
... In the EPA Aldeia-Beberibe in Pernambuco, one collected specimen, found perched on a leaf in the forest understory, showed black elytron and dark-green pronotum, both shiny, which is an atypical melanic variation in this species. Silva et al. (2015), specimens of the D. alpercata can be distinguished from other Deltochilum by having the anterior part of head (between clypeogenal sutures) prolonged, similar to a small 'muzzle' (Silva et al. 2015: figs 60, 61); head with small clypeal teeth external to central teeth (Silva et al. 2015: fig. 61); dorsal interocular distance three to four times eye width (Silva et al. 2015: fig. ...
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Dichotomius valoisae new species (Coleoptera: Scarabaeidae: Scarabaeinae: Dichotomiini) from Santa Luzia do Itanhi, Sergipe, Brazil, is described based on external differences from its congeners. Diagnostic characters and a brief discussion about its conservation status are provided, as well as an updated identification key to the Dichotomius sericeus (Harold) species group.
... Deltochilum are roller beetles typical of tropical, subtropical and some temperate habitats (Génier, 2012;Halffter, 2003). Species in Deltochilum have a wide variety of trophic habits as coprophagy and necrophagy, with some species attracted to rotting fruits and fungi, or specialized to prey on diplopods (Cano, 1998;Capello & Halffter, 2019;Halffter, 2003;Halffter & Halffter, 2009;Larsen et al., 2009;Silva et al., 2012Silva et al., , 2015Silva et al., , 2017. ...
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... aedeagus) and internal (i.e. sclerites of the internal sac of the aedeagus) parts of the male genitalia as morphological characters useful for diagnosing clades at various taxonomic levels (López-Guerrero et al., 2009;Tarasov & Solodovnikov, 2011;Manjarrés & Molano, 2015;Silva et al., 2015;Tarasov & Génier, 2015;Génier & Moretto, 2017;Rossini et al., 2018). The external morphology of the aedeagus has helped to separate closely related species, and even subspecies, of the genus Canthon (e.g. ...
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... The list of species obtained was reviewed and updated according to the suprageneric designation proposed by Bouchard et al. (2011) and although there are 11 recognized tribes, only seven are found in Mexico. Supra-specific revisions of the genera Canthon (Rivera-Cervantes and Halffter 1999), Coprophanaeus (Edmonds and Zídek 2010), Deltochilum (Génier 2012;González-Alvarado and Vaz-de-Mello 2014;Silva et al. 2015), Dichotomius (López-Guerrero 2005), Martinezidium (Vazde-Mello 2008) and Phanaeus (Edmonds and Zídek 2012), were also taken into account because they include changes of status for several species on the list. Some species were omitted from the list and those records were considered erroneous or corresponded to incorrect geographic records (see discussion). ...
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... Species of Deltochilum, as is the case with some other tropical dung beetles, are associated with carrion and have been widely used as indicators of habitat transformations ( Cultid et al. 2012). Deltochilum has not yet been fully reviewed, but some subgenera have been revised recently ( González et al. 2009; Génier 2012; González-Alvarado and Vaz-de-Mello 2014; Silva et al. 2015). Génier (2012) Calhyboma is the most heterogeneous subgenus of Deltochilum, with 12 valid species after this work. ...
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Streblopus Van Lansberge, 1874 has been one of the most mysterious dung beetle groups of the Neotropical fauna, having a rather peculiar morphology, very few known specimens in collections and a difficult placement among the scarabaeine lineages. In this work, based on the examination of a recently collected series of specimens and a synthesis of some scattered, but deeply valuable, information available in the literature, we readdress many of the questions posed by past authors. It is shown that Streblopus is a relict genus composed of two currently living species of widely disjunct distribution, namely S. opatroides Van Lansberge, 1874, from patches of Atlantic Forest in the Brazilian states of Bahia and Espírito Santo, and S. punctatus (Balthasar, 1938), known from a few localities across Sub-Andean humid forests in the Peruvian and Ecuadorian Amazon Forest. We redescribe both and present in detail the evidence pointing to their validity as two independent species; a discussion of their remarkable sexual dimorphism is also given. The biogeography of Streblopus in South America is addressed, and we conclude that the present disjunct distribution of the genus is a consequence of the retreat of the tropical forest corridors that once connected the Atlantic Forest to the Amazon Basin through the South American Dry Diagonal during several periods of the Neogene, particularly until the Middle Miocene. Finally, we propose an African origin for the genus based on its close phylogenetic relationship with a group of Old World taxa ‒ particularly Circellium Latreille, 1825 and Scarabaeini. Having diverged from those groups in the late Upper Cretaceous, we argue that the ancestor of Streblopus arrived in South America crossing the Atlantic Ocean by rafting. We present a synthesis of data from a wide variety of biological groups to support our ideas and contend that long-distance dispersal hypotheses should be taken more seriously by scarab beetle specialists.