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The genus Gromphas Dejean, 1836 (Coleoptera, Scarabaeinae): nomenclature,

distribution, and conservation, including a contribution to the debate on
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 The genus Gromphas Dejean, 1836
(Coleoptera, Scarabaeinae): nomenclature, distribution,
and conservation, including a contribution to the debate
on electronic publications in zoology

Mario CUPELLO
Laboratório de Sistemática e Bioecologia de Coleoptera, Departamento de Zoologia,
Universidade Federal do Paraná, Caixa Postal 19020, 81531-980, Curitiba, Paraná (Brazil);
and Department of Entomology, Texas A&M University,
College Station, TX 77843 (United States)
[email protected]

Submitted on 11 April 2023 | Accepted on 26 July 2023 | Published on 9 February 2024

urn:lsid:zoobank.org:pub:4B49C1D9-1196-4942-969F-2E923B1FC12C

Cupello M. 2024. — The genus Gromphas Dejean, 1836 (Coleoptera, Scarabaeinae): nomenclature, distribution, and
conservation, including a contribution to the debate on electronic publications in zoology. Zoosystema 46 (2): 23-59.
https://doi.org/10.5252/zoosystema2024v46a2. http://zoosystema.com/46/2

ABSTRACT
Previously overlooked literature now brought to my attention has resulted in the following nomen-
clatural conclusions: 1) The species known since 2013 as Gromphas inermis Harold, 1869 must be
called G. lacordairii (Oken, 1834), whose original combination was Coprobius lacordairii. Gromphas
inermis is a new junior subjective synonym of G. lacordairii; 2) Gromphas was made available by De-
jean, in 1836, not Brullé, in 1837; the former, therefore, should be credited with the authorship; and
3) the type species of Gromphas is Coprobius lacordairii Oken, 1834 by original monotypy, not Onitis
aerugionosus Perty, 1830 by subsequent monotypy. Also discussed is the publication year of G. jardim
Cupello & Vaz-de-Mello, 2015. This leads me to address the problem of zoological works first pub-
lished in electronic-only versions with their own pagination and which are later reissued integrated
into a journal’s volume and repaginated. It is here argued that these two versions – the detached and
KEY WORDS the volume-integrated ones – should be deemed separate available works, and that new nomenclatu-
Gromphas dichroa,
Gromphas lacordairii, ral acts can be made available from detached versions. If this is accepted, the later publication of the
Gromphas lacordairei, volume-integrated versions has no bearing on the availability of the earlier detached versions. I also
Gromphas inermis, introduce new data on the type series of Onitis aeruginosus (currently, Gromphas aerugionosa) and
publication date,
preprint, G. inermis, new specimens of the rare G. jardim, mislabelled specimens of G. amazonica Bates, 1870,
available work, and newly discovered specimens of the vanished G. dichroa Blanchard, 1846. The latter include the
extinction,
disappeared species, new record from Santa Catarina state, Brazil. The disappearance of G. dichroa since 1954, including
South America. the possibility that it might be extinct, is discussed.

ZOOSYSTEMA • 2024 • 46 (2) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 23
 Cupello M.

RÉSUMÉ
Le genre Gromphas Dejean, 1836 (Coleoptera, Scarabaeinae) : nomenclature, distribution et conservation,
avec une contribution au débat sur les publications électroniques en zoologie.
La littérature précédemment ignorée et maintenant portée à mon attention a abouti aux conclusions
nomenclaturales suivantes : 1) l’espèce connue depuis 2013 sous le nom de Gromphas inermis Harold,
1869 doit être appelée G. lacordairii (Oken, 1834), dont la combinaison originale était Coprobius
lacordairii. Gromphas inermis est un nouveau synonyme subjectif junior de G. lacordairii ; 2) Gromphas
a été rendu disponible par Dejean, en 1836, et non par Brullé, en 1837 ; c’est donc le premier qui
doit être crédité de la paternité; 3) l’espèce type de Gromphas est Coprobius lacordairii Oken, 1834
selon la monotypie originale, et non Onitis aerugionosus Perty, 1830 selon la monotypie ultérieure.
L’année de publication de G. jardim Cupello & Vaz-de-Mello, 2015, est également discutée. Ceci
m’amène à aborder le problème des travaux zoologiques publiés pour la première fois en version
électronique avec leur propre pagination et qui sont ensuite réédités, intégrés dans le volume d’une
revue et repaginés. L’argument avancé ici est que ces deux versions – la version détachée et la version
intégrée au volume – devraient être considérées comme des œuvres disponibles distinctes, et que de
MOTS CLÉS nouveaux actes nomenclaturaux peuvent être rendus disponibles à partir des versions détachées. Si
Gromphas dichroa,
Gromphas lacordairii, l’on accepte ce point de vue, la publication ultérieure des versions intégrées au volume n’a aucune
Gromphas lacordairei, incidence sur la disponibilité des versions détachées antérieures. Je présente également de nouvelles
Gromphas inermis, données sur les séries types d’Onitis aeruginosus (présentement Gromphas aerugionosa) et de G. iner-
date de publication,
preprint, mis, de nouveaux spécimens du rare G. jardim, des spécimens mal étiquetés de G. amazonica Bates,
travaux disponibles, 1870 et des spécimens nouvellement découverts de G. dichroa Blanchard, 1846, qui a disparu. Ces
extinction,
espèces disparues, derniers comprennent le nouveau signalement de l’État de Santa Catarina, Brésil. La disparition de
Amérique du Sud. G. dichroa depuis 1954, y compris la possibilité qu’il soit éteint, est discutée.

INTRODUCTION In another part of the paper, I shall address the publication

date of Gromphas jardim Cupello & Vaz-de-Mello, a problem
Some years ago, Fernando Vaz-de-Mello and I published a that has to do with the controversy over the availability of
series of papers dealing with the systematics of the South electronic-only versions of works that appear with pre-volume
American dung beetle genus Gromphas (Cupello 2013; Cu- pagination (Dubois et al. 2013, 2015a, b; Krell 2015). I will
pello & Vaz-de-Mello 2013, 2014, 2015). All the main sys- offer a solution to this problem that has apparently not been
tematic aspects were addressed, from nomenclature to species put forward before. In the continuation, I will present data
delimitation and intraspecific variation to the phylogeny of on specimens newly studied by me in collections around
the genus and its relationships with other scarabaeine groups. the world, including the type series of Onitis aeruginosus
Also addressed were the ecology and distribution of the spe- (currently Gromphas aeruginosa), some new specimens of the
cies. Yet, unsurprisingly, new discoveries have accumulated rare G. jardim, mislabelled specimens of G. amazonica Bates,
since then and some have made me reconsider a few of the 1870, and a few new specimens of the vanished G. dichroa
previous conclusions. This paper is a compilation of these Blanchard, 1846. The latter include a new state record for
new findings. Brazil and the most recent collection date for the species,
I shall first address nomenclatural issues that change three 1954. The reasons for the disappearance of the species over
aspects of our revision. Firstly, the valid name of the spe- the past 70 years, including the possibility of extinction, are
cies we called Gromphas inermis Harold, 1869, a common examined.
element of middle South America, is actually Gromphas
lacordairii (Oken, 1834). Secondly, the authorship of the
genus-group name Gromphas does not belong to Brullé MATERIAL AND METHODS
(1838) as we said, but to Dejean (1836). Thirdly, the type
species of Gromphas is not Onitis aeruginosus Perty, 1830 Collections
by subsequent monotypy by Sturm (1843), but Coprobius The following collections were visited during the research for
lacordairii Oken, 1834 by original monotypy. These novelties this paper (curator or another contact in parentheses):
are all based on my reevaluation of the nomenclatural history
of the genus and its species in light of my recent reading CAPC Carlos Aguilar personal collection, Capiatá;
of previously overlooked literature. The two key references CEAH Coleção Entomológica Adolph Hempel, Instituto
that had been overlooked are Lacordaire’s (1830) memoir of Biológico, São Paulo (Sergio Ide);
his beetle explorations in South America and Oken’s (1834) CEMT Coleção Entomológica de Mato Grosso Eurides
Furtado, Universidade Federal de Mato Grosso,
review of Lacodaire’s paper. Both were recently brought to Cuiabá (Fernando Z. Vaz-de-Mello);
my attention by my colleague and ICZN Commissioner CMNC Canadian Museum of Nature, Ottawa (François
Patrice Bouchard. Génier);

24 ZOOSYSTEMA • 2024 • 46 (2)

CNCI Canadian National Collection of Insects, Arachnids Material housed in the following collection is included on
and Nematodes, Ottawa (Patrice Bouchard and the list of material examined from label data kindly provided
Serge Laplante); by Patricia González-Vainer:
DZUP Coleção Entomológica Padre Jesus Santiago Moure,
Departamento de Zoologia, Universidade Federal
do Paraná, Curitiba (Lucia Massutti de Almeida);
FSCA Florida State Collection of Arthropods, Gainesville, FCUR Faculdad de Ciencias, Universidad de la República,
Florida (Paul Skelley); Montevideo (Enrique Morelli).
INPA Instituto Nacional de Pesquisas da Amazônia, Manaus
(Marcio Luiz de Oliveira);
LSUK The Linnean Society of London, London (Suzanne Finally, the geographical provenance of the Gromphas
Ryder); specimens housed in the following collections was com-
MCNZ Museu de Ciências Naturais da Fundação Zoobotânica piled by Pedro Giovâni da Silva in 2011 and graciously
do Rio Grande do Sul, Porto Alegre (Luciano de shared with me in March 2023 for inclusion in this work.
Azevedo Moura); These collections are all located in the Brazilian state of
MCZC Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge, Massachusetts (Charles Farnum, Rio Grande do Sul. Note that, in his Herculean effort of
Crystal Maier and Rachel Hawkins); cataloguing all the Scarabaeinae from these regional col-
MFNB Museum für Naturkunde Berlin, Leibniz-Institut lections, da Silva did not have time to take note of the
für Evolutions- und Biodiversitätsforschung, Ber- exact number of specimens existing from each locality
lin (Joachim Willers, Bernd Jaeger, and Johannes and each date. Therefore, this information is not provided
Frisch);
MGAP Museu Anchieta de Ciências Naturais, Colégio in Figure 14 or in Table 1, where it would be expected
Anchieta, Porto Alegre (Fernando Meyer); to be found.
MHNG Muséum d’Histoire naturelle de Genève, Geneva
(Giulio Cuccodoro); LBEV Coleção Entomológica do Laboratório de Biologia
MNHN Muséum national d’Histoire naturelle, Paris (Olivier Evolutiva, Universidade Federal de Santa Maria,
Montreuil and Antoine Mantilleri); Santa Maria;
MNHNP Museo Nacional de Historia Natural del Paraguay, MCCR Acervo entomológico do Museu de Ciências Naturais
San Lorenzo (Bolívar Garcete); Carlos Ritter, Pelotas (currently part of the holdings
MPEG Museu Paraense Emílio Goeldi, Belém (Orlando of Laboratório de Zoologia, Departamento de Zoo-
Tobias Silveira); logia e Genética, Instituto de Biologia, Universidade
MZSP Museu de Zoologia, Universidade de São Paulo, São Federal de Pelotas, Pelotas, Brazil);
Paulo (Sonia Casari and Carlos Campaner); MECB Coleção do Museu de Entomologia Ceslau Maria
MZUFPA Museu de Zoologia, Instituto de Ciências Biológi- Biezanko, Pelotas;
cas, Universidade Federal do Pará, Belém (Fernando MCTP Coleção de Insetos do Museu de Ciências e Tecno-
A. B. Silva); logia da Pontifícia Universidade Católica do Rio
NHM The Natural History Museum, London (Maxwell Grande do Sul, Porto Alegre;
Barclay), formerly BMNH; MHNU Coleção do Museu de História Natural da Univer-
NHMB Naturhistorisches Museum Basel, Basel (Isabelle sidade Católica de Pelotas, Pelotas;
Zürcher and Matthias Borer); MRGC Acervo do Museu Professor Ramiro Gomes Costa,
NHMW Naturhistorisches Museum Wien, Vienna (Harald Porto Alegre.
Schillhammer);
NMPC National Museum (Natural History), Prague (Jiří
Hájek); Nomenclatural Principles
OUMNH Hope Entomological Collections, Oxford Univer-
sity Museum of Natural History, Oxford (Darren
Given the convoluted nomenclatural problems associated
Mann); with Gromphas and its species (Cupello 2013; Cupello &
RBINS Institut royal des Sciences naturelles de Belgique, Vaz-de-Mello 2013, 2014; herein), I gave special attention
Brussels (Alain Drumont); to the dating of nomenclaturally relevant publications.
RMNH Naturalis Biodiversity Centre, Leiden (Hans Hui- I based this dating primarily on Evenhuis (1997a, b; 2003;
jbregts); 2015; personal communication), Bousquet (2016), and
SDEI Senckenberg Deutsches Entomologisches Institut,
Müncheberg (Stephan Blank); the literature cited therein. For Edgar von Harold’s papers
SMF Forschungsinstitut und Naturmuseum Senckenberg, published in Coleopterologische Hefte, I followed Bouchard
Frankfurt-am-Main (Andrea Hastenpflug-Vesmanis; et al. (2011) and based their dating on when the journal’s
current curator: Marianna V. P. Simões); respective volumes were recorded as received in donation
SMTD Museum für Tierkunde, Senckenberg Naturhis- by the Entomological Society of London (current Royal
torische Sammlungen Dresden, Dresden (Olaf Jäger
and Klaus-Dieter Klass); Entomological Society) in the society’s Proceedings. See in
TAMU Texas A&M University, College Station, Texas (John the references section comments on the publication date
D. Oswald, Mario Cupello, and Edward G. Riley); of each 19th-century publication (and some 20th-century
ZFMK Zoologisches Forschungsinstitut Alexander Koenig, ones) relevant to the nomenclatural issues discussed herein.
Bonn (Dirk Ahrens); All nomenclatural discussions and decisions are based on
ZMSC Zoologische Staatssammlung München, Munich
(Michael Balke and Ditta A. Balke); the provisions of the International Code of Zoological
ZMUK Zoologische Museum, Universität Kiel, Kiel (Michael Nomenclature (ICZN 1999), hereafter referred to simply
Kuhlmann). as ‘the Code’.

25
 Cupello M.

ON THE AUTHORSHIP OF THE GENUS-GROUP 4) The third and final edition of Dejean’s catalogue was
NAME GROMPHAS, AND THE AUTHORSHIP, published in parts between 1836 and 1837. The page list-
ORIGINAL COMBINATION, AND SPELLING ing ‘Gromphas. Dejean’ and ‘[Gromphas] lacordairei. Dej.’,
OF THE SPECIES-GROUP NAME page 159, appeared in 1836 (Madge 1988; Bousquet 2016)
‘GROMPHAS LACORDAIREI’ (Fig. 2). Like in the previous edition, no description or diag-
nosis was provided for either the genus or the species.
Nomenclatural history 5) Brullé (1838: 304-305) (Figs 1; 2) considered ‘Gromphas,
The nomenclatural history related to the establishment of Dej.’ a subgenus of Copris Geoffroy, 1762 and described the
the names Gromphas and ‘Gromphas lacordairei’, as the latter taxon by saying: ‘This subgenus is composed of one sole spe-
has been commonly spelled, can be summarised as follows: cies that has the appearance of the Phanées [i.e., the subgenus
1) Lacordaire (1830) (Figs 1; 2), in a memoir of his South Phanaeus of Copris] and their broad sternal plate [i.e., metaven-
American coleopterological observations (1824-1830), listed, trite], as well as their antennae with arched, cup-shaped club
family by family, the many species that he had seen in the articles; it is distinguished only by the presence of anterior tarsi
southern continent. Following four paragraphs on pages in males as well as in females’ (Fig. 2). Even though Brullé
260 and 261 discussing the species of Coprobius Latreille, explicitly said that Gromphas included only one species, the
1829 (current junior synonym of Canthon Hoffmannsegg, name of that species was never mentioned by him. While it
1817), he introduced a new paragraph on a new dung is evident from the context that he was referring to Dejean’s
beetle species from Buenos Aires, ‘of a rather large size, of ‘Gromphas lacordairei’, this has no nomenclatural bearing.
metallic color’, which Dejean, the person responsible for In a footnote to page 304, Brullé added that the subgenus
identifying the Coleoptera that Lacordaire had brought Gromphas was still new (‘Sous-genre encore inédit’), alluding
back to Europe (Lacordaire 1830: 187), decided to name to the fact that Dejean had never described it.
after him (Fig. 2). The name of this new species was not 6) Sturm (1843: 108), in a catalogue of his beetle collec-
expressly given, however. This new species, according to tion, listed ‘Gromphas. Dej.’ with a single species, Onitis
Lacordaire, and certainly writing from information pro- aeruginosus Perty, 1830.
vided by Dejean, ‘must belong to a new genus’. Lacordaire 7) Arguing that, although already ‘characterised’ by Brullé
ended the paragraph by adding that the new species is (1838), no species had been described for Gromphas until then,
‘fairly commonly found in the excrement of horses, and Blanchard (1846: 181-182) described ‘Gromphas Lacordairei,
digs, like Copris, deep holes in the earth, a habit foreign to Dej., Catal.’ from specimens collected in Santa Cruz de la
the aforementioned Ateuchus’. In the following paragraph, Sierra, Bolivia, and G. dichroa, from Montevideo, Uruguay.
he addressed another new species, ‘Ateuchus arachnoides’, 8) Harold (1869b), after explaining that the dung beetles
from Tucumán, which reminded him of a spider and was described by Blanchard as G. lacordairei referred not to La-
said to also belong to a new genus. cordaire’s Buenos Aires species, but to the same species as
2) Three years later, Dejean (1833) (Figs 1; 2), in the sec- Gromphas aeruginosa (= Onitis aeruginosus), and that Dejean’s
ond edition of his Coleoptera collection’s catalogue, the first name, by not being accompanied with descriptions, could not
published after Lacordaire’s return, listed on page 143, among be accepted, concluded that Lacordaire’s species had never been
several dung beetle groups, the new genus ‘Gromphas. Dejean’, formally named. To fix the situation, he established the name
with the new species ‘Lacordairei Dej.’, from ‘Buenos-Ayres’ Gromphas inermis Harold, 1869 for it. As for the authorship
(Fig. 2). Because this is the only Buenos Aires scarabaeine of Gromphas, Harold (1869a) credited it to Brullé, the first
named by Dejean after Lacordaire and, moreover, because to describe the genus.
it belongs to a new genus, it is obvious that this is the new 9) Burmeister (1874) disagreed with Harold (1869b) and
species that Dejean named after Lacordaire that Lacordaire considered Dejean’s names Gromphas and Gromphas lacordai-
referred to in 1830. No description or diagnosis was provided rii (his spelling) available from the latter’s 1830s collection
by Dejean for either the genus or the species. catalogues. By being senior to both Blanchard’s G. lacordairei
3) A review of Lacordaire’s memoir was published by Lor- and Harold’s G. inermis, G. lacordairii Dejean was, in Bur-
enz Oken (1834) (Figs 1; 2). Oken enumerated the species meister’s view, the proper valid name of Lacordaire’s species,
addressed by Lacordaire one by one; in a paragraph on the with G. inermis as its junior synonym, whereas Blanchard’s
Coprobius, besides several names mentioned for this genus G. lacordairei was a junior synonym of Perty’s G. aeruginosa.
by Lacordaire, Oken (1834: 1117) also included ‘lacordairii, 10) A period of instability reigned during the late 19th
digs deep; arachnoides, like spider in cow dung’ (‘Coprobius: and the early 20th centuries, when authors transitioned from
[…] lacordairii, gräbt tief; arachnoides, wie Spinne im Kuh- using Harold’s G. inermis (e.g., Preudhomme-de-Borre 1886;
mist’) (Fig. 2). Though Lacordaire expressly explained that Judulien 1899; Gahan & Arrow 1903; Heyne & Taschenberg
they belonged to new genera, not to Coprobius, it is clear 1908), to ‘G. lacordairei Dejean’ (e.g., Tremoleras 1910), and
from the context and the associated biological and morpho- then to ‘G. lacordairei Brullé’ (e.g., Bruch 1911) as the valid
logical information that the two species referred to by Oken name of Lacordaire’s Buenos Aires species. With the adop-
as Coprobius lacordairii and C. arachnoides are, respectively, tion of the latter name and authorship by Gillet’s (1911) and
Lacordaire’s new Buenos Aires species and his Ateuchus arach- Blackwelder’s (1944) catalogues and by d’Olsoufieff’s (1924)
noides from Tucumán. landmark revision of the phanaeines, a consensus seemed to

26 ZOOSYSTEMA • 2024 • 46 (2)

 A B

C D

Fig. 1. — Authors in the early taxonomic history of Gromphas: A, Jean Théodore Lacordaire (1801-1870), the Belgian-French entomologist who collected in
Argentina the first known specimens of the genus. He was also the first to record aspects of its behaviour in the literature (Lacordaire 1830). From an engraving
reproduced by Candèze (1872); B, Pierre François Marie Auguste Dejean (1780-1845), who recognised that the specimens collected by Lacordaire belonged to a
then-new genus and species, which he named ­– first as a nomen in litteris and later as a nomen nudum (Dejean 1833) – ‘Gromphas lacordairei’. From Wikipedia;
C, Lorenz Oken (1779-1851), a leader of the Naturphilosophie movement in Germany and editor of the journal Isis. In a 1834 literature review for the journal, Oken
combined the morphological and behavioural information introduced by Lacordaire (1830) with the specific name coined by Dejean (1833) and made the name
Coprobius lacordairii available. From Ecker (1883); D, Gaspard Auguste Brullé (1809-1873), who has been erroneously regarded as the author who made both
Gromphas and ‘Gromphas lacordairei’ available. From Wikipedia.

27
 Cupello M.

Lacordaire (1830)

Dejean (1833)

Dejean (1836) Oken (1834)

Fig. 2. — The early taxonomic literature on Gromphas Dejean, 1836 and G. lacordairii (Oken, 1834). Lacordaire’s (1830) was the first publication where the spe-
cies and genus were ever mentioned (green underline), and where a description was provided, though their names were not given. Notice that the discussion on
this new genus and species follows four paragraphs on Coprobius (red underlined). In the second edition of his catalogue, Dejean (1833) finally mentioned the
names, but did not accompany them with either a description, a diagnosis, or an indication. Oken (1834), in his review of the papers appearing in several volumes
of Annales des sciences naturelles, including Lacordaire’s, listed the name lacordairii followed by a short description (‘digs deep’), but erroneously assigned it to
Coprobius Latreille, 1829. This short description, plus the indication of Lacordaire’s paper and expanded description, made the species-group name Coprobius
lacordairii available under Oken’s (1834) authorship. When Dejean (1836), in the third edition of his catalogue, combined it with the then-nomen nudum Gromphas,
he made the latter available and lacordairii its type species. See the text for more details.

have been reached and all authors until 2013 adopted ‘G. la- 11) In their revision of Gromphas, Cupello & Vaz-de-Mello
cordairei Brullé’ as the valid name and treated G. inermis as (2013) argued that Brullé cannot be deemed the author who
its invalid junior synonym. The authorship of Gromphas was gave availability to G. lacordairei for he did not mention the
likewise consistently attributed to Brullé. And though never name in his work. They considered that Blanchard (1846)
expressly addressed in the literature, it is clear that ‘G. lacor- had been the first author to use G. lacordairei in a way that
dairei Brullé’, having been supposedly established in the same complies with the availability criteria of the Code and that,
work as Gromphas, was tacitly regarded as the type species. therefore, the authorship of the name would be his. How-

28 ZOOSYSTEMA • 2024 • 46 (2)

ever, as Blanchard’s name referred to an Amazonian species Analysis of previous history
(namely, G. aeruginosa), it was not available to denote the Now aware of Lacordaire’s (1830) and Oken’s (1834) texts,
species then known as ‘G. lacordairei Brullé’ (i.e., Lacordaire’s I challenge some of the previous interpretations:
Buenos Aires species). The oldest available name for the latter 1) Because Lacordaire (1830) mentioned neither the ge-
was, as far as they were aware, G. inermis. Hence, Cupello & neric name nor the full binomen of his new Buenos Aires
Vaz-de-Mello adopted this name for Lacordaire’s Buenos species, he did not make either available. Nevertheless, by
Aires species in all of their papers dealing with Gromphas listing three taxonomic characters ‒ viz., colouration, size,
(Cupello & Vaz-de-Mello 2013, 2014, 2015). Moreover, and digging behaviour ‒ and the respective states shown by
because Burmeister (1874) explicitly said that he was using the species ‒ viz., metallic, ‘large’, and ‘digs deep holes in the
‘G. lacordairii Dejean’ as a different name from ‘G. lacord- earth like Copris but unlike any Ateuchus’ ‒, he presented what
airei Blanchard’, Cupello & Vaz-de-Mello interpreted that the Code would term a description or definition, however
he also established a new available name under his own brief it might be considered (the Code cites ‘taxonomic char-
authorship, G. lacordairii Burmeister, 1874, a junior syno- acters’ in the definition of ‘description’, but does not explain
nym of G. inermis Harold, 1869. As for the authorship of what the term means, only plain ‘character’; I follow Mayr
Gromphas, since Cupello & Vaz-de-Mello believed that Brullé et al.’s [1953], Mayr’s [1969], and Mayr & Ashlock’s [1991]
had been the first to publish the name in a way to comply definition). When, four years later, Oken (1834: 1117), in a
with all the availability criteria of the Code, they agreed review of Lacordaire’s paper, used the name Coprobius lacor-
with the consensus of the time and attributed it to Brullé. dairii to refer to this new species of Lacordaire’s, he made the
Since no nominal species had been originally included in the name available by a) using it as valid (complying with Article
genus, Cupello & Vaz-de-Mello initially designated one of 11.5), b) accompanying it with a rather short description
the two nominal species subsequently assigned to the genus (‘it digs deep’, ‘gräbt tief’) (Art. 12.1), and c) accompanying
by Blanchard (1846), G. dichroa, as the type species. Later, it with an indication, namely, an indication to Lacordaire’s
however, they came across Sturm’s (1843) earlier inclusion of text he was expressly reviewing, where the slightly expanded
Onitis aeruginosus in the genus and interpreted that this fixed description can be found (Art. 12.2.1). Therefore, the oldest
it as the type species by subsequent monotypy (Cupello & name available for Lacordaire’s Buenos Aires species is neither
Vaz-de-Mello 2014). ‘Gromphas lacordairei Dejean, 1833 [or 1836]’, ‘Gromphas
12) The following taxonomy and nomenclature resulted lacordairii Dejean’, ‘Gromphas lacordairei Brullé, 1837 [or
from all this activity and have until now been considered 1834, or 1838]’, nor G. inermis Harold, 1869, but Coprobius
valid (only names so far discussed and deemed available lacordairii Oken, 1834.
included): 2) Since every author who has ever used the name Grom-
phas lacordairei, regardless of the spelling employed and the
Gromphas Brullé, 1837 recognized taxonomic limits of the species, has always referred
[sic; see Bousquet (2016) for the correct publication year of directly or indirectly to the name coined by Dejean and, thus,
Brullé’s work, 1838]. to the specimens collected by Lacordaire, they were all refer-
ring to the same nominal species. For this reason, no one has
Type species. — Onitis aeruginosus Perty, 1830, by subsequent established new homonyms ‘lacordairei/lacordairii’ in Gromphas,
monotypy by Sturm (1843: 108).
and no separate available name ‘G. lacordairei Blanchard’ or
‘G. lacordairii Burmeister’ exists as Cupello & Vaz-de-Mello
Gromphas aeruginosa (Perty, 1830) (2013) argued. What we actually have is, in retrospect, in
Blanchard’s case, an erroneous attribution of the authorship of
Onitis aeruginosus Perty, 1830: 39, pl. 8, fig. 8. the name (to Dejean) and identification (i.e., his application
Gromphas aeruginosa – Sturm 1843: 108. to specimens belonging to an Amazonian species, G. aerugi-
nosa), and, in Burmeister’s, only an erroneous attribution of
Gromphas lacordairei Blanchard, 1846: 181 (subjective synonymy the authorship (to Dejean). Using Smith & Smith’s (1972) apt
firstly established by Harold (1859). term, ‘lacordairei Blanchard’ and ‘lacordairii Burmeister’ are
chresonyms, not homonyms, of Coprobius lacordairii Oken,
1834 (hetero- and orthochresonyms, respectively; see Dubois
Gromphas inermis Harold, 1869 2000), and thus have no separate availability. To make it clear:
they are simple subsequent citations of the name originally
Gromphas inermis Harold, 1869b: 62. made available by Oken, not separate available names in
Gromphas lacordairii Burmeister, 1874: 130 (objective synonymy their own right. They must be accordingly removed from the
firstly recognised by Cupello & Vaz-de-Mello [2013]). synonymic list of the Gromphas species.
3) When Dejean (1836) listed ‘lacordairei Dej.’ under
Gromphas in the third edition of his catalogue, he was using
Gromphas dichroa Blanchard, 1846 the same species-group name that i) had been indirectly men-
tioned as a nomen in litteris by Lacordaire (1830); ii) appeared
Gromphas dichroa Blanchard, 1846: 182. as a nomen nudum in the second edition of his own catalogue

29
 Cupello M.

(Dejean 1833); and iii) had been made available two years I was unable to recognise any Bakewell specimens in the
earlier by Oken (1834). Hence, by including an already avail- Gromphas material of the MNHN, or Dejean scarabs in general.
able name under Gromphas, Dejean (1836) complied with As for any potential La Ferté Gromphas, the only specimen
the availability criterion for genus-group names established I located there is the G. lemoinei Waterhouse, 1891 depicted
in Article 12.2.5 and made Gromphas available. The author- in Figure 4. But because this species lives only in Colombia
ship of this name does not belong to Brullé (1838), after all, and Venezuela (Cupello & Vaz-de-Mello 2013, 2015) (Fig. 5),
but to Dejean (1836), something rather fitting since it was this specimen cannot be part of the series collected by Lacor-
indeed Dejean who coined the name. daire in Buenos Aires and later acquired by Dejean. As for the
4) Coprobius lacordairii (currently, Gromphas lacordairii) NHM, sometimes it is relatively easy to identify Bakewell’s
is the type species of Gromphas Dejean, 1836 by original Dejean scarabs relying on Dejean’s labelling style (see Horn
monotypy, since, as just explained, this was the only nomi- et al. 1990b for an example). Unfortunately, however, I found
nal species included in Gromphas (through an indication to no Dejean or Bakewell Gromphas in the museum.
its name) when the genus-group name was made available. Two further possibilities explored by me were the Oxford Uni-
5) The type series of Coprobius lacordairii is composed versity Museum of Natural History, Oxford (OUMNH), and
of the specimens observed and collected by Lacordaire in the Museo Regionale di Scienze Naturali, Turin, Italy (MRSN).
Buenos Aires, for they were the ones which provided the Drawing from letters exchanged between the two entomologists
characters mentioned in his 1830 description indicated in the early 1820s, it is known that Frederick William Hope
by Oken (1834) and it is clear that Oken himself did not (1797-1862) received, in 1823, around 50 Coleoptera from
see additional specimens. Where are these specimens? My Dejean (Smith 1986). Even though syntypes of C. lacordairii
conclusion is that they are now lost. Lacordaire’s Coleoptera cannot be among these c. 50 specimens given that Lacordaire
collection was sold in parts through the years, and some only came back to Europe in 1830, it is possible that Dejean
of its material is currently housed in the Paris (MNHN), and Hope continued to exchange material over the years. All the
Brussels (RBINS), and London (NHM) museums (Horn specimens received by Hope, if still surviving, must be in the
et al. 1990b; Cambefort 2006). Unfortunately, nothing is OUMNH, where the Hope-Westwood collection is preserved.
known particularly about the whereabouts of Lacordaire’s Unfortunately, none of the Gromphas that Cupello & Vaz-de-
scarabs, nor did I find in any of the searched museums a Mello (2015) found there bears any link to either Dejean or
specimen that could have belonged to Lacordaire’s Buenos Lacordaire. Three of them are from areas other than Buenos
Aires series (see the list of collections studied in the Material Aires ‒ two from Brazil and one from Bolivia ‒ another five
and Methods). The specimen illustrated in figure 4, plate have no provenance data, and none is from Argentina.
27, of a later work by Lacordaire (1855) (Fig. 3B), which As for the Turin museum, the Dejean material housed
belongs to the species nowadays known as G. inermis sensu there originates from the private collection of Massimiliano
Cupello & Vaz-de-Mello, is not part of that series, for La- Spinola (1780-1857) and, except for some Chrysomelidae
cordaire (1855) mentioned in the ‘explanation’ on page 10 still preserved in original Dejean boxes, is now scattered in
that this specimen came from Tucumán (Argentina), not the general collection among specimens from several other
Buenos Aires (Fig. 3A). sources (Giachino 1982). Box 94 of the general collection
In addition to the syntypes that may have been housed contains two cabinet labels reading ‘Gromphos [sic] lacord-
in Lacordaire’s collection, there were also those deposited airei’, one of them ascribing the authorship of the name to
in Dejean’s. We know they existed because, as we have Dejean and standing beside two specimens, the other citing
seen, Dejean listed the species as present in his collection no author and standing beside a single specimen (Giachino
in his 1830s catalogues (Dejean 1833, 1836) (but not 1982). However, because both labels state Brazil as the place
in the 1821 first edition, preceding Lacordaire’s South of origin of the species, it is unlikely that the specimens as-
American travel) and indicated their provenance precisely sociated with them came from Dejean’s material and certainly
from the place where Lacordaire collected them, Buenos not from Lacordaire.
Aires. Like Lacordaire’s, Dejean’s beetle collection was sold Having failed to find the specimens, I deem the type series
in parts in 1840 and scattered across Europe (Horn et al. lost. But is the allocation of the name Coprobius lacordairii
1990a; Cambefort 2006; Bousquet & Bouchard 2013; (and, by extension, of Gromphas) affected by that? It has been
Maldaner et al. 2017). His scarabs seem to have been split traditionally considered Lacordaire’s specimens belonged to
mainly between François Thibault de la Carte, marquis de the species currently known as Gromphas inermis (see Cu-
La Ferté-Senectère (1808-1886), in France, and Robert pello & Vaz-de-Mello 2013). Arguably, the earliest indication
Bakewell (1810-1867), in Britain. Eventually, La Ferté’s that they did belong at least to the genus Gromphas as the
specimens were mostly (i.e., excluding the Cetoniinae) in- name is currently applied is from Brullé (1838: 298), who
corporated into the MNHN via the Oberthür collection in expressly ascribed to this genus ‒ but, as we have seen, without
1952 (Cambefort 2006), whereas Bakewell’s were divided mentioning its name ‒ the species that Lacordaire observed
in 1867, with one part going to the NHM that same year, digging deep holes below horse dung.
while the other (apparently a ‘second choice’) ended up Yet, the best evidence regarding the identity of Lacordaire’s
at the MNHN via the Oberthür collection along with La specimens is found in a work by Lacordaire himself (Lacor-
Ferté’s (Horn et al. 1990a; Cambefort 2006). daire 1855). In this work, Lacordaire recognised (though

30 ZOOSYSTEMA • 2024 • 46 (2)

 A

Fig. 3. — Lacordaire’s (1855) illustration of ‘Gromphas lacordairei Blanchard’. Note that in the ‘explanation’ (i.e., the captions) of his plate 27 (A), Lacordaire indi-
cated Tucumán, Argentina, as the locality of origin of G. lacordairei. I assume he was referring to the provenance of the individual specimen figured in his plate
(B), not of the species as a whole, since, in the text, he said that the species ranged from Peru to Argentina. Being from Tucumán, the specimen cannot be part
of the series collected by Lacordaire in Buenos Aires and that later became the type series of G. lacordairii (Oken, 1834).

hesitantly) two species for Gromphas, G. dichroa Blanchard, (1855) plate 27 as ‘Gromphas lacordairei Blanch.’ (Fig. 3), his
1846 and ‘G. lacordairei Blanchard’, the latter said to be the Buenos Aires specimens must indeed have belonged to G. in-
one he discovered in Buenos Aires. Since the only Gromphas ermis. That said, Lacordaire’s (1855) text clearly shows that
species other than G. dichroa to occur in Buenos Aires is G. in- what he considered to be ‘G. lacordairei’ (i.e., his taxonomic
ermis sensu Cupello & Vaz-de-Mello (2013, 2015), and, as species ‘G. lacordairei’) also included specimens of G. aer-
said above, this is clearly the species depicted in Lacordaire’s uginosa. This is evident because he says that the distribution

31
 Cupello M.

Fig. 4. — The only specimen of Gromphas from the La Ferté collection. This specimen, a G. lemoinei Waterhouse, 1891, like the majority of the La Ferté speci-
mens, is now preserved in the MNHN. Since La Ferté owned specimens from the Dejean collection, one could suggest that this specimen might be one of the
syntypes of G. lacordairii (Oken, 1834). But this is not the case. Gromphas lemoinei is endemic to Colombia and Venezuela, while the syntypes were all collected
in Buenos Aires, Argentina. Photographs courtesy of Christophe Rivier (MNHN). Scale bar: 5 mm.

of ‘G. lacordairei’ reaches Peru and males are different from 7) The complete revised nomenclature of the genus Grom-
females in bearing cephalic and pronotal horns. It is evident phas and its species is as follows (see the list of references for
that he mistook real G. inermis/G. lacordairii (both sexes details on the publication dates).
hornless and lacking in Peru) for females and G. aeruginosa
specimens (both sexes horned and occurring in Peru) for Revised nomenclature of the genus Gomphras
males of a single species, a common misconception among
coleopterists working until the mid-19th century (Cupello & Gromphas Dejean, 1836
Vaz-de-Mello 2013). But since G. aeruginosa does not occur
in Argentina, there are no doubts that all the specimens he Gromphas Dejean, 1836: 159 [30th July 1836].
collected in Buenos Aires did belong to G. inermis. As the
Type species. — Coprobius lacordairii Oken, 1834, by original
identity and geographical provenance of the lost syntypes are monotypy.
unambiguous, the allocation of the name is straightforward
and no neotype designation is necessary or justified under Etymology. — From the Latinisation of the Ancient Greek noun
Article 75.3 of the Code. γρομφάς (gromfás), meaning ‘an old sow’, feminine in gender (Brullé
6) Having concluded that the syntypes of G. lacordairii Oken, 1838; Harold 1869a; Bailly 1895; Liddell & Scott 1897; Brown
1954; Cupello & Vaz-de-Mello 2013). The name likely draws a
1834 belong to the same species as the lectotype designated parallel between the stout, hornless body shape of both pigs and the
by Cupello & Vaz-de-Mello (2013) for Gromphas inermis only species originally included in the genus, G. lacordairii, while
Harold, 1869, the two names are now subjective synonyms, also making a probable reference to the ‘filthy’ coprophagous habits
with the former being valid due to its 35-year seniority. of these dung beetles.

32 ZOOSYSTEMA • 2024 • 46 (2)

 N

VENEZUELA

COLOMBIA

ECUADOR

PERU
BRAZIL

BOLIVIA

South Atlantic Ocean

South Pacific Ocean

PARAGUAY

Gromphas Dejean, 1836

G. lemoinei Waterhouse, 1891

G. aeruginosa (Perty, 1830)

URUGUAY G. amazonica Bates, 1870

G. jardim Cupello &
Vaz-de-Mello, 2015
ARGENTINA
G. lacordairii (Oken, 1834)
1000 km
G. dichroa Blanchard, 1846

Fig. 5. — Updated distribution map of the six species of Gromphas. Data retrieved from Cupello & Vaz-de-Mello (2013, 2015), Damborsky et al. (2015), Cajaiba
et al. (2017), Gámez & Acconcia (2018), Alonso et al. (2019), Uchoa & Rodrigues (2019), Bitencourt et al. (2019), Noriega et al. (2021), and the material examined
for this work.

Gromphas aeruginosa (Perty, 1830) ish’, ‘verdigris green’ or ‘dark green’ (Lewis & Short 1891; Brown
1954; Glare 1968-1982; Papavero 1994). Though not originally
Onitis aeruginosus Perty, 1830: 39, pl. 8, fig. 8 [31st December 1830]. explained, the name makes an obvious reference to the greenish
metallic colouration of many individuals of the species, including
Gromphas aeruginosa – Sturm 1843: 108. those in the type series, a characteristic that Perty (1830) described
Name-bearing type. — Lectotype (male), designated by Scherer as ‘obscure viridi-metallicus’.
(1983: 298), ZMS. Distribution. — The Amazon Basin in Colombia, Ecuador, Peru,
Type locality. — Brazil: the Amazon rainforest, somewhere visited Brazil, and Bolivia, especially in sandbanks, floodplains and natural
by Spix and Martius along the Amazon River, the Japurá, the lower savannas, with southern incursions into the dry Chiquitano forests
Madeira, or the Rio Negro (see Papavero 1971 for a map with these of eastern Bolivia and the Bañados de Izozog wetlands (‘Río Para-
places indicated) (see discussion below). petí’) of the southern Bolivian Dry Chaco. The species is also known
from a sole locality in the upper Magdalena River Valley, the town
Etymology. — A Latin first-class adjective in the nominative case of Gigante (Huila, Colombia), separated from the Amazon by the
meaning ‘rusty’, ‘covered with verdigris’, and, by extension, ‘green- Cordillera Oriental, the eastern branch of the Colombian Andes. It

33
 Cupello M.

is unclear whether this population is somehow connected to those Distribution. — Open environments of middle South America in
in the Amazon across the Andes or represents a geographical isolate. Brazil, Bolivia, Paraguay, Argentina, and Uruguay, inhabiting savan-
If the latter, then the question is raised as to whether this Gigante nas, grasslands, floodplains, river sandbanks, pastures, and restingas
population is a relict from a time when the species was more widely as well as associated forest edges and gallery forests.
ranged and skirted the Cordillera Oriental or whether it is somehow
the result of jump dispersal across the Andean mountains.
Gromphas dichroa Blanchard, 1846

Gromphas lacordairii (Oken, 1834) Gromphas dichroa Blanchard, 1846: 182 [18th December 1846].
Name-bearing type. — Holotype by monotypy (female), MNHN.
Coprobius lacordairii Oken, 1834: 1117 [30th November 1834].
Type locality. — Uruguay: Montevideo, ‘near the sea’ (Blanchard
Gromphas inermis Harold, 1869b: 62 [1st November 1869], n. syn. 1846).
Gromphas lacordairii bipunctata d’Olsoufieff, 1924: 59 [31st De- Etymology. — A New Latin first-class adjective in the nomina-
cember 1924], n. syn. [subjective synonymy with G. inermis firstly tive case meaning ‘having two colours’, ‘bicoloured’ (Wiktionary
established by Cupello & Vaz-de-Mello (2013: 464); here extended 2022). The word ultimately derives from the combination of the
to G. lacordairii (Oken, 1834)]. Ancient Greek prefix δῐ- (di-), meaning ‘twice’, ‘double’ (Brown
1954; Wiktionary 2023a), and the Ancient Greek noun χρῶμᾰ
Name-bearing type. — Coprobius lacordairii: syntypes (unknown (khrôma), meaning ‘colour’, ‘pigment’, especially of the skin or
sexes), unknown whereabouts, likely destroyed; allocation remains body surface (Bailly 1895; Liddell & Scott 1897; Brown 1954),
unambiguous (see the text above). Latinized by the addition of the Latin adjectival suffix -us (-a, -um)
Gromphas inermis: lectotype (male), designated by Cupello & Vaz- (Wiktionary 2023b). The name makes an obvious reference to the
de-Mello (2013: 464), MNHN (ex Edgar von Harold and René dorsal colouration of the holotype, with a centrally red pronotum
Oberthür collections). and blue elytra, a pattern seen in many ‒ but not all ‒ individuals
Gromphas lacordairii bipunctata: lectotype (male), designated by of the species (Cupello & Vaz-de-Mello 2013, 2015; see more de-
Cupello & Vaz-de-Mello (2013: 464), MNHN (ex René Oberthür tails later in this work). Blanchard (1846) called the species in his
collection). vernacular French the ‘gromphas bicolore’.
Type locality. — Coprobius lacordairii and Gromphas inermis: Distribution. — The Pampas biome and peripheral areas of the
Argentina: either somewhere in the province of Buenos Aires or southern Atlantic Forest (e.g., Itapiranga, Santa Catarina, and Nova
the adjacent city of Buenos Aires. Petrópolis, Rio Grande do Sul) in southern Brazil, Argentina, and
Gromphas lacordairii bipunctata: Brazil: Mato Grosso. Uruguay; possibly also present in the southern Humid Chaco of
Paraguay. Nothing is known about the habitats occupied by the
Etymology. — Coprobius lacordairii: the masculine singular genitive species in this vast area. One possibility, based on what is known
of the proper noun Lacordairius, the Latinised form of the French of the biology of the other Gromphas, is that the species occupies
surname Lacordaire. It is an eponym after Belgian-French natural- more humid areas, especially wetlands, including the floodplains
ist Jean Théodore Lacordaire (1801-1870), the collector of the type and sandbanks of the Uruguay River along the border of Argentina
material. The name is spelled as ‘lacordairii’, and not ‘lacordairei’ as and the Brazilian state of Rio Grande do Sul and other such water
originally intended by its coiner Dejean (1833, 1836) and so far cited bodies like Lake Guaíba (e.g., in Porto Alegre) and the River Plate
in most of the literature, because Oken derived it not directly from (e.g., Montevideo and Buenos Aires).
Lacordaire’s name in modern French as Dejean did (i.e., the name
Lacordaire + the Latin masculine genitive suffix -i), but, as said, from
its Latinized form, Lacordairius (i.e., by adding the masculine suffix -i Gromphas amazonica Bates, 1870
to the stem Lacordairi-; see Article 31.1.1 of the Code for more details).
The ‘lacordairei’ spelling is regarded here as an incorrect subsequent Gromphas amazonica Bates, 1870: 175 [30th June 1870].
spelling in the terms of Article 33.3 of the Code; even though it is
in prevailing usage, because this spelling has never been attributed to Name-bearing type. — Lectotype (female), designated by Cupello &
the publication of the original spelling (i.e., to Oken 1834), it cannot Vaz-de-Mello (2013: 462), MNHN (ex Henry Walter Bates and
be deemed a correct original spelling under Art. 33.3.1. René Oberthür collections).
Gromphas inermis: A Latin adjective in the nominative case mean-
ing ‘unarmed’, ‘having no weapons’ (Lewis & Short 1891; Brown Type locality. — Brazil: Amazonas: Tefé.
1954; Glare 1968-1982), a reference to the hornless condition of
this species in contrast to the horned congeneric G. aeruginosa. Etymology. — A New Latin first-class adjective in the nominative
Gromphas lacordairii bipunctata: A New Latin first-class adjective case meaning ‘Amazonian’, ‘from the Amazon rainforest’ (Wiktion-
in the nominative case meaning ‘bi-punctate’, ‘bearing two punc- ary 2019b). A toponym after the place famously explored by H. W.
tures’, the combination of the Latin prefix bi- for ‘two’, ‘occurring Bates between 1848 and 1859 (Bates 1863; Papavero 1973), where
twice’, and the New Latin adjective punctatus (-a, -um) for ‘punctate’ he collected the type series and where the species is endemic.
(Brown 1954; Wiktionary 2019a). It alludes to the pair of tenuous Distribution. — Floodplains and sandbanks of the Amazon and
posterior pronotal fossae that characterized the purported variety of some of its upper tributaries and headwaters, namely the Juruá,
G. lacordairii for which d’Olsoufieff (1924) established the name and Huallaga, Ucayali, and Pisqui Rivers, across Peru, the southern tip
which he knew exclusively from Mato Grosso (Brazil). Cupello & of Colombia, and Brazil.
Vaz-de-Mello (2013) observed that specimens bearing this feature
are always small and actually found throughout the range of the
species, always in polymorphism with the much more abundant
smooth individuals of all sizes. Due to this, and in keeping with Gromphas lemoinei Waterhouse, 1891
the authors’ taxonomic criteria, they were prompted to invalidate
d’Olsoufieff’s taxon and name. Gromphas lemoinei Waterhouse, 1891: 60 [1st July 1891].

34 ZOOSYSTEMA • 2024 • 46 (2)

Name-bearing type. — Holotype by monotypy (unknown sex), and five females. Their labels are transcribed below (italics for
NHM. handwriting, slashes for line breaks). As Scherer (1983) already
Type locality. — Venezuela: La Guaira: La Guaira. noted, none of the former syntypes bears original labels in
Perty’s handwriting, those having been replaced by printed
Etymology. — The masculine singular genitive of the proper noun labels by later curators (Scherer 1983).
Lemoine, a French surname meaning ‘the monk’ (‘Le Moine’).
Waterhouse (1891) did not explain after whom this eponym was
As explained by Cupello & Vaz-de-Mello (2013), the
formed, but I suspect it may have been the French entomologist geographical provenance pointed out by Perty (1830) for
Victor Lemoine (b. ?-1897), the only Lemoine who I found associ- O. aeruginosus, the then Brazilian provinces and now states of
ated with entomology in the late 19th century. Alas, apart from the São Paulo and Minas Gerais, is certainly wrong, as the species
fact that he was based in Paris and was a pioneer in the application is exclusively Amazonian. The provenance indicated on the
of radiography in the study of arthropods (Lemoine 1897), almost
nothing is known about Victor Lemoine (Anonymous 1897; Con-
labels of the type specimens, ‘Brasilien’ or ‘Brasilia’, is not of
stantin 2012; Groll 2016). great help either for it is already widely known (and, indeed,
stated in the work itself ) that Perty’s (1830) book was based on
Distribution. — Open savannas, pastures and river sandbanks in material collected exclusively in Brazil during Johann Baptist
the Colombian and Venezuelan Llanos and associated areas.
von Spix and Karl Friedrich Philip von Martius’ expedition
across the country (1817-1820). But where exactly in Brazil
Gromphas jardim Cupello & Vaz-de-Mello, 2015 did the naturalists find the species? Studying the itinerary of
Spix and Martius’ journey, especially the list of places visited
Gromphas jardim Cupello & Vaz-de-Mello, 2015: 4 [12th October by them in the Amazon, can at least limit the possibilities. In
2015; see discussion below].
addition to other areas of Brazil, the two naturalists visited
Name-bearing type. — Holotype by original designation (male), together numerous Amazonian localities along the Amazon
OUMNH. River, from its mouth in the Atlantic Ocean upstream to
Type locality. — Bolivia: Beni: Moxos, Río Ichiguita, 155 m, Benjamin Constant, on the border of Brazil, Colombia, and
15°08’S, 65°18’W, savanna habitat. Peru, as well as along the lower Madeira River (Spix & Martius
1823-1831; Papavero 1971). They also explored separately
Etymology. — The masculine singular nominative of the proper the lower c. 420 km of the Amazon’s main left tributary, the
noun Jardim, a Portuguese surname meaning ‘garden’, used in ap-
position. It is an eponym after my beloved maternal grandfather and Rio Negro, from Manaus to Barcelos (Spix), and the entire
Brazilian airline pilot Arlindo da Silva Jardim (1923-2014), who Brazilian extension of another mighty left tributary, the
died in the same week I realised this species was new while examin- Japurá (Martius). Gromphas aeruginosa is abundant in many
ing specimens at the Oxford and London museums. of these places (Cupello & Vaz-de-Mello 2013, 2015), and
Distribution. — Open savannas, pastures and perhaps river sand- the type series, and particularly relevant for the type locality,
banks in the Beni Savanna of Bolivia (in the Mamoré River Basin) the lectotype, can have been collected in any of them. Fol-
and the western Brazilian Cerrado (on the northern fringes of the lowing Recommendation 76A.2 of the Code, I correct Perty’s
Pantanal wetlands). erroneous type locality statement (‘Habitat in mediterraneis
Prov. S. Pauli et Minarum’) to ‘Brazil: the Amazon rainforest,
somewhere visited by Spix and Martius along the Amazon
ON THE TYPE SERIES OF GROMPHAS AERUGINOSA River, the Japurá, the lower Madeira, or the Rio Negro’.
(PERTY, 1830)
Type material examined. — Lectotype • ♂; ZSMC; designated
by Scherer (1983: 298): green disk, ‘Brasilien’, ‘Type von / grom-
For our revision of Gromphas (Cupello & Vaz-de-Mello 2013), phas / aeruginosa / Perty’ [Hans Kulzer’s handwriting], ‘Type’, ‘alte
Vaz-de-Mello and I could examine the lectotype of Onitis / Sammlung’, ‘HOLOTYPUS / Onitis Perty / aeruginosus / det.
aeruginosus Perty, 1830 (i.e., G. aeruginosa) only through Dr.G.Scherer 1981’ [Scherer’s handwriting], ‘Gromphas / aeruginosus
photos provided by Michael Balke, the Coleoptera curator / (Perty) / det. G.Scherer, 1981’ [Scherer’s handwriting].
at the Zoologische Staatssammlung München, Munich, Ger- Paralectotypes • ♂; ZSMC: green disk, ‘Brasilien’, ‘a te / Samm ung’
[sic; ‘l’ in both ‘Alte’ and ‘Sammlung’ faded], ‘Lecto- / PARATYPUS
many (ZSMC), where the specimen has been housed since / Onitis / aeruginosus / Perty / Dr.G.Scherer, 1981’ [Scherer’s hand-
its arrival in Europe in the early 19th century (Scherer 1983; writing] • ♂; ZSMC: ‘Lecto- / PARATYPUS / Onitis / aeruginosus
Cupello & Vaz-de-Mello 2013). The lectotype was designated / Perty / Dr. G.Scherer, 1981’ [Scherer’s handwriting] • ♂, ZSMC:
by Scherer (1983) from a series of eight syntypes, all of which green disk, ‘Brasilien’, ‘alte / Sammlung’, ‘Lecto- / PARATYPUS /
I was able to locate during a three-day visit to the ZSMC Onitis / aeruginosus / Perty / Dr.G.Scherer, 1981’ [Scherer’s handwrit-
ing] • ♂; ZSMC: green disk bordered in black, ‘Brasilia’ [unknown
in November 2019. As originally suspected by Cupello & handwriting], ‘alte / Sammlung’, ‘Lecto- / PARATYPUS / Onitis /
Vaz-de-Mello (2013), Scherer (1983), like Perty (1830) be- aeruginosus / Perty / Dr.G.Scherer, 1981’ [Scherer’s handwriting] •
fore him, mistakenly took every major specimen (i.e., those ♂, ZSMC: green disk bordered in black, ‘alte / Sammlung’, ‘Lecto- /
with well-developed cephalic and pronotal horns) for males PARATYPUS / Onitis / aeruginosus / Perty / Dr.G.Scherer, 1981’
and every minor one for females. His information that the [Scherer’s handwriting] • ♂; ZSMC: green disk, ‘Brasilien’, ‘alte
/ Sammlung’, ‘Lecto- / PARATYPUS / Onitis / aeruginosus / Perty
type series was composed of four males and four females was / Dr.G.Scherer, 1981’ [Scherer’s handwriting] • ♂; ZSMC: ‘9. /
consequently bound to be incorrect; in fact, there are three Brasilia / O. ♂ ♀ / aerugino- / sus / P.’ [unknown handwriting;
males (including the lectotype, as correctly sexed by Scherer) same style as Max Gemminger’s labels as shown in Scherer (1983)],

35
 Cupello M.

‘Lecto- / PARATYPUS / Onitis / aeruginosus / Perty / Dr.G.Scherer, described G. inermis in a work dealing specifically with the
1981’ [Scherer’s handwriting]. material that he had seen during a visit to the MNHN (see
Additional non-type material examined. — Bolivia. Cocha- Cupello 2020), and the specimen bears a label identifying it
bamba • 1 ♂; [genitalia dissected], 1 ♀; Carrasco, Puerto Villarroel, in Harold’s handwriting as G. inermis (Fig. 7). Another label
Valle Sacta; 17°07’S, 64°45’W; 230 m; 9.V.2000; M. Aliaga leg.; borne by the specimen, a rectangular, green one typical of
human faeces trap; Amazon Forest; TAMU. South American specimens collected by Alcide d’Orbigny
Santa Cruz • 1 ♂; 1 ♀; Andrés Ibáñez, Lomas de Arena; 17°55.360’S,
63°09.637’W; 413 m; 5.XII.2008; W. D. Edmonds and T. Vidaurre
housed in the MNHN, reads ‘G. aeneus, / Blanch. / Corrientes
leg.; human faeces; forest remnant; TAMU • 3 ♂, 3 ♀; Ichilo, Buena / M. D’Orbigny.’. This nomen in litteris was curiously never
Vista; 380 m; XI.1971; F. Steinbach leg.; TAMU • 1 ♂; Ichilo, Buena mentioned by Blanchard (1846) in his study of the d’Orbigny
Vista; 17°27’49”S, 63°35’58”W; 360 m; no date; W. D. Edmonds South American material or anyone else in the literature except
leg.; horse dung; pasture; TAMU. for Harold (1869b), who cited it as one of the (to use modern
Brazil. Acre • 1 ♀; Cruzeiro do Sul; VI.1952; J. Correia leg.; TAMU
• 1 ♂, 3 ♀; same data as for the preceding; ex Alvarenga collection; jargon) rejected unavailable synonyms of G. inermis along with
DZUP • 1 ♀; Cruzeiro do Sul; IX.1952; J. Correia leg.; ex Alva- ‘G. larcordairei Dejean’. The only way that Harold could have
renga collection; DZUP • 3 ♂; Rio Branco; 15-20.XI.1961; F. M. learnt of the existence of this name was from this label, thus
Oliveira leg.; DZUP. making clear that he did examine this d’Orbigny specimen for
Amazonas • 2 ♂, 1 ♀; Benjamin Constant; IX.1960; L. G. Pereira the description of G. inermis.
leg.; DZUP • 2 ♂ [genitalia dissected], 3 ♀; Benjamin Constant;
XI.1962; A. Silva leg.; TAMU • 2 ♂, 2 ♀; Borba, ‘Guajará’ (see Iack- Type material examined. — Lectotype • ♂; MNHN (ex Edgar
Ximenes et al. [2005: 60] for a discussion on the dubious identity von Harold and René Oberthür collections); designated by Cupello
of this locality recorded on labels of Parko specimens); VII.1943; & Vaz-de-Mello (2013: 464): “Buen. Aires” [Harold’s handwriting],
A. Parko leg.; ex Alvarenga collection; DZUP • 4 ♂, 5 ♀; Careiro, “inermis / Harold.” [Harold’s handwriting], “Ex-Musæo / E.Harold”,
Janauacá; XII.1988; J. Dellone leg; DZUP • 2 ♂, 3 ♀; Tabatinga; “LECTOTYPE”, “MNHN, Paris / EC12059 / [QR code]”, “HO-
VII.1956; M. Alvarenga leg; DZUP. LOLECTOTYPE / Gromphas / inermis / Har. / F.Z. Vaz-de-Mello
Pará • 1 ♂ [genitalia dissected]; Óbidos; XI.1954; J. Brazilino leg; 2013” [Fernando Z. Vaz-de-Mello’s handwriting] (Fig. 6).
ex Alvarenga collection; DZUP • 3 ♀; Óbidos; VIII.1963; no collec- Paralectotype • unsexed; MNHN: green disk [obverse] / not seen,
tor; DZUP • 1 ♂, 1 ♀; Óbidos; 4.V.2002; Reinhard F. leg.; CAPC. but probably aged white, stating an accession number, likely from
Rondônia • 1 ♂; Porto Velho; XII.1955; M. Alvarenga leg.; ex 1834 given that it is a d’Orbigny specimen [reverse], “G. æneus, /
Alvarenga collection; DZUP • 1 ♂, 1 ♀; Porto Velho, Madeira River Blanch / Corrientes. / M. D’Orbigny.” [unknown handwriting], “558”
(‘Amazonas / P. Velho – Rio / Madeira.’); XII.1938; no collector; ex [unknown handwriting], “Gromphas / inermis / Harold” [Harold’s
F. Justus Júnior collection; DZUP. handwriting] (Fig. 7).
Peru. Loreto • 2 ♂; Maynas, Iquitos; 106 m; 1-15.V.1948; C. Bolí-
var leg.; at light; TAMU.
San Martín • 1 ♀; Juanjuí; 10.III.2010; M. Tafur Novda (?) leg.; CAPC NEWLY DISCOVERED SPECIMENS OF THE RARE
• 1 ♂, 1 ♀; Juanjuí, ‘zona sur’; 7.VIII.2007; no collector; CAPC. GROMPHAS JARDIM CUPELLO & VAZ-DE-MELLO,
Ucayali • 1 ♀; Coronel Portillo, Pucallpa; 200 m; 22.VIII.1963;
J. Schunke L. leg.; under pig dung; TAMU • 1 ♂, 1 ♀; Coronel 2015
Portillo, Pucallpa; 200 m; 15.IX.1963; J. Schunke L. leg.; under pig
dung; TAMU • 1 ♂; Coronel Portillo, Pucallpa; 200 m; 17.IX.1963; Cupello & Vaz-de-Mello (2015) described G. jardim based on
J. Schunke L. leg.; TAMU • 1 ♂, 3 ♀; Coronel Portillo, Pucallpa; two males and three females from Brazil and Bolivia. These
200 m; 17.IX.1963; J. Schunke L. leg.; under pig dung; TAMU specimens were originally discovered in the collections of
• 1 ♂; Coronel Portillo, Pucallpa; 200 m; 18.IX.1963; J. Schunke
L. leg.; under cow dung; TAMU • 1 ♀; Coronel Portillo, Pucallpa; the Natural History Museum, London (NHM), the Oxford
200 m; 20.IX.1963; J. Schunke L. leg.; under cow dung; TAMU • University Museum of Natural History (OUMNH), and
2 ♀; Coronel Portillo, Pucallpa; 200 m; 21.IX.1963; J. Schunke L. the Universidade Federal de Mato Grosso, Brazil (CEMT).
leg.; under cow dung; in meadow; TAMU • 2 ♀; Coronel Portillo, Darren Mann, the OUMNH Coleoptera curator, allowed
Pucallpa, Pueblo Aguaytia; V.2009; no collector; CAPC • 4 ♂,
5 ♀; Coronel Portillo, Pucallpa, San Lorenzo; 17-20.VII.1992; no
me to take one of the two OUMNH female paratypes for
collector; TAMU. the museum where I was then based, the Museu Nacional,
Erroneous data Brazil. Minas Gerais • 2 ♂; Manhumirim (‘NA- in Rio de Janeiro (MNRJ). Unfortunately, three years later,
NHUMIRIM’ [sic]); 30.III.1937; Zellior leg.; DZUP. this paratype was destroyed. It was consumed along with
c. 5 million other insects housed in the MNRJ by the great
fire that devastated the main building of the institution on
THE TYPE SERIES OF GROMPHAS INERMIS the evening of 2nd September 2018 (Escobar 2018). This
HAROLD, 1869 would have meant that only four specimens are now known
to exist in collections, but this is not the case. A few months
Cupello & Vaz-de-Mello (2013) designated the lectotype earlier, in July 2018, I visited the Florida State Collection
of G. inermis from the only syntype found by them in the of Arthropods, in Gainesville, Florida (FSCA), and there
MNHN, but failed to illustrate it. I take the opportunity to I found a sixth specimen of this rare species, a male (Fig. 8).
do this here in Figure 6. Since then, I have located one of the Then, more recently, after joining Texas A&M University as
missing paralectotypes. It belongs to the MNHN, but has been the new Assistant Curator of Entomology in October 2023,
on loan to my colleague Federico Ocampo, in Argentina, since I found in the collection three more specimens, a male and
the 2000s. Many are the aspects that confirm that this speci- two females. Altogether, rather than decreasing, the number of
men is indeed part of Harold’s type series. Harold (1869b) known specimens has risen to eight, half males, half females.

36 ZOOSYSTEMA • 2024 • 46 (2)

 A B

C D

E F

Fig. 6. — The lectotype of Gromphas inermis Harold, 1869, a junior subjective synonym of G. lacordairii (Oken, 1834): A, dorsal view; B, ventral view; C, lateral
view; D, labels, note that the ‘Buen. Aires’ and ‘inermis / Harold’ labels are in Harold’s handwriting; E, posterior view; F, dorsal view of the head and protibiae.
Photographs courtesy of Christophe Rivier (MNHN). Scale bars: A-C: 5 mm; E, F, 2 mm.

37
 Cupello M.

The four new specimens also confirm that most of the dif-
ferences that Cupello & Vaz-de-Mello listed between G. jardim
and G. amazonica, its sister species (Cupello & Vaz-de-Mello
2015), are indeed accurate, particularly in the shape of the
cephalic projection and the apical tubercle of the protibiae,
as well as the distinctions concerning colouration and the
ventral carina of the protibiae (unarmed in G. jardim, armed
with a row of sharp tubercles in G. amazonica). The only main
character then deemed distinct between the two species that
I now think does not vary in a non-overlapping way between
them is the pronotal granulation. Cupello & Vaz-de-Mello
(2015) said that, while in G. jardim the pronotal granulation,
in lateral view, reaches the posterior edge of the pronotum, it
is ‘absent or rudimentary in [the] posterolateral region after
[the] lateral fossa’ in G. amazonica. This description is accurate
enough for G. jardim; all known specimens, including the
ones from FSCA and TAMU, have the entire lateral region
of the pronotum covered with granules. But a reanalysis of
the species showed that G. amazonica is more variable in this
regard than previously considered to be. While most of the
specimens are indeed smooth after the lateral fossa of the pro-
Fig. 7. — The labels of the only known (and newly found) paralectotype of notum, a few may show the integument rugose and, for this
Gromphas inermis Harold, 1869. Notice that it is a d’Orbigny specimen from reason, may be mistaken for G. jardim if no material of the
Corrientes (Argentina) and bears Blanchard’s nomen in litteris ‘G. aeneus’, which
was only mentioned in the literature by Harold (1869b). The label at the top
latter is available for comparison. Therefore, I caution readers
bearing the identification ‘Gromphas / inermis / Harold’ is in Harold’s handwrit- not to rely solely on this character to tell these species apart.
ing. Courtesy of Federico Ocampo. Attention is also needed as to the way that the apical tuft of
setae of the male protibiae of G. amazonica is described in
Cupello & Vaz-de-Mello’s identification key. The form de-
The TAMU specimens are part of the same series as the scribed refers to large- and medium-sized individuals only;
OUMNH holotype and paratypes from Bolivia, and origi- in small males, the tuft tends to be separated from the apical
nally belonged to the W. D. Edmonds collection; they came tubercle in a similar way as in G. jardim of all sizes.
to TAMU after Edmonds’ material was donated to the uni-
Additional material examined. — Bolivia. Beni • 1 ♂; Cercado,
versity in 2012 (Streit 2012). As explained by Cupello & Trinidad (‘BOLIVIA - REGION / AMAZONICA / TRINIDAD’);
Vaz-de-Mello (2015), all these specimens were misidentified X.1951; no collector; ex E. N. Kellesvig-Waering collection; FSCA •
as Gromphas lacordairii (cited as ‘lacordairei’) in Hamel- 1 ♀; Moxos, San Ignacio de Moxos, Río Ichiguita; 15°08’S, 65°18’W;
Leigue et al.’s (2006, 2009) study of the Bolivian scarabae- 155 m; 18.V.2005; C. Hamel and T. Vidaurre leg; human faeces
ine fauna. The male from FSCA, in turn, the second oldest trap, savanna; ex W. D. Edmonds colletion; TAMU • 1 ♂, 1 ♀;
Moxos, San Ignacio de Moxos, Río Ichiguita; 15°08’S, 65°18’W;
ever recorded for G. jardim, was collected in October 1951 155 m; 19.V.2005; C. Hamel and T. Vidaurre leg; human faeces
in the town of Trinidad (Cercado, Beni, Bolivia), located trap, savanna; ex W. D. Edmonds colletion; TAMU.
only about 50 km from the type locality in San Ignacio de
Moxos (Moxos, Beni). It is here addressed in the literature
for the first time. ON THE PUBLICATION DATE
Apart from the body size of the FSCA male – it is the OF GROMPHAS JARDIM
largest specimen so far recorded [total length: 17.2 mm CUPELLO & VAZ-DE-MELLO, 2015
(versus 16.3 mm previous maximum); length without head:
15.0 mm (vs 13); pronotal width: 10.1 mm (vs 9 mm); elytral In a previous publication (Vaz-de-Mello & Cupello 2018a),
width: 10.5 mm [not measured before)] –, the newly dis- I followed Dubois et al.’s (2013; 2015a, b) arguments and
covered specimens show no noticeable differences from the considered that an electronic work is only available for no-
previously known material and fit the original description menclatural purposes once it is published with the pagination
of the species well. The sexual dimorphism that Cupello & and other bibliographical information of its version issued as
Vaz-de-Mello (2015), relying on their small original series, part of a journal’s volume. For that reason, electronic-only
suspected to exist in the size and degree of impression of the versions of articles that are originally published with their
posterior pronotal fossae – clearly marked and easily visible own pagination but are later (re)issued as part of a particular
to the naked eye in males, much less marked and almost volume of a journal, and, consequently, are given new pagi-
imperceptible in females even under the microscope – is also nation corresponding to the sequence of works published in
observed in this new material, suggesting that it is indeed a that volume, were previously considered unavailable in their
real sexual difference. ‘detached’ versions. Nomenclatural novelties contained in

38 ZOOSYSTEMA • 2024 • 46 (2)

Fig. 8. — The male specimen of the rare Gromphas jardim Cupello & Vaz-de-Mello, 2015 newly discovered in the Florida State Collection of Arthropods, Gaines-
ville, Florida, USA. With the destruction of one of the paratypes in the 2018 Museu Nacional fire, it is one of the eight specimens known to survive in collections.
Photographs courtesy of Paul Skelley (FSCA).

them ‒ i.e., new names and nomenclatural acts (Dubois versions as referred to by Articles 9.9 and 21.8.3. If they are
et al. 2013) ‒ would only be given availability when their dated, they can be interpreted as preprints as defined in the
volume-integrated versions finally appeared. Nevertheless, Code’s glossary. Provided that they comply with the other
after additional thought and reading, I changed my stance demands of Article 8, these preprints, i.e., the detached ver-
on this matter. sions, are available publications and can be the place where
My new rationale is that when a work is registered in Zoo- nomenclatural novelties are made available.
bank and its first electronic-only ‘detached version’ (i.e., not Evidence that the journal has the intention of effectively
incorporated into a volume) is clearly intended by its jour- publishing the work and not merely posting a preliminary
nal to be a proper publication (i.e., not just a ‘preliminary version on its website ‒ i.e., that the journal issued the PDF
version’, ‘preview’, ‘early version’, ‘uncorrected proof ’), this with the clear ‘purpose of providing a public and permanent
detached version should be deemed the work where new scientific record’ of the content of the work (Article 8.1.1
nomenclatural novelties are given availability. In turn, the of the Code) ‒ will most usually and clearly come from the
later, re-paginated version integrated into a journal’s volume journal’s use of the unambiguous expression ‘published on-
should be interpreted in the same way as second printings line on [date]’ or ‘published on [date]’ instead of alternatives
or second editions of printed works are; i.e., not as a modi- like ‘available on-line since/from [date]’ to refer to the date
fied version of a previous ‘unfinished’ work, but instead as a when the detached version of the work was made public.
new, independent available version of a work that is already Likewise, the use of the term ‘Version of Record’ (see Krell
available. The detached version with its own pagination and 2015) confirms the intention of permanently publishing the
the one with volume pagination should, therefore, be seen work, whereas terms like ‘early view’ may indicate a prelimi-
as two independent available works; instead of seeing the nary version and others such as ‘accepted’ certainly denotes
pagination of the detached version as preliminary and the a preliminary version.
one of the volume version as definitive, it should be inter- This whole argument and my new conclusion are particularly
preted that the pagination in the former is definitive and the relevant in regard to the paper published by me and Vaz-de-
one in the latter is a new version. If this whole argument is Mello in the Journal of Natural History on a new species and the
accepted, the early electronic-only detached versions comply phylogeny of Gromphas. Its electronic-only detached version
with Article 8.1.1 and are thus not to be deemed preliminary was uploaded to the journal’s website on 12th October 2015,

39
 Cupello M.

B C

Fig. 9. — The publication date of Gromphas jardim Cupello & Vaz-de-Mello, part I; A, the webpage of Cupello & Vaz-de-Mello’s article in the Journal of Natural
History. Notice that the information mixes data from the detached and the volume-integrated versions. The green arrow points to the publication date of the
detached version as indicated by the journal, 12th October 2015, whereas the red arrow points to the bibliographical information concerning solely the volume-
integrated version published in 2016. Note also that the journal uses the expression ‘published online’ (not ‘ahead of print’, ‘early view’, or the like) to refer to
the detached version, denoting that it was not considered ‘preliminary’, but a proper publication; B, the upper part of the first page of the PDF of the detached
version. The yellow arrow points to the publication year of the PDF, 2015; the white arrow indicates the ‘article history’. Note that, while the full date, including
the day and month, of the submission and acceptance are given (white arrow), only the year of the publication is mentioned (yellow arrow); C, the same page
in the PDF of the volume-integrated version. The pink arrow points to the bibliographical data of this version (publication year, volume, number, and pagination),
whereas the black arrow shows the updated ‘article history’. Observe that here, the full publication date of the detached version (‘Online’) is given. The full date
of the publication of the volume-integrated edition itself, however, is not given anywhere either in the PDF or on the journal’s website. I was able to obtain the
date when this version was sent to print, 15th March 2016, through direct inquiry with the publisher.

whilst its volume-integrated version was sent to print on 15th A similar, though not identical, proposal was put forward
March 2016 (Henrietta Thomson, Taylor & Francis Group, by ICZN Commissioner Frank-Thorsten Krell a few years
personal communication, 6 February 2020). Considering that ago (Krell 2015). In a few words, applying the NISO/ALPSP
the journal uses the expression ‘published online’ to refer to Recommendations, Krell sees the detached and volume-in-
its 2015 detached version (Fig. 9A) and that, according to its tegrated versions not as a ‘first’ and a ‘second’ independently
Editor in Chief Andrew Polaszek (personal communication, available edition of a publication as I do here, but as the very
5th February 2020), this is meant to signify that it is the de- same work and, indeed, the very same version of the work,
finitive version of the paper (i.e., in his words, the Version of for changes in pagination and information about volume
Record), I follow them and treat the 2015 detached version and issue (‘bibliographical metadata’) are not considered as
as definitive (i.e., not ‘preliminary’). It is, therefore, the first changes in the content. Similar to my proposal, availability is
available work wherein the species-group name G. jardim gained upon the publication of what I call here the detached
Cupello & Vaz-de-Mello appeared and where it was made version provided that it is considered the Version of Record
available. The work’s 2016 volume-integrated version is, in by the publisher and is not further modified without warning.
turn, a new edition with modified bibliographical information I decided not to follow Krell’s exact proposal, however, because
and no special nomenclatural relevance. In conclusion, the I judge that mine avoids several uncertainties associated with
publication date of G. jardim is that of the detached version: his. These include determining whether a journal follows the
12th October 2015. NISO/ALPSP Recommendations (e.g., do amateur or small

40 ZOOSYSTEMA • 2024 • 46 (2)

 A

Fig. 10. — The publication date of Gromphas jardim Cupello & Vaz-de-Mello, part II; A, footer of the first page of the PDF of the detached version showing the
Zoobank link of the article (yellow arrow); B, footer of the first page of the PDF of the volume-integrated version. Notice that, although this version was published
in 2016 (and is dated as such at the upper part of the same page; see Fig. 6C), the footer is dated 2015 (pink arrow). The gray arrow shows that it is the full date
of the detached version that is given in the volume-integrated version; C, the webpage of Cupello & Vaz-de-Mello’s article in Zoobank. Note that even though
the publication date stated on the website correctly refers to the detached version (green arrow), the information in the ‘Journal Article’ section is inaccurate
(red arrow). It mixes the publication year of the detached edition (‘2015’) with the bibliographical information of the 2016 volume-integrated version (‘50 (15-16):
943-969’). The correct pagination of the 2015 detached version – the one wherein G. jardim was made available – is 1-27.

museum or society journals follow them?) or whether a Version 946 and 949-950 of the volume-integrated version. When
of Record has been later modified to produce a Corrected or citing the name G. jardim in a work, its authorship and year
an Enhanced Version of Record without explicit indication, in of publication must be given as ‘Cupello & Vaz-de-Mello,
which case the original detached version becomes unavailable 2015’, for 2015 was the year when the name was made
under Krell’s guidelines but not under mine. How can one available under Krell’s criteria (as well as under mine). But
ever know – and be assured – that a Version of Record will should the paper be dated 2015 or 2016 in the references?
not be later modified without warning and, thereby, make a If 2015, then this will agree with the year of the authorship
previously available work unavailable? of the name as cited in the text; but which pagination to
Krell’s proposal also creates difficulties during the preparation cite in the same reference? If the one of the detached ver-
of lists of references and catalogues. How should one cite a sion, then the volume-integrated version will essentially be
work first issued as a detached version? By the pagination and ignored (just like I propose). But if, instead, the pagination
publication date of this detached version or by those of the of the volume-integrated version is to be cited, then it will
later volume-integrated version? Take the case of G. jardim, for be inconsistent with its publication date being cited as 2015,
instance: both versions consist of 27 pages of text (the PDF of for this is not the pagination of the PDF published in 2015
the volume version has an additional unpaginated cover page (this inconsistency is present, for example, on the Zoobank
containing basically bibliographical information), but their webpage of Cupello & Vaz-de-Mello’s article; see Fig. 10C).
pagination is not the same. While the 2015 detached version If, to fix this problem, one decides to cite the article in the
is paginated [1]-27, the 2016 volume version is paginated references based entirely on the information of the volume-
[943]-969. The information making the name available, in integrated version, including its year of publication (2016),
particular, appears on pages 4 (information on the holotype) then the inconsistency will be with the year of publication
and 7-8 (description) of the detached version, and on pages mentioned in the text for the authorship of the name (2015).

41
 Cupello M.

In one way or the other, considering the detached and the threatening the availability of the original work), and so it is
volume-integrated versions as the same as proposed by Krell easier (and faster) for new names and nomenclatural acts to be
(2015) is, as aptly pointed out by Dubois et al. (2015b), an available than under Dubois et al.’s, and all while still respect-
artificial procedure and creates a lot of problems. Under my ing the principle of permanency of the zoological literature.
proposal, in contrast, the date and pagination to be cited on It must be noted that Dubois et al. (2013, 2015a, b) have
the list of references are unambiguous: those of the detached analysed the alternative that I present here ‒ viz., to treat the
version, as it was there where the name was made available. detached and volume-integrated PDFs as different works and
Dubois et al. (2015a, b) have already made similar comments to deem them independently available ‒, but came to a dif-
and stressed the importance of page citation in taxonomic ferent conclusion. While they agree with the first part of the
works, a claim that I fully endorse. The fact that pagination argument ‒ that the two versions are distinct publications, not
is not regulated by the Code, as Krell (2015) argued, does the same as argued by Krell (2015) ‒, they consider that the
not make it less relevant for taxonomists and cataloguers or first version, the detached version, is unavailable because, in
its stability less desirable. On the contrary, anyone who has their opinion, it fails to comply with Articles 8.1.1 and 9.9.
carried out a taxonomic project, particularly the preparation Their point is that because the PDF of the detached version
of catalogues and comprehensive systematic revisions ‒ or, is removed from the journal’s website and replaced with the
to be more general, anyone who has needed to check in the volume version once the latter is published, this would indi-
original publication a quotation or specific information cited cate that the journal regarded the detached version as only
by a third author ‒ is aware of the importance of knowing, preliminary and so that it did not have the ‘purpose of pro-
and being able to cite, the exact and unambiguous place in viding a public and permanent scientific record’ as demanded
a publication where a piece of information is to be found. by Article 8.1.1. I disagree.
In contrast to Krell’s (2015), my procedure, I believe, is The term ‘published’, if used by the journal to refer to the
more straightforward and in line with the way the Code and detached version, clearly denotes the stated purpose of the
zoologists have been approaching printed publications for editors and publishers that the content of this version is pub-
decades: once a work has been published in a way to satisfy lic, citable, and a permanent part of the scientific literature.
the Code’s criteria of availability, that version is the defini- However, if, in practice, they replace the PDF of the detached
tive one where new nomenclatural acts are established, no version with that of the volume-integrated version and the
matter how many new modified versions are later reissued or former thus ceases to be accessible from the journal’s website,
whether the original version is no longer obtainable from the this does not deny the original intention (the purpose) behind
publisher. Concerning the latter point – viz., the detached their action. Rather, this only shows either that the editors are
version not being publicly obtainable after being replaced bona fide ignorant of the intricacies of the Code (they believe
by the volume-paginated version on the journal’s website –, that replacing the original PDF with another nearly identical,
it could be argued (see, e.g., Dubois et al. 2015b) as a dif- differing only in the bibliographical data, would not inter-
ficulty for implementing my proposal. However, if Krell is fere with the availability of the original work; i.e., it would
correct (and I believe he is) that the later volume-integrated not interfere with its original purpose of being a permanent
versions are almost always identical to the early detached ones scientific record), or that they simply disagree with Dubois
except for their bibliographical content (but see Dubois et al. et al.’s interpretation of what a published work is and that, in
2013), then there should be no problem: in practice, these their interpretation, replacing the PDFs would not go against
volume-integrated versions can be used as surrogates for the Article 8.1.1 (e.g., if they follow Krell’s proposal). The point is
detached versions when these are not obtainable. Nevertheless, not whether the PDF of the detached version remains obtain-
if Dubois is correct and many journals do produce volume- able from the journal’s website or not, but rather, what the
integrated versions with additional modifications, then my original intention of the editors was in preparing the PDF of
procedure will safeguard the availability of the early works, the detached version. Article 8.1.1 does not demand a work
which would become unavailable under Krell’s, and avoid the to be permanently public, but only that it being permanently
chaos of never knowing whether a work has been – or will public has to be the publishers’ original intention1.
be – made unavailable due to modifications included without Being, in practice, permanently obtainable from the pub-
warning in subsequent versions produced by the publisher. lisher has indeed never been a requirement for a work to be
Despite my new stance on electronic publications, I still available or to be regarded as a permanent scientific record in
consider Dubois et al.’s (2013, 2015a, b) arguments to be in zoology. For instance, is the 10th edition of Systema Naturae
most parts quite sensible, as indeed recognised by Krell (2015) still obtainable from Lars Salvius? Of course not. It is currently
himself, and should be taken into consideration by readers. only accessible through original copies in a handful of public
I only prefer my new stance over Dubois et al.’s because I con- and private libraries and a very few antique bookshops around
sider mine protects more widely the availability and priority the world, through photocopies or facsimile reproductions,
of new names and nomenclatural acts and leaves authors less or through digitised copies obtainable online from specialised
exposed to bad editorial procedures over which they are usu- websites (e.g., Biodiversity Heritage Library, Google Books).
ally powerless. Under my proposal, editors have to comply This situation is analogous to that of detached versions that
with fewer requirements for a work to be available (e.g., they have been removed from their respective journals’ websites:
can produce as many modified versions as they like without they remain in existence only on the computers (digital library)

42 ZOOSYSTEMA • 2024 • 46 (2)

 A B

Fig. 11. — Newly (re)discovered specimens of the potentially extinct Gromphas dichroa Blanchard, 1846: A, labels of the holotype female and oldest known speci-
men, property of the Muséum national d’Histoire naturelle, Paris. It had been deemed lost in the last revision of the genus (Cupello & Vaz-de-Mello 2013). The
black arrow indicates a label in Harold’s handwriting; Harold (1869b) indeed reported the study of the specimen during his visit to the MNHN, when he confirmed
that the species belongs to Gromphas – as originally maintained by Blanchard (1846) and Lacordaire (1855) – instead of Bolbites as he had previously suggested
(Harold 1869a; see Cupello et al. 2021). Courtesy of Federico Ocampo; B, C, the specimen from the Institut royal des Sciences naturelles de Belgique, Brussels.
Courtesy of Alain Drumont (RBINS); D, the four-specimen series housed in the Museu Anchieta de Ciências Naturais, Porto Alegre, Brazil. This series includes the
most recently collected individual of the species, caught in Santa Catarina (Brazil) in 1954 (second from the left). Courtesy of Luísa Menezes da Silveira (MGAP).

43
 Cupello M.

of those who downloaded them before removal, or of those Nomenclature and is registered in ZooBank under number [exact
who had the PDF shared with them by colleagues, as well as Zoobank registration number]’. Text between square brackets is to be
in physical libraries in the form of house-made printed cop- replaced with information relating to the work in question.
ies, or stored on academic social networks like ResearchGate
or Academia. In all these cases, modern or old, new copies
ceased to be obtainable from the original publisher. ON SOME MISLABELLED SPECIMENS
A similar thing could also be said of many printed journals OF GROMPHAS AMAZONICA HOUSED
published (currently or in the past) by small amateur socie- IN THE BRUSSELS MUSEUM (RBINS)
ties, small museums, or even privately by amateurs around
the world, particularly in Europe. Anyone who has worked While examining the collection of the Institut royal des Sci-
on the taxonomy of Coleoptera, for instance, knows how dif- ences naturelles de Belgique, Brussels (RBINS), for this work,
ficult it can be to obtain copies of papers published by many I realised that, quite surprisingly, all the specimens of G. ama-
of these small journals. While this situation is far from desir- zonica housed there either are certainly mislabelled or bear
able, rarity and difficulty of obtaining have never prevented a suspicious provenance data. To prevent future misunderstand-
printed work from being available. Why should it be the case ings and the potential publication of erroneous geographical
for electronic publications? Although it would undoubtedly records, it is relevant to address these specimens here. Two
be preferable for journals to maintain the detached version on males gifted along with 759 268 other insects to the RBINS
their websites alongside the volume-integrated version, I see in 1939 or before (one with accession number 12.595) and
no reason why them failing to do so would be any different originating from the collection of the French insect dealer
from printed works that ceased to be obtainable from the Eugène Le Moult (1882-1967) bear information saying they
original publisher, nor why considering otherwise would be were collected in the municipality of Uberaba, state of Minas
a natural (or even desirable) interpretation of the Code and Gerais, in central Brazil and within the Cerrado ecoregion.
its 2012 Amendment (ICZN 2012). This would only create A series of three males and seven females from the same gift
more uncertainties. So, I disagree with Dubois et al. and argue (same accession number) are labelled as if coming from the
for the proposal that I put forward above. town of Muzo, in the Colombian department of Boyacá, and
Yet, I concede that, sometimes, it can be difficult to ascer- within the Magdalena Valley. Finally, a male and a female,
tain the purpose of a detached version uploaded to a journal’s these from a lot of 26 488 insects acquired by the RBINS from
website. Such ambiguity may trigger endless quarrels about J. J. Gillet on 5th October 1935 (accession number 10.640),
the publication date of a work under the Code, and detract are said to have been caught in Paraguay without further data.
zoologists from the actual object of their studies, animals and Since, based on trustworthy data, G. amazonica is known only
their biology. It would greatly serve stability and universality in from localities along the Huallaga, Pisqui, Ucayali, Juruá,
zoological nomenclature and the science of zoology overall if and Amazon Rivers, in the Amazon rainforest (Cupello &
journals and their publishers stated fully unambiguously their Vaz-de-Mello 2013, 2015; see Fig. 5), these three records,
intention as related to Art. 8.1.1 when posting the PDFs of all of which lying outside that area, are most likely incorrect.
works containing zoological names and new nomenclatural Janssens (1940) examined the two Gillett specimens sup-
acts on their websites2. Currently, there is nothing in the Code posedly from Paraguay and considered that they indicated
enforcing this. All I can do here is urge editors and publish- that G. amazonica is much more widely distributed than its
ers to expressly state on the first page of the earliest version previously thought Amazonian restriction (e.g., in Bates 1870;
of a work that they deem published that it has the ‘purpose d’Olsoufieff 1924). In my opinion, he was wrong in accept-
of providing a public and permanent scientific record’. This ing the label content at face value. In regard to the Muzo
will prevent any doubts about the work’s compliance with (Colombia) record, which could be argued to be the least
Art. 8.1.1 and, provided that the other criteria established in improbable given the strong biotic influence from Amazonia
Chapter 3 of the Code are met, ensure its availability. At the in the Magdalena Valley, Alejandro Lopera (personal com-
same time, to solve this problem definitively in a formal way, munication to MC, 7th February 2020), a specialist in the
I put forward the following proposal to the Commission: in Colombian dung beetle fauna, informed me that the area near
the forthcoming 5th edition of the Code, Article 8.5.2 should Muzo and the rest of the Magdalena River Basin ‘have been
be amended as follows: well sampled, so if it were a correct location it should have
been reported by now’ (see, e.g., Medina et al. 2012). This
8.5. Works issued and distributed electronically strengthens my confidence that the Muzo record is indeed
To be considered published, a work issued and distributed electroni- incorrect. It is noteworthy, however, that G. aeruginosa, the
cally must other typically Amazonian species of Gromphas, is known from
reliable specimens to have been collected at least three times
[...]
between 1979 and 1982 at a locality in the upper Magdalena
8.5.2. state the following sentence on its first or last page: ‘This work Valley, the Colombian town of Gigante (Huila) (Cupello &
is issued on [exact date of publication, with day, month, year] for Vaz-de-Mello 2013). Therefore, if, against my expectation,
the purpose of providing a public and permanent scientific record as G. amazonica does prove to be present there, it will not be
established in Article 8.1.1 of the International Code of Zoological unprecedented for an Amazonian Gromphas.

44 ZOOSYSTEMA • 2024 • 46 (2)

Material examined. — Brazil. Amapá • 1 ♂ [genitalia dissected], I found one male and three females collected between 1928
1 ♀; Macapá, BR-156, Km 14; 26.XI.1981 [in cow dung]; I. S. and 1954 in the Brazilian states of Rio Grande do Sul and
Gorayeb and team leg.; MPEG. — Amazonas • 1 ♂ [genitalia dis- Santa Catarina (Fig. 11D). These data represent the most
sected]; Benjamin Constant; IX.1955; I. C. Lima leg.; ex Alvarenga
collection; DZUP. recent record for the species (seven years later than the pre-
Peru. Loreto • 3 ♂ [all with genitalia dissected], 2 ♀; vious most recent record from 1947) and the first report for
Ucayali, Contamana; XII.2001; no collector; TAMU. Santa Catarina; furthermore, the Rio Grande do Sul speci-
Erroneous data: Brazil. — Minas Gerais • 2 ♂ [both with genitalia men caught in Nova Petrópolis represents a new municipal-
dissected]; Uberaba; no further data; ex Le Moult collection (one
with accession number 12.595); RBINS. — Colombia. Boyacá •
ity record in the state. These two new records also confirm
3 ♂ [all with genitalia dissected], 7 ♀; Occidente, Muzo; no further that the species at least used to occupy areas not only in the
data; ex Le Moult collection (all with accession number 12.595); Uruguayan Savanna ecoregion (also known as the Pampas),
RBINS. — Paraguay • 1 ♂, 1 ♀; no further data; ex J. J. Gillet col- from where most of the specimens are known, but also in
lection (accession number 10.640); RBINS. the bordering southern fringe of the Atlantic Forest; whether
they were caught in the rainforests themselves, in transitional
NEW RELEVANT RECORDS FOR THE VANISHED areas, or in deforested farmlands is unclear. The four MGAP
GROMPHAS DICHROA BLANCHARD, 1846 specimens encompass much of the colour variation shown by
G. dichroa: both Rio Grande do Sul females have the typical
Cupello & Vaz-de-Mello (2013, 2015) discussed the rarity bicoloured pattern of the species, with a shiny red head and
of G. dichroa in collections. Only 9 males and 9 females were pronotum and blue elytra, whereas the Itapiranga individu-
known to us apart from the missing holotype3, the lost six als show either the body entirely green (the male) or its head
specimens studied by Barattini & Sáenz (1961, 1964), and and pronotum red at centre and green over the lateral edges
the two equally lost specimens mentioned by Burmeister and its elytra uniformly green (the female). As discussed and
(1874). No individuals were known to have been caught illustrated by Cupello & Vaz-de-Mello (2013, 2015), this
since 1947 despite intensive collecting efforts across the whole variation can occur within a single population, and
species’ range from southern Paraguay to the Argentinian the bicoloured pattern evolved in parallel in at least two other
province of Buenos Aires4. Even Antonio Martínez (1922- sympatric lineages of dung beetles, namely Bolbites onitoides
1993), who collected and studied the Argentinian dung beetle Harold, 1868 (Cupello et al. 2021) and the clade formed by
fauna so extensively through the mid- and late-20th century, Canthon lividus Blanchard, 1846 and C. auricollis Redten-
never collected a G. dichroa or obtained a specimen caught bacher, 1868 (Vaz-de-Mello & Cupello 2018b).
by others after the 1940s. The same is reported by Federico The MGAP is only one of the two museums in Porto Alegre,
Ocampo (personal communication, 21 June 2021), who has the capital city of the state of Rio Grande do Sul and which
done fieldwork more recently in areas within the historical is (or was) part of the range of G. dichroa, the other being the
range of G. dichroa, including Uruguay and Buenos Aires. Museu de Ciências Naturais da Fundação Zoobotânica do
For these reasons, I started to wonder whether G. dichroa Rio Grande do Sul (MCNZ). While the MGAP collection
may have become extinct. grew rapidly during the first half of the 20th century, it has
Since the publication of Cupello & Vaz-de-Mello’s revi- been mostly inert since the death of its founder, Father Pio
sion, I had the opportunity to visit several other collections Buck (1883-1972), and houses few specimens collected after
in Europe and the Americas, including all the main ones in the 1950s. It is not surprising, therefore, that no G. dichroa
southern Brazil and Paraguay, and in just two of them did collected after that decade is held there. On the other hand,
I find additional specimens of G. dichroa. One is the Brussels the MCNZ was founded as a separate museum only in 1954
museum (RBINS), which houses a single specimen. As can (Nedel 2005) and owns scarabaeine material collected mostly
be seen in Figure 11B, C, this specimen shows the bicoloured thereafter. Its dung beetle collection is particularly rich and
colouration pattern of the species (similar to fig. 14a of Cu- includes material collected all over Rio Grande do Sul, includ-
pello & Vaz-de-Mello 2013) and bears only two labels, neither ing several rare species native to the state and other regions of
informing where and when the specimen was collected. But Brazil (personal observation in 2018). Were G. dichroa still
this certainly happened before 1935, for the accession number living in that region, it would be expected that at least some
‘10.640’ stated on one of its labels refers to material acquired specimens would be housed in the MCNZ. But this is not
on 5th October that year by the RBINS from the Belgian the case. Why? This becomes even more intriguing because
coleopterist Joseph Jean Edouard Gillet (1865-1937) (Alain Gromphas lacordairii, a closely related species (Cupello &
Drumont, Coleoptera curator at RBINS, personal communica- Vaz-de-Mello 2015), is abundant both in the MGAP and in
tion, 16th March 2020). This same specimen was mentioned the MCZN and continues to be commonly caught through-
five years later by Janssens (1940) in his study of the RBINS out the region where G. dichroa used to be found, including
phanaeines, a work overlooked by Cupello & Vaz-de-Mello. specific localities such as Porto Alegre (last G. dichroa from
Regrettably, I did not have time to sex the specimen during 1928; last G. lacordairii from 1993) and the greater Buenos
my stay in Brussels, nor did Janssens mention it in his paper. Aires area (1935 and 2006, respectively) (Figs 12-14; Table 1).
The second collection where I found additional specimens But despite its decades-long disappearance and my suspicion
of G. dichroa is that of the Museu Anchieta de Ciências Natu- of possible extinction, I concede that no formal assessment
rais, Colégio Anchieta, Porto Alegre, Brazil (MGAP). There, of the conservation status of G. dichroa is currently possi-

45
 Cupello M.

Fig. 12. — The disappearance of Gromphas dichroa Blanchard, 1846. The map shows the date of the last record at each known collecting point for the species.
The different colours represent the decade of the last record at each locality. Notice that G. dichroa has not been found since the 1950s. Compare with the map
for G. lacordairii in Figure 13. Observe, for instance, that G. lacordairii continues to be found even in Porto Alegre and the greater Buenos Aires area, places
where G. dichroa was last collected in the 1920s and 1930s, respectively. Data gathered from the material examined by Cupello & Vaz-de-Mello (2013, 2015)
and herein, as well as from additional records found in the literature (Blanchard 1846; Martínez 1959; Barattini & Sáenz 1961, 1964). See Table 1 for more details.

ble under the criteria set out by the IUCN Species Survival denotes a general lack of collection effort. These include,
Commission (2012). At least the historical extent of its in Argentina, Santo Tomé (last G. dichroa from 1928; last
geographical occurrence, beyond 600 000 km2, is way too G. lacordairii from 1926) and Santa Maria (last G. dichroa
large for any of the IUCN threatened categories (maximum from 1947, no G. lacordairii), and, in Brazil, Cerro Largo
of 20 000 km2 for vulnerable), and nothing in terms of the (1941 and 1949, respectively), Nova Petrópolis (both species
species’ population sizes and trends, habitat preferences, last recorded in 1928), and Itapiranga (idem in 1954). Also
current area of occupancy, and life habits is available for a worthy of attention are the Campanha Gaúcha and Serras
proper judgement. Formally classifying the species as ‘extinct’, de Sudeste regions of the Brazilian state of Rio Grande do
in particular, would be, for the moment, too precipitate Sul. Although G. dichroa has never been recorded from
as G. dichroa does not match the IUCN’s requirement of there, these regions are apparently within its range and still
having been the special target of exhaustive surveys in all encompass vast extensions of relatively pristine areas of the
places where it is known to have been historically present Pampas, particularly around the nature preserve Área de
as well as others where it might be expected to occur. Un- Proteção Ambiental Ibirapuitã, on the border of Brazil and
til these are performed using all the collecting techniques Uruguay (see Souza et al. 2020’s map). Should all these future
available for dung beetles (not only pitfall traps baited with collections be performed and, yet, G. dichroa continue failing
excrement; see below), the species is better classified as ‘data to appear, then we can finally ‒ and unfortunately ‒ deem
deficient’. Critical localities are those where even G. lacor- the species extinct. Nevertheless, if any of the localities does
dairii has not been recorded over the last decades, for this retrieve the species, a study of the re-discovered populations

46 ZOOSYSTEMA • 2024 • 46 (2)

 New discoveries about Gromphas

Fig. 13. — Gromphas lacordairii (Oken, 1834) records in the general area of occurrence of G. dichroa. Like Figure 12 for G. dichroa, this map shows the date of
the last record at each known collecting point for G. lacordairii. The ‘general area of occurrence of G. dichroa’ (the pale yellow area on the maps) is here inter-
preted to include southern Paraguay, southern Brazil (Rio Grande do Sul and western Santa Catarina), northeastern Argentina (Misiones, Corrientes, Entre Ríos,
the Buenos Aires city and northern and northeastern Buenos Aires province), and Uruguay. The different colours of the locality symbols represent the decade
of the last record. Data gathered from the material examined by Cupello & Vaz-de-Mello (2013, 2015) and herein, as well as from additional records found in the
literature (Barattini & Sáenz 1961, 1964; Cabrera-Walsh & Cordo 1997; Morelli et al. 2002; Sánchez & Genise 2008). See Table 1 for more details.

ZOOSYSTEMA • 2024 • 46 (2) 47

 Cupello M.

53

50

45
45

40

35
35 34

30

25 24
23
21
20
17
15
13

10 9

5
5
2 3
2
1 1 1
0
1890s 1900s 1910s 1920s 1930s 1940s 1950s 1960s 1970s 1980s 1990s 2000s 2010s

Gromphas lacordairii Gromphas dichroa

Fig. 14. — Number of specimens of Gromphas lacordairii (Oken, 1834) and G. dichroa Blanchard, 1846 collected in the ‘general area of occurrence of G. dichroa’
(see captions of Fig. 13 for explanation) in each decade since 1890 (refer to Table 1 for details. Localities in the table from personal communication with P. G. Da
Silva not included here; see explanation in ‘Material and Methods’). These figures indicate that although never common, G. dichroa was consistently collected
during the first half of the 20th century. The most recent specimen, however, dates from 1954, 70 years ago (from Itapiranga, Santa Catarina). At the same time,
G. lacordairii has always been much more common and has never ceased to be collected.

will enable a formal assignment to one of the IUCN catego- the 1950s), this species was recently rediscovered living
ries. Another dung beetle species occupying much the same about 1000 km farther south in the municipality of Santa
range as G. dichroa, Anisocanthon pygmaeus (Gillet, 1911), Maria in the Rio Grande do Sul state (da Silva et al. 2011,
and which has, too, vanished since the 1950s, has also been 2012a, 2013) and, subsequently, a little farther north in the
classed as data deficient by Vaz-de-Mello et al. (2014). My municipality of Anitápolis, Santa Catarina state (Simões-Cl-
treatment thus follows the established convention. ivatti & Hernández 2022). These rediscoveries of long-lost
This prudence is indeed justified. A number of Brazilian dung beetles are, in fact, not limited to the Brazilian fauna,
dung beetles that, like G. dichroa, had vanished during the as recently shown by Deschodt et al. (2021) in Madagascar
second half of the 20th century have been recently re-dis- and Hielkema (2023) in Suriname.
covered either in new populations or in the same area where But, if not extinction, what else could explain the wide-
they were previously known. A representative of the latter spread disappearance of G. dichroa whereas G. lacordairii
case is Paracryptocanthon borgmeieri (Vulcano, Pereira & continues to be abundant? Part of the explanation may be
Martinez, 1976), which until recently was known from that the populations of G. dichroa have always been naturally
just seven females collected in the 1960s and 1970s and small and sparse. The number of specimens collected before
which was rediscovered in 2013 through the collection of the 1950s, for example, was already much smaller than that
a large series of specimens of both sexes at the type locality of G. lacordairii (Fig. 14). Kohlmann (1991) and Price &
(Pacheco & Vaz-de-Mello 2017). Its almost 40-year disap- May (2009) have shown that pairs of sympatric phanaeine
pearance is particularly remarkable because the species’ entire species with similar body size like the dichroa/lacordairii pair
range lies in the heart of the heavily collected city of Rio de usually differ drastically in their local relative abundance,
Janeiro. In turn, a case of a vanished species that was later from a ratio of c. 3 individuals per 7 to 1/100. A similar
rediscovered living far from its previously known range is proportion was found by Cupello & Vaz-de-Mello (2018)
Sulcophanaeus rhadamanthus (Harold, 1875). After being for the deltochilines Sylvicanthon seag Cupello & Vaz-de-
known for decades from a few old specimens collected in Mello, 2018 and S. securus (Schmidt, 1920) in northern
the Serra da Mantiqueira and Serra dos Órgãos mountain Amazonia, although the exact figures vary from place to
ranges in southeastern Brazil (Edmonds 2000; the author place. More impressive still, Feer (2000) observed the same
does not provide precise label data of the specimens he ex- phenomenon across an entire Scarabaeinae community in
amined, but it is likely that they were all collected before French Guiana. This persistent pattern may be the result of

48 ZOOSYSTEMA • 2024 • 46 (2)

the competitive exclusion principle (Hardin 1960). One of complete local extinction. This in itself may already explain
the competitors, being more efficient in a given context, why G. dichroa has always been rarer: it is less competitive
multiplies and pushes the other to lower abundances or even than its close ally and always sympatric G. lacordairii.

Table 1. — Number of specimens of G. dichroa Blanchard, 1846 and G. lacordairii (Oken, 1834) collected in the range of G. dichroa (see definition in the captions
of Fig. 13). Data compiled from the material examined by Cupello & Vaz-de-Mello (2013, 2015) and herein, from information provided by colleagues (F. Ocampo
and P. G. da Silva, the latter indicated by the collection abbreviations LBEV, MCCR, MCTP, MECB, MHNU, MRGC), and from additional records found in the
literature (Blanchard 1846; Martínez 1959; Barattini & Sáenz 1961, 1964; Cabrera-Walsh & Cordo 1997; Morelli et al. 2002; Sánchez & Genise 2008; Canziani &
González-Vainer 2022). Bold denotes places where both species have been recorded.

Country State/
Department Locality Date (number of specimens)
Gromphas dichroa
Argentina
Buenos Aires Anchorena X.1935 (1)
Corrientes Santo Tomé XII.1922 (1), IX.1926 (1), II.1927 (4), X.1928 (1)
Misiones Santa Maria XI.1947 (1)
Brazil
Rio Grande do Cerro Largo I.1931 (or 1933?) (1), III.1941 (1)
Sul Nova Petrópolis I.1928 (1)
Porto Alegre X.1928 (1)
Santa Catarina Itapiranga XI.1934 (1), X.1954 (1)
Paraguay
? Southern Paraguay? ? (unknown number of specimens; reference: Martínez 1959, who merely listed the species for
the country. The presence of G. dichroa in Paraguay, either historical or present, still has to
be confirmed by physical specimens or any more substantial evidence)
Uruguay
No specific locality ? (4), XII.1894 (1)
Florida No specific locality ? (unknown number of specimens; reference: Barattini & Sáenz 1961, 1964)
Salto (?) ? (1)
Soriano No specific locality ? (unknown number of specimens; reference: Barattini & Sáenz 1961, 1964)
Montevideo Montevideo ? (1), 1826-1827 or 1829 (1; reference: Blanchard 1846; this record refers to the holotype, the sole
specimen known from Montevideo. Blanchard (1846) stated that the specimen was collected
by Alcide d’Orbigny (1802-1857) in Montevideo ‘near the sea’. D’Orbigny stayed in Montevideo
during three different periods: 1) 29th October to 17th November 1826; 2) 2nd December
1826 to 10th January 1827; and 3) 12th to 27th December 1829 (d’Orbigny 1835, 1839-1843;
Papavero 1971). He may have collected the holotype in any of these three periods.)
Tacuarembó No specific locality ? (unknown number of specimens; reference: Barattini & Sáenz 1961, 1964).
No data 1
Gromphas lacordairii
Argentina
Autonomous City Buenos Aires city IV.1943 (1), I.1993 (1)
of Buenos Aires
Buenos Aires ‘17 km south 8.I.1980 (1)
of Buenos Aires’
Boulogne Sur Mer XI.1941 (1)
Florencio Varela I.1978 (1)
Isla Martín García IV.1937 (1)
La Plata ? (9)
Martínez 16.II.1924 (1)
Navarro 2006 (6 nests; reference: Sánchez & Genise 2008)
Pergamino II.1949 (1)
San Isidro 14.IV.1956 (1), XII.1967 (1), XI.1973 (1),
San Pedro ? (1)
William C. Morris 1988-1991 (unknown number of specimens; reference: Cabrera Walsh & Cordo [1997])
Corrientes No specific locality ? (11)
‘5 km W Ituzaingó’ IX.1979 (1)
Ituzaingó III.1976 (2)
San Roque I.1920 (1)
Santo Tomé ? (10), X.1925 (5), XII.1925 (1), IX.1926 (1)
Entre Ríos No specific locality ? (2)
Pronunciamiento I-III.1963 (23)
Villa Paranacito ? (1)
Misiones No specific locality ? (15)
Dos de Mayo I.1966 (1)
Puerto Iguazú X.1927 (1)
San Ignacio 1928-1929 (2), 21.X.1929 (1)
Upper Paraná River 01-18.XI.1933 (1)

49
 Cupello M.

Table 1. — Continuation.

Brazil
Rio Grande do No specific locality ? (5), III.1915 (1), 7.XI.1959 (1), 20.X.1961 (1), I.1995 (1)
Sul Aceguá 08-14.XI.2011 (7)
Bagé 6.XII.2006 (1), 17.I.2007 (1), 07-13.I.2012 (4)
Bossoroca III.1971 (LBEV)
Cachoeira do Sul II.1996 (4)
Canoas 3.I.1958 (MRGC)
Capão da Canoa 22.I.1978 (1), 10.IX.1987 (MCTP), 9.X.1989 (1), 18.XI.1989 (7)
Caxias do Sul 17.IX.1987 (MECB)
Cerro Largo I.1931 (1), I.1939 (1), I.1940 (10), 1949 (2), I.1949 (2)
Charqueadas 12.I.1989 (26)
Chiapetta 16.XI.1974 (LBEV)
Cidreira 2.XI.1992 (4)
General Câmara 16.IX.1982 (1)
Imbé II.1961 (9), II.1973 (2), 6.XI.1999 (1)
Mostardas I.1945 (4)
Nova Petrópolis 28.I.? (MRGC), I.1928 (16)
Osório 3.I.1950 (5), 24.I.1958 (9), 25.I.1958 (2), 2.II.1964 (1)
Pareci Novo 20.X.1927 (1), VIII.1935 (1), XI.1938 (1)
Pelotas XII.1934 (2), I.1935 (2), XII.1935 (MHNU), 17.II.1941 (MHNU), 10.XI.1953 (1), XII.1955 (MHNU),
1959 (MCCR), 12.II.1962 (MECB), II.1963 (1), II.1968 (MECB), 2.IV.1968 (MECB), 2.I.1970
(MECB), 13.X.1975 (MECB), 16.X.1978 (MECB), 2.IX.1982 (MECB), 14.X.1983 (MECB),
III.1988 (MECB), IX.1995 (MCCR), X.1995 (MCCR)
Porto Alegre ? (1), VI.1927 (1), 16.X.1928 (1), 29.IX.1929 (1), 29.V.1934 (1), 15.XI.1935 (1), 1941 (1), 1943
(1), 25.IX.1949 (1), 3.X.1956 (1), I.1959 (3), 25.XI.1962 (MRGC), 6.XII.1962 (1), 7.X.1963 (1),
22.X.1963 (1), 1964 (1), 26.I.1965 (1), 7.II.1965 (1), 7.X.1965 (1), 26.X.1966 (1), 18.VIII.1970
(1), 8.XI.1993 (1)
Quaraí I.1945 (1)
Rio Grande 26.IV.2005 (MCCR)
Santa Maria 7.IV.1971 (LBEV), 2.II.1973 (LBEV), 23.X.1973 (LBEV), 1.X.1974 (LBEV),
30.X.1974 (LBEV)
Santo Augusto 12.X.1966 (1)
São Jerônimo 13.I.1989 (2), 11.I.1989 (6),
São Leopoldo XI.1958 (1)
Torres X.1956 (1), 2.II.1960 (MCTP), 9.XII.1964 (1), II.1965 (MRGC)
Tramandaí I.1979 (1)
Triunfo 21.IX.1989 (1), 14-15.I.1997 (1)
Uruguaiana 18.XII.1996 (2)
Viamão X.1956 (2)
Xangri-Lá 20.I.1968 (1)
Santa Catarina Itapiranga IX.1953 (1), V.1954 (1)
Seara: Nova Teutônia 25.IX.1963 (1), 23.X.1963 (1), 2.I.1964 (1), 30.I.1964 (1), I.1966 (1), II.1970 (1), XII.1970 (6)
Seara: Pinhal IV.1958 (1), XII.1958 (1)
Paraguay
Caaguazú Caaguazú XII.1948 (1)
Caazapá Caazapá 01-8.XII.1990 (1)
Capital District Asunción IX.1922-IV.1923 (1), 12.X.1990 (1), 7.I.1991 (1), 26.II.1992 (1)
Central Ypacaraí 23.X.1996 (2)
Guairá Colonia Independencia I.1950 (1), III.1950 (1)
Cordillera del Ybytyruzú, 24.XII.1994 (1)
Carpa Cué
Cordillera del 19.II.2004 (2)
Ybytyruzú, Cerro Akatí
Villarrica X.1941 (1)
Itapúa Coronel Bogado I.1944 (1)
Isla Yacyretá ? (3), 14.XII.1999 (1)
Paraguarí Parque Nacional Ybycuí 13.I.1996 (1)
Sapucai 1903 (2)
Uruguay
No specific locality ? (7)
Artigas Arroyo Tres Cruces 18.I-20.II.1958 (1)
Grande
Canelones Atlántida ? (1)
Campo Experimental 10.XI.2020 (6)
Instituto de Higiene
Pando 10.I.2005 (1)
Cerro Largo Bañado de Medina 13-20.XII.2011 (8)
Melo 13-20.XII.2011 (2)

50 ZOOSYSTEMA • 2024 • 46 (2)

 Table 1. — Continuation.

Colonia No specific locality ? (1)

Riachuelo ? (1)
Durazno No specific locality ? (unknown number of specimens; reference: Barattini & Sáenz 1961, 1964)
Florida Cerro Colorado 1992-1994 (44; reference: Morelli et al. 2002)
Puntas de Sauce 107 (1999; reference: Canziani & González-Vainer 2022. Date of collection not stated in the paper,
de Maciel but year informed by Patricia González-Vainer, pers. comm., 13th September 2022)
Maldonado No specific locality ? (unknown number of specimens; reference: Barattini & Sáenz 1961, 1964)
10 km SW of José I.2014 (unknown number of specimens; reference: Federico Ocampo, pers. comm.)
Ignácio
Montevideo Montevideo ? (4), 20.XII.1929 (1), 10.X.1933 (1)
Rivera Vichadero 21-27.XI.2011 (2)
Tacuarembó near San Gregorio 20.X.2019 (1)
de Polanco

And there may be more to this. In evolutionary time, this 2012; Canziani & González-Vainer 2022; but see da Silva
continuous pressure from competition may eventually lead the 2011; da Silva et al. 2012a, b, 2013; da Silva & Bogoni 2014)
rarer, less competitive species to niche specialisation through may have been inefficient in attracting the species. In sum, it
ecological character displacement (Brown & Wilson 1956; may be that the disappearance of G. dichroa since the 1950s
Pfennig & Pfennig 2009). Perhaps the competition with the is simply the consequence of a peculiar biology (this itself a
presumably more efficient G. lacordairii drove G. dichroa to result of the competition with the more ecologically aggres-
stenotopy. And if, in this specialisation, G. dichroa departed sive G. lacordairii) that keeps population densities low and
from the usual coprophagous, non-inquiline behaviour of spatial presence limited, coupled with the fact that modern
most dung beetles, including G. lacordairii, maybe becom- collectors are not looking in the right places or applying the
ing, for example, mycetophagous or an inquiline of insect right collecting techniques that earlier naturalists, by chance
or vertebrate nests, this would explain why it has not been or design, employed.
attracted to regular dung traps. Edmonds (2000) indeed sug- But having now examined all the difficulties inherent in the
gested that S. rhadamanthus, with a body size similar to its idea and explored its alternatives, we can at least recognise
sympatric and possibly sister species S. menelas (Castelnau, the plausibility of my initial hypothesis, extinction. If this is
1840), was rarer for possibly having such an idiosyncratic indeed the case, causes that could have driven G. dichroa to
biology, and Cupello & Vaz-de-Mello (2018) made a simi- extinction are certainly multifactorial and likely included at
lar suggestion to explain why S. securus has been collected so least two of the elements discussed above, competition with
less frequently than S. seag. Perhaps this is also the case with sympatric species such as G. lacordairii and potential strict
G. dichroa and G. lacordairii. Indicating that G. lacordairii ecological specialisations. But chief among the factors is likely
is indeed more eurytopic and competitive than G. dichroa is the negative impact of the ongoing anthropogenic conversion
that it has a much broader geographical range and inhabits of habitats in southern and central South America. It may be,
a more diverse set of biomes than its rarer relative (Fig. 5; as briefly explored by Cupello & Vaz-de-Mello (2013), that
Cupello & Vaz-de-Mello 2013, 2015). the intense human pressure converting the forests and natural
But low abundance and stenotopy in themselves, while grasslands from Paraguay and southern Brazil to Argentina
explaining rarity in the field and collections, would still and Uruguay into urban space, pastures, and agricultural fields
leave unanswered the question of why G. dichroa has not over the second half of the 20th century, particularly from the
been collected since the 1950s as these factors did not pre- 1960s Green Revolution onwards (Roesch et al. 2009; Souza
vent the species from being collected before. An explanation et al. 2020; Ribeiro et al. 2021), impacted the populations of
could be that early-20th-century collectors applied collection the putatively stenotopic G. dichroa, and this drove a specialised
procedures more suitable for the capture of G. dichroa than and already fragile species to extinction or at least to a drastic
modern collectors do, such as the search for specimens in ant reduction in abundance and range. Gromphas lacordairii, in
nests. Even though, judging from its morphology, G. dichroa turn, a more eurytopic and, thus, adaptable species, has not
does not seem to be a myrmecophilous species, we know, for only resisted but continued to flourish. For the time being,
instance, that the likely collector of the MGAP series, Father however, the fate of G. dichroa will remain uncertain. Future
Pio Buck, used to search for beetles in ant nests (e.g., the fieldwork may, after all, rediscover the species living in some
type series of Ateuchus myrmecophilus (Boucomont, 1935) of the region’s last areas of natural grasslands, wetlands, river
was collected by him in an Acromyrmex lobicornis (Emery, sandbanks, floodplains, shrublands, and gallery forests, which,
1888) nest; see Boucomont 1935). It may also be that G. di- based on its widespread presence in the northern Pampas and
chroa is a saprophagous species attracted to decaying fruits the biology of congeneric species, are the candidate preferred
or fungi, and the current preferred usage of mammalian habitats of G. dichroa. This would be an exciting discovery,
excrement and carrion as bait (e.g., Morelli et al. 2002; da finally allowing us to better understand the biology and
Silva 2017; da Silva et al. 2008, 2009; da Silva & Di Mare conservation status of this intriguing and beautiful species.

51
 Cupello M.

Material examined. — Gromphas dichroa: Brazil. Rio Grande Largo; I.1939; no collector; MGAP • 1 unsexed specimen; Cerro
do Sul • 1 ♀; Cerro Largo; III.1941; no collector (but likely Pio Largo; I.1940; no collector; MGAP • 2 unsexed specimens; Cerro
Buck leg.); MGAP • 1 ♀; Nova Petrópolis; I.1928; no collector (but Largo; 1949; no collector; MGAP • 2 unsexed specimens; Cerro
likely Pio Buck leg.); MGAP. — Santa Catarina • 1 ♀; Itapiranga; Largo; I.1949; no collector; MGAP • 12 ♂, 14 ♀; Charqueadas, Fa-
XI.1934; no collector (but likely Pio Buck leg.); MGAP • 1 ♂; zenda Águas Claras; 12.I.1989; A. Ferreira leg.; MCNZ • 2 ♂, 2 ♀;
Itapiranga; X.1954; no collector (but likely Pio Buck leg.); MGAP. Cidreira, Salinas; 2.XI.1992; Lascombe leg.; MCNZ • 1 ♂; General
No data: • 1 unsexed specimen; ex J. J. Gillet collection; RBINS. Câmara (‘Gen Câmara’); 16.IX.1982; A. Lise leg.; MCNZ • 4 ♂,
Gromphas lacordairii: Argentina. Autonomous City of Buenos Aires 5 ♀; Imbé; II.1961; Buckup leg.; MCNZ • 1 ♂, 1 ♀; Imbé, Santa
• 1 unsexed specimen; Palermo; IV.1943; no collector; ex Martínez Teresinha Beach; II.1973; M. H. Galileo leg.; MCNZ; • 1 ♂; Imbé,
collection; CMNC. — Buenos Aires • 1 ♂; Boulogne Sur Mer; Santa Teresinha Beach; 6.XI.1999; L. Moura and I. Heydrich leg.;
XI.1941; no collector; ex Martínez collection; CMNC • 1 unsexed MCNZ • 3 unsexed specimens; Mostardas; I.1945; no collector;
specimen; Florencio Varela; I.1978; Giacomozzi leg.; CMNC • MGAP • 16 unsexed specimens; Nova Petrópolis; I.1928; no collec-
1 unsexed specimen; San Isidro; 14.IV.1956; H. J. Molinari leg; tor; MGAP • 5 unsexed specimens; Osório; 3.I.1950; no collector;
CMNC • 1 unsexed specimen; San Isidro; XII.1967; no collector; MGAP • 8 unsexed specimens; Osório; 24.I.1958; no collector;
ex Martínez collection; CMNC • 1 unsexed specimen; San Isidro; MGAP • 1 unsexed specimen; Osório; 24.I.1958; Pio Buck leg.;
XI.1973; no collector; ex Martínez collection; CMNC. — Chaco MGAP • 1 unsexed specimen; Osório; 25.I.1958; no collector;
• 1 ♂, 3 ♀; Presidencia de la Plaza; 26°59’S, 59°50’W; 7.X.2006; MGAP • 1 unsexed specimen; Osório; 25.I.1958; Pio Buck leg.;
M. Damborsky leg.; TAMU. — Corrientes • 1 unsexed specimen; MGAP • 1 ♀; Osório; 2.II.1964; P. Braun leg.; MCNZ • 1 unsexed
‘5 km W Ituzaingó’; IX.1979; no collector; ex Martínez collection; specimen; Pareci Novo; 20.X.1927; no collector; MGAP • 1 unsexed
CMNC • 2 unsexed specimens; Ituzaingó; III.1976; no collector; ex specimen; Pareci Novo; VIII.1935; no collector; MGAP • 1 unsexed
Martínez collection; CMNC. — Entre Ríos • 16 ♂, 7 ♀; Uruguay, specimen; Pareci Novo; XI.1938; no collector; MGAP • 1 unsexed
Molino, Pronunciamiento; I-III.1963; J. Foerster leg.; cow dung; specimen; Pelotas; II.1963; no collector; MGAP • 1 unsexed spe-
TAMU. — Misiones • 1 ♀; Cainguás, Dos de Mayo; I.1966; J. Fo- cimen; Porto Alegre; 16.X.1928; no collector; MGAP • 1 unsexed
erster leg.; carrion; TAMU. — Salta • 1 unsexed specimen; Embalse specimen; Porto Alegre; 29.IX.1929; no collector; MGAP • 1 un-
de Cabra Corral; 18.I.1984; Bordón leg.; ex Martínez collection; sexed specimen; Porto Alegre; 29.V.1934; no collector; MGAP • 1
CMNC • Guachipas; I.1948; no collector; ex Martínez collection; unsexed specimen; Porto Alegre; 15.XI.1935; no collector; MGAP
CMNC • 1 unsexed specimen; Guachipas, Alemanía; III.1973; no • 1 unsexed specimen; Porto Alegre; 1941; ‘T. L.’ leg.; MGAP; •
collector; ex Martínez collection; CMNC. — Santiago del Estero • 2 1 unsexed specimen; Porto Alegre; 1943; no collector; MGAP • 1
unsexed specimens; Río Hondo, Termas de Río Hondo; 16.II.1982; unsexed specimen; Porto Alegre; 25.IX.1949; no collector; MGAP
Henry Howden and Anne Howden leg.; CMNC. — Tucumán • • 1 unsexed specimen; Porto Alegre; 3.X.1956; no collector; MGAP
1 ♂; Yerba Buena, San Javier, 1200 m; 20.I.1970; Mielke leg.; DZUP. • 1 ♀; Porto Alegre; 6.XII.1962; P. Braun leg.; MCNZ • 1 unsexed
Bolivia. Santa Cruz • 1 unsexed specimen; Chiquitos, Roboré, San- specimen; Porto Alegre; 7.X.1963; no collector; MGAP • 1 un-
tiago de Chiquitos; XI.1959; no collector; ex Martínez collection; sexed specimen; Porto Alegre; 22.X.1963; no collector; MGAP •
CMNC. — Tarija • 1 ♀; between Yaguacua and Caiza; 21°50’52”S, 1 unsexed specimen; Porto Alegre; 1964; no collector; MGAP • 1
63°36’26”W; 620 m; 3.I.2005; Mann, Hamel and Herzog leg.; unsexed specimen; Porto Alegre; 26.I.1965; no collector; MGAP • 1
horse dung; Chaco transition zone, low lying pasture, open habitat, unsexed specimen; Porto Alegre; 7.II.1965; no collector; MGAP • 1
sandy-loam; TAMU. unsexed specimen; Porto Alegre; 7.X.1965; no collector; MGAP • 1
Brazil. Espírito Santo • 4 ♂, 1 ♀; Linhares; XI.1965; A. Maller leg.; unsexed specimen; Porto Alegre; 26.X.1966; no collector; MGAP •
DZUP • 2 ♂; Linhares; XII.1965; A. Maller leg.; DZUP. — Mato 1 ♀; Porto Alegre; 18.VIII.1970; no collector; MCNZ • 1 ♂; Porto
Grosso • 1 ♀; Santo Antonio de Leverger; XII.1965; W. D. Ed- Alegre; 8.XI.1993; V. L. C. Lascombe leg.; MCNZ • 3 unsexed spe-
monds leg.; meat trap; ‘capoeira’ vegetation; TAMU • 4 ♂ [1 with cimens; Porto Alegre, Belém Novo; I.1959; no collector; MGAP • 1
genitalia dissected], 1 ♀; Santo Antonio de Leverger; 4.XII.1965; unsexed specimen; Quaraí; I.1945; no collector; MGAP • 1 unsexed
W. D. Edmonds leg.; cattle dung; TAMU. — Mato Grosso do specimen; Santo Augusto; 12.X.1966; O. Roppa leg.; CMNC • 2 ♂;
Sul • 1 ♂; Corumbá; 17.X.1961; F. M. Oliveira leg.; DZUP. — São Jerônimo, Fazenda Santa Hermenêutica; 13.I.1989; A. Ferreira
Paraná • 1 ♂, 1 ♀; Araucária, Barigui; V.1941; no collector; ex F. leg.; MCNZ • 3 ♂, 3 ♀; São Jerônimo, Instituto Rio Grandense do
Justus Júnior collection; DZUP • 1 ♀; Curitiba; X.1961; S. Laroca Arroz (‘IRGA’); 11.I.1989; on dung; A. Ferreira leg.; MCNZ • 1
leg.; DZUP • 5 ♂, 7 ♀; Curitiba; 25.I.1962; S. Laroca leg.; DZUP unsexed specimen; São Leopoldo; XI.1958; no collector; MGAP
• 5 ♂, 5 ♀; Curitiba; 28.X.1964; Sebastião Laroca leg; DZUP • • 1 ♂; Torres; X.1956; T. de Lema leg.; MCZN • 1 ♂; Triunfo,
1 ♂, 1 ♀; Curitiba; 21.XI.1965; Mitchell and Graf leg.; DZUP • Braskem plant (stated on the label by its former name ‘COPESUL’);
1 ♂; Curitiba; 11.IV.1968; Gonçalves leg.; DZUP • 1 ♀; Curitiba; 21.IX.1989; M. Hoffmann leg.; MCNZ • 1 ♀; Triunfo, Braskem
900 m; 2.XI.1963; Departamento de Zoologia da Universidade plant (‘COPESUL’); 14-15.I.1997; A. Franceschini leg.; MCNZ •
Federal do Paraná leg. (‘Dept. ZOO. leg’); DZUP • 1 unsexed 2 ♀; Uruguaiana, Estância Corumbá; 18.XII.1996; N. S. Macedo leg.;
specimen; Curitiba, Cajuru; X.1938; Claretiano leg; MGAP • 1 MCNZ • 1 ♂, 1 ♀; Viamão; X.1956; L. Buckup and E. Buckup leg.;
unsexed specimen; Curitiba, Parolin (‘Parolim’); I.1937; Claretiano MCNZ • 1 ♀; Xangri-Lá, Atlântida Beach; 20.I.1968; no collector;
leg; MGAP • 1 ♀; Guaíra; X.1953; C. R. Leite leg.; ex Alvarenga MCNZ. — Santa Catarina • 1 unsexed specimen; ‘Emboebas’ (?);
collection; DZUP • 1 ♀; Ponta Grossa; VIII.1942; no collector; ex 3.IV.1953; no collector; MGAP • 1 ♀; Canoinhas, Pinhal; XII.1952;
F. Justus Júnior collection; DZUP • 1 ♂, 3 ♀; São José dos Pinhais; A. Maller leg.; ex Campos Seabra collection; TAMU• 1 unsexed
18.XI.1966; Laroca and Antoni leg.; DZUP. — Rio de Janeiro • specimen; Itapiranga; IX.1953; MGPA • 1 unsexed specimen;
1 ♀; Duque de Caxias, São Bento (‘S Bento’); XII.1954; P. A. Tales Itapiranga; V.1954; no collector; MGAP • 1 unsexed specimen;
leg.; ex Alvarenga collection; DZUP • 1 ♀; Rio de Janeiro, Botafogo; Joinville; VI.1949; no collector; MGAP • 1 unsexed specimen; São
X.1957; Alvarenga leg.; ex Alvarenga collection; DZUP • 1 unsexed Bento do Sul; I.1989; no collector; CMNC • 1 unsexed specimen;
specimen; Rio de Janeiro, Copacabana; XII.1994; R. L. Vaz de Mello Seara, Nova Teutônia; 25.IX.1963; F. Plaumann leg.; CMNC • 1
leg.; CMNC. — Rio Grande do Sul • 3 ♂, 1 ♀; Cachoeira do Sul; unsexed specimen; Seara, Nova Teutônia; 23.X.1963; F. Plaumann
II.1996; H. Schaurich leg.; MCNZ • 1 ♂; Capão da Canoa, Curu- leg; CMNC • 1 unsexed specimen; Seara Nova Teutônia; 2.I.1964;
mim; 22.I.1978; C. J. Becker leg.; MCNZ • 1 ♀; Capão da Canoa; F. Plaumann leg.; CMNC • 1 unsexed specimen; Seara, Nova Teu-
9.X.1989; M. A. Santos leg.; MCNZ • 3 ♂, 4 ♀; Capão da Canoa; tônia; 30.I.1964; F. Plaumann leg.; CMNC • 6 unsexed specimens;
18.XI.1989; M. A. Santos leg.; MCNZ • 1 unsexed specimen; Cerro Seara, Nova Teutônia; XII.1970; no collector; CMNC • 1 unsexed
Largo; I.1931; no collector; MGAP • 1 unsexed specimen; Cerro specimen; Seara, Nova Teutônia; 300-500 m; II.1970; no collector;

52 ZOOSYSTEMA • 2024 • 46 (2)

CMNC • 1 ♂; Seara, Pinhal, 700 m; IV.1958; no collector; DZUP MCNZ Gromphas and illuminated my discussion on the dis-
• 1 ♂; Seara, Pinhal, 700 m; XII.1958; no collector; DZUP. — São appearance of G. dichroa with some important facts about the
Paulo • 1 unsexed specimen; ‘CIPO’ (?); 6.II.1973; V. N. Alin leg.;
CMNC • 1 unsexed specimen; ‘CIPO’ (?); 12.XII.1973; V. N. Alin
geography of southern Brazil. Patricia González-Vainer shared
leg.; CMNC • São Paulo, Santo Amaro (‘S. Amaro’); 20.XII.1953; information on Uruguayan populations of G. lacordairii. Luísa
no collector; MGAP. Menezes da Silveira was responsible for digitising the label data
Paraguay. Alto Paraguay • 1 ♀; Puerto la Esperanza; 2.XII.2006; of the MGAP G. lacordairii and for producing Figure 11D.
C. Aguilar Julio leg.; CAPC. — Caaguazú • 1 unsexed specimen; Paul Skelley is the author of Figure 8, and Olivier Montreuil,
Caaguazú; XII.1948; F. H. Schade leg.; CMNC. — Caazapá
• 1 unsexed specimen; Caazapá; 1-8.XII.1990; G. Arriagada leg.;
Antoine Mantilleri and Christophe Rivier are responsible for
CMNC. — Capital District • 1 ♂; Asunción; 26.II.1992; J. A. Ko- Figures 4, 6. Alvaro Mones, Miguel A. Monné and Walter Sosa
chalka leg.; MNHNP • 1 unsexed specimen; Asunción, Mburicaó; assisted me in my ultimately fruitless search for the G. dichroa
12.X.1990; G. Arriagada leg.; CMNC • 1 unsexed specimen; Asun- specimens studied by Luis Barattini and Aberlado Sáenz for
ción, Mburicaó; 7.I.1991; G. Arriagada leg.; CMNC. — Central their 1960s revision of the genus. Federico Ocampo was gen-
• 1 ♂; Ypacaraí (‘1000 Mts. al E. de / YPACARAÍ’); 23.X.1996; D.
Moyano leg.; CAPC • 1 ♂; Ypacaraí (‘1000 Mts. al E. de / YPA- erous enough to allow me to publish data from specimens on
CARAÍ’); 23.X.1996; D. Moyano leg.; MNHNP. — Concepción loan to him, including the holotype of G. dichroa, and to cite
• 1 ♀; San Lázaro, Vallemí; 26.XI.2006; C. Aguilar leg.; CAPC. — his unpublished field notes. Alejandro Lopera confirmed my
Guairá • 1 ♂; Cordillera del Ybytyruzú, Carpa Cué; 24.XII.1994; suspicion about the likely erroneous state of the Muzo record
Aguilar leg.; MNHNP • 2 ♀; Cordillera del Ybytyruzú, Cerro Akatí; for G. amazonica, and Jorge A. Noriega and Juan Pablo Botero
19.II.2004; no collector; CAPC • 1 unsexed specimen; Mbocayaty
(‘Bcayaty’); 23.IX.1946; Schade leg.; CMNC. • 1 unsexed specimen; clarified some further dubious Colombian records. Stéphane
Villarrica; X.1941; Schade leg.; CMNC. — Itapúa • 1 ♀; eastern Boucher and Olivier Montreuil advised me on the history of
portion of Isla Yacyretá, Paraná River [Butia savanna]; 14.XII.1999; the MNHN collections, while Alain Drumont photographed
J. A. Kochalka leg.; MNHNP. — Paraguarí • 1 unsexed specimen; the sole RBINS G. dichroa illustrated in Figure 11B, C and dis-
Parque Nacional Ybycuí; 13.I.1996; C. Aguilar J. leg.; CMNC. — pelled some of my doubts about Lacordaire’s beetle collection.
San Pedro • 2 unsexed specimens; Cororo, Río Ypané; II.1979;
no collector; Martínez leg.; CMNC • 2 ♂; Nueva Germania; 15- Max Barclay provided information on the possible identity
20.I.1992; Aguilar leg.; MNHNP. of the eponymous Lemoine of C. O. Waterhouse’s Gromphas
Uruguay. Canelones • 1 ♂; Pando; 10.I.2005; no collector; CAPC lemoinei. Pedro Giovâni da Silva allowed me to use the data of
• 6 unsexed specimens; Ruta 82, Km 46 500, Campo Experimental G. lacordairii specimens he gathered in small collections across
Instituto de Higiene; 10.XI.2020; unknown collector; cow dung; Rio do Grande do Sul state. Finally, Andrew Polaszek (Editor
FCUR. — Cerro Largo • 1 ♀; no further locality or date; Sáenz
leg.; TAMU. — Montevideo • 1 unsexed specimen; no further in Chief, Journal of Natural History) and Henrietta Thomson
data; MGAP. — Tacuarembó • 1 unsexed specimen; Ruta 43, on (Taylor and Francis Group) are thanked for the information
the banks of Río Negro near San Gregorio de Polanco; 32°38’43”S, about the publication date of the online and printed versions
55°47’45”W; 20.X.2019; unknown collector; cow dung; FCUR. of my JNH Gromphas paper. My travel to Ottawa, when I had
Likely erroneous data: Bolivia. Santa Cruz • 1 ♀; Ichilo, Buena the opportunity to meet Patrice Bouchard at the CNCI, was
Vista; 380 m; XI.1968; F. Steinbach leg.; TAMU.
Unknown wild origin • 1 ♂; reared from a laboratory colony at funded by a Canadian Museum of Nature Visiting Scientist
Texas A&M University, College Station, TX, USA; summer 1981; Award, and François Génier, Robert Anderson, and Andrew
ex G. T. Fincher collection; TAMU • 1 ♂; same data as for preced- Smith warmly hosted me while at the CMNC. My visits to
ing but the date; XI.1981; ex G. T. Fincher collection; TAMU; 1 ♀ the MNHN as well as to several other European museums
• same data as for preceding but the date; 1982; ex G. T. Fincher were partly funded by an Ernst Mayr Travel Grant (Museum
collection; TAMU.
No data • 2 ♂, 8 ♀; DZUP • 12 unsexed specimens; MGAP. of Comparative Zoology, Harvard University, USA) and an
Oxford University Museum of Natural History Visiting Fel-
lowship. I extend my gratitude to all the curators who received
Acknowledgements me during my studies of the collections under their care. The
I am especially grateful to Patrice Bouchard for bringing to my librarians at the NHM and CNCI as well as Vinicius S. Fer-
attention Lacordaire’s memoir of his South American beetle explo- reira, Alvaro Mones, Juan Pablo Botero, and the Biodiversity
rations and Oken’s review of it. Patrice also had the interest and Heritage Library are thanked for granting me access to literature
patience to discuss most of the nomenclatural issues addressed in that would be otherwise unavailable. I deeply appreciate the
this paper and to point out my mistakes. Equally appreciated is careful reading and criticism of different parts and versions of
the help provided by Neal Evenhuis, Miguel Alonso-Zarazaga, this work by Frank-Thorsten Krell, Patrice Bouchard, Michele
and François Génier in interpreting the intricate nomenclatural Rossini, Auke Hielkema, W. D. Edmonds, Juan Pablo Botero,
problems I was dealing with, even though some of them may Victor Moctezuma, Cibele S. Ribeiro-Costa, Pedro Giovâni
not agree with my conclusions. Any mistakes are naturally my da Silva, Elie Saliba, an anonymous reviewer, and the editor
own. Neal Evenhuis also kindly shared with me his unpublished Annemarie Ohler. I was funded by a PhD scholarship granted
dating of Dictionnaire universel d’Histoire naturelle, Voyage dans by the Brazilian Conselho Nacional de Desenvolvimento Científico
l’Amérique méridionale, and Isis von Oken. Many other colleagues e Tecnológico (CNPq) during the writing of the manuscript. I
assisted me in completing this work. Adriano B. Kury and Álvaro am further thankful to my post-doctoral supervisor, prof. Ci-
Alfredo Bragança Júnior helped me investigate the publication bele S. Ribeiro-Costa, for allowing me to enjoy the facilities of
date and editorship of Perty’s Delectus animalium articulatorum. the Laboratório de Sistemática e Bioecologia de Coleoptera, at the
Luciano de Azevedo Moura sent me detailed label data of the Federal University of Paraná.

53
 Cupello M.

REFERENCES with publication dates and notes. Zookeys 583: 1-776. https://
doi.org/10.3897/zookeys.583.7084
Alonso C. B. G., Zurita G. A. & Bellocq M. I. 2019. — Live- Bousquet Y. & Bouchard P. 2013. — The genera in the second
stock areas with canopy cover sustain dung beetle diversity in catalogue (1833-1836) of Dejean’s Coleoptera collection. Zookeys
the humid subtropical Chaco forest. Insect Conservation and 282: 1-219. https://doi.org/10.3897/zookeys.282.4401
Diversity 12 (4): 296-308. https://doi.org/10.1111/icad.12340 Brown R. W. 1954. — Composition of scientific words. A manual of
Anonymous 1897. — Séance du 14 avril 1897. Présidence M. A. methods and a lexicon of materials for the practice of logotechnics.
Grouvelle. Nécrologie. Bulletin de la Société entomologique de Smithsonian Books, Washington, D. C., 882 p. https://archive.
France [year 1897]: [129]. https://www.biodiversitylibrary.org/ org/details/compositionofsci00brow
page/9210617 Brown Jr. W. L. & Wilson E. O. 1956. — Character displacement.
Bailly A. 1895. — Dictionnaire grec-français. Hachette, Paris, 2193 p. Systematic Zoology 5: 49-64. https://doi.org/10.2307/2411924
Barattini L. P. & Sáenz A. 1961. — Contribución al conocimiento Bruch C. 1911. — Catálogo sistemático de los coleópteros de
de las especies del género Gromphas Brullé 1854 [sic] (Col. la República Argentina. Pars IV. Familias Lucanidae, Scara-
Scarab.). Actas y Trabajos del Primer Congreso Sudamericano de baeidae (Coprini-Cetonini), Passalidae. Revista del Museo de
Zoología 3: 21-29. [publication date according to the colophon: La Plata XVII (IV): 181-225. https://www.biodiversitylibrary.
8th September 1961 (Alvaro Mones, personal communication, org/page/53896861
24th April 2021)]. Brullé A. 1838. — [livraison 8: p. 225-448], in Brullé A. 1837-
Barratini [sic] L. P. & Sáenz A. 1964. — Contribution a l’étude 1838. Histoire naturelle des insectes, traitant de leur organisation
des espèces du genre Gromphas (Brullé) [Col. Scarabaeidae]. et de leurs moeurs en général, par M. V. Audouin, et comprenant
Revue française d’Entomologie 31 (3): 173-181. leur classification et la description des espèces, par M. A. Brullé. Le
Bates H. W. 1863. — The Naturalist on the River Amazons, a Record tout accompagné de planches gravées sur acier, d’après des peintures
of Adventures, Habits of Animals, Sketches of Brazilian and Indian exécutées pour cette édition sur la collection du Muséum de Paris.
Life, and Aspects of Nature under the Equator, During Eleven Years Tome VI. Coléoptères III. F. D. Pillot, Paris, 448 p. + pls. 18, 19,
of Travel. John Murray, London, in two volumes, Vol. 1, 351 p., 19bis, 20, 25 [publication date for nomenclatural purposes: 31st
Vol. 2, 423 p. https://doi.org/10.5962/bhl.title.21335 of December 1838 (based on Bousquet 2016)] https://www.
Bates H. W. 1869. — [proceedings from November 1, 1869]. biodiversitylibrary.org/page/25546370
Proceedings of the Entomological Society of London [year 1869]: Burmeister H. 1874. — Lamellicornia Argentina. Entomologische
xix-xxiii. https://www.biodiversitylibrary.org/page/14787693 Zeitung 35: 120-133. [publication date for nomenclatural pur-
Bates H. W. 1870. — On a new genus and some new species of poses: 30th June 1874 (based on the range ‘April-Juni’ provided
Copridæ (Coleoptera-Lamellicornia). The Transactions of the at the end of the volume on page 444)] https://www.biodiversi-
Entomological Society of London 18: 173-180. [publication date tylibrary.org/page/9236010
for nomenclatural purposes: 30th June 1870 (dated ‘June’ on Cabrera-Walsh G. & Cordo H. A. 1997. — Coprophilous
the article’s first page; I follow Article 21.3.1 of the Code to arthropod community from Argentina with species of potential
determine the precise day of publication for nomenclatural use as biocontrol agents against pest flies. Environmental Ento-
purposes)] https://www.biodiversitylibrary.org/page/14719377 mology 26 (2): 191-200. https://doi.org/10.1093/ee/26.2.191
Bitencourt B. S., Dimas T. M., da Silva P. G. & Morato E. F. Cajaiba R. L., Périco E., Dalzochio M. S., da Silva W. B., Bas-
2019. — Forest complexity drives dung beetle assemblages along tos R., Cabral J. A. & Santos M. 2017. — Does the composi-
an edge-interior gradient in the southwest Amazon rainforest. tion of Scarabaeidae (Coleoptera) communities reflect the extent
Ecological Entomology electronic-only detached version: 1-10. of land use changes in the Brazilian Amazon? Ecological Indicators
https://doi.org/10.1111/een.12795 74: 285-294. https://doi.org/10.1016/j.ecolind.2016.11.018
Blackwelder R. E. 1944. — Checklist of the coleopterous insects Cambefort Y. 2006. — Des coléoptères, des collections et des hommes.
of Mexico, Central America, the West Indies and South America. Paris, Muséum national d’Histoire naturelle, 2006, 375 p.
Part 2. United States National Museum Bulletin 185: 189-341. (Archives ; 10).
https://www.biodiversitylibrary.org/page/7876519 Candèze [E. C. A.] 1872. — Notice sur J.-Théodore Lacordaire,
Blackwelder R. E. 1957. — Checklist of the coleopterous insects associé de l’Académie, né à Recey-sur-Ource (France) le 1er février
of Mexico, Central America, West Indies and South America. 1801, mort à Liége le 18 juillet 1871. Annuaire de l’Académie
Part 6. United States National Museum Bulletin 185: [v]-vii, royale des Sciences, des Lettres et des Beaux-Arts de Belgique 38:
927-1492. https://www.biodiversitylibrary.org/page/7877283 unpaged [Lacordaire’s engraving] + [139]-160.
Blanchard É. 1846. — [livraison 80: 169-184, pl. 31], in Blan- Canziani C. & González-Vainer P. 2022. — Structure and
chard É. & Brullé G. A. 1835-1846. Voyage dans L’Amérique composition of dung beetle assemblages (Coleoptera: Scara-
Méridionale (le Brésil, la République orientale de L’Uruguay, la baeidae) in a livestock ranch in central Uruguay: Responses
République Argentine, la Patagonie, la République du Chili, la of functional groups and species to local habitats and trophic
République de Bolivia, la République du Pérou), exécuté pendant resources. The Coleopterists Bulletin 76 (3): 407-418. https://doi.
les années 1826, 1827, 1828, 1829, 1830, 1831, 1832 et 1833, org/10.1649/0010-065X-76.3.407
par Alcide d’Orbigny. Tome Sixième. 2e partie : Insectes. P. Bertrand, Constantin R. 2012. — Mémorial des coléoptéristes français.
Paris, and Ve. Lerrault, Strasbourg, 222 p. [publication date Retirage de décembre 2012. Bulletin de Liaison de l’Association des
for nomenclatural purposes: 18th December 1846 (Evenhuis, Coléoptéristes de la Région Parisienne 14 (supplement): [1]-[116].
personal communication, 3rd September 2019)] https://www. Cupello M. 2013. — Case 3612. Onitis aeruginosus Klug, 1855
biodiversitylibrary.org/page/2531073 (Insecta, Coleoptera, Scarabaeidae): proposed conservation of
Bouchard P., Bousquet Y., Davies A. E., Alonso-Zarazaga the specific name. Bulletin of Zoological Nomenclature 70: 15-18.
M. A., Lawrence J. F., Lyal C. H. C., Newton A. F., Reid https://doi.org/10.21805/bzn.v70i1.a4
C. A. M., Schmitt M., ŚlipiŃski A. & Smith A. B. T. 2011. — Cupello M. 2020. — The discovery of Edgar von Harold type mate-
Family-group names in Coleoptera (Insecta). Zookeys 88: 1-972. rial in the Museum of Zoology, Dresden. Scarabaeus 1: 15-24.
https://doi.org/10.3897/zookeys.88.807 Cupello M. & Vaz-de-Mello F. Z. 2013. — Taxonomic revi-
Boucomont A. 1935. — Description de trois Coprides de l’Amérique sion of the South American dung beetle genus Gromphas Brullé,
du Sud. Miscellanea Entomologica 36 (2): 9-10. 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini: Grom-
Bousquet Y. 2016. — Litteratura Coleopterologica (1758-1900): phadina). Zootaxa 3722 (4): 439-482. https://doi.org/10.11646/
a guide to selected books related to the taxonomy of Coleoptera zootaxa.3722.4.2

54 ZOOSYSTEMA • 2024 • 46 (2)

Cupello M. & Vaz-de-Mello F. Z. 2014. — Correction of the Dubois A., Bour R. & Ohler A. 2015a. — What is an online
type species of the South American genus Gromphas Brullé, 1837 ‘preliminary version’ of a publication in the meaning of Article
(Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Zootaxa 9.9 of the Code? – One more step on the trail of the Asian ele-
3790 (2): 399-400. https://doi.org/10.11646/zootaxa.3790.2.7 phant. Bulletin of Zoological Nomenclature 72 (1): 6-18. https://
Cupello M. & Vaz-de-Mello F. Z. 2015. — A new species and doi.org/10.21805/bzn.v72i1.a13
the phylogeny of the South American genus Gromphas Brullé, Dubois A., Bour R. & Ohler A. 2015b. — Nomenclatural avail-
1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Journal ability of preliminary electronic versions of taxonomic papers:
of Natural History electronic-only detached version: 1-27. [pub- in need of a clear definition. Bulletin of Zoological Nomenclature
lication data for nomenclatural purposes: 12th October 2015 72 (3): 252-265. https://doi.org/10.21805/bzn.v72i3.a1
(information available at the journal’s website: https://doi.org/1 Ecker A. 1883. — Lorenz Oken, a biographical sketch. Or ‘in memo-
0.1080/00222933.2015.1091099)] [volume-integrated version riam’ of the centenary of his birth read before the fifty-second meeting
published in 2016 ‒ on or after 15th March, the day it was sent of the German Association for the Advancement of Science at Baden-
to print (Henrietta Thomson, Taylor & Francis Group, personal Baden, September 20, 1879. With explanatory notes selections from
communication to MC, 6th February 2020) ‒, as volume 50 (15- Oken’s correspondence and a portrait of the professor. Kegan Paul,
16): 943-969] https://doi.org/10.1080/00222933.2015.1091099 Trench & Co., London, xxiv + 183 + 47 p.
Cupello M. & Vaz-de-Mello F. Z. 2018. — A monographic Edmonds W. D. 2000. — Revision of the Neotropical dung beetle
revision of the Neotropical dung beetle genus Sylvicanthon genus Sulcophanaeus (Coleoptera: Scarabaeidae: Scarabaeinae).
Halffter & Martínez, 1977 (Coleoptera: Scarabaeidae: Scarabaei- Folia Heyrovskyana, Supplementum 6: 1-60.
nae: Deltochilini), with a reappraisal of the taxonomic history of Escobar H. 2018 — In a ‘foretold tragedy’, fire consumes Brazil
‘Canthon sensu lato’. European Journal of Taxonomy 467: 1-205. museum. Science 361 (6406): 960. https://doi.org/10.1126/
https://doi.org/10.5852/ejt.2018.467 science.361.6406.960
Cupello M., Ribeiro-Costa C.S & Vaz-de-Mello F. Z. 2021. — Evenhuis N. L. 1997a. — Litteratura taxonomica dipterorum (1758-
The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: 1930). Vol. I. A-K. Backhuys, Leiden, 426 p.
Phanaeini): its phylogenetic significance, geographical polychroma- Evenhuis N. L. 1997b. — Litteratura taxonomica dipterorum (1758-
tism and the subspecies problem. Zoological Journal of the Linnean 1930). Volume II. L-Z. Backhuys, Leiden: 427-871.
Society electronic-only detached version: 1-62 [volume-integrated Evenhuis N. L. 2003. — Publication and dating of the jour-
version published in March 2022, volume 194 (3): 973-1034.] nals forming the Annals and Magazine of Natural History and
https://doi.org/10.1093/zoolinnean/zlab015 the Journal of Natural History. Zootaxa 385: 1-68. https://doi.
Cupello M., Iannuzzi L. & Vaz-de-Mello F. Z. 2022. — On org/10.11646/zootaxa.385.1.1
the authorship of Canthon rutilans cyanescens ‘Harold, 1868’ Evenhuis N. L. 2015. — [dating of] Berliner Entomologische
(Coleoptera, Scarabaeidae, Scarabaeinae, Deltochilini): the treat- Zeitschrift (1857-1876). Available online at: http://hbs.bish-
ment of varieties and nomina nuda by a 19th-century zoologist. opmuseum.org/dating/berlentzeit.html (last accessed on 4th
Bionomina 27: 67-87. https://doi.org/10.11646/bionomina.27.1.4 February 2020)
Damborsky M. P., Bohle M. C. A., Polesel M. G. I., Porcel Feer F. 2000. — Les coléoptères coprophages et nécrophages (Scara-
E. A. & Fontana J. L. 2015. — Spatial and temporal variation baeidae s. str. et Aphodiidae) de la forêt de Guyane Française:
of dung beetle assemblages in a fragmented landscape at eastern Composition spécifique et structure des peuplements. Annales de
humid chaco. Neotropical Entomology 44: 30-39. https://doi. la Société Entomologique de France (N. S.) 36 (1): 29-43.
org/10.1007/s13744-014-0257-2 Gahan C. J. & Arrow G. J. 1903. — List of the Coleoptera col-
Dejean P. F. M. A. 1821. — Catalogue de la collection de coléoptères de lected by Mr. A. Robert at Chapada, Matto Grosso (Percy Sladen
M. le Baron Dejean. Crevot, Paris, I-VIII + 136 p. + 2 p. [errata] Expedition to Central Brazil). Proceedings of the Zoological Society
[publication date for nomenclatural purposes: 12th May 1821 of London 1903 (2): 244-258.
(see Bousquet 2016)] https://doi.org/10.5962/bhl.title.11259 Gámez J. & Acconcia R. 2018. — Escarabajos coprófagos y
Dejean P. F. M. A. 1833. — [livraison 2: 97-176], in Dejean necrófagos (Scarabaeidae: Scarabaeinae: Phanaeini) del flanco
P. F. M. A. 1833-1836. Catalogue des Coléoptères de la collection andino llanero, piedemonte y llanos, con el reporte de nuevos
de M. le Comte Dejean. Méquignon-Marvis Père et Fils, Paris, registros para los estados de Barinas y Mérida (Venezuela). Revista
443 p. [publication date for nomenclatural purposes: 27th July Gaditana de Entomología 9 (1): 65-75.
1833 (see Bousquet 2016)] https://doi.org/10.5962/bhl.title.8771 Glare P. G. W. (ed.) 1968-1982. — Oxford Latin Dictionary.
Dejean P. F. M. A. 1836. — [livraisons 1-4: 1-384], in Dejean Clarendon Press, Oxford, xxiii + 2126 p.
P. F. M. A. 1836-1837. Catalogue des Coléoptères de la collection Giachino P. M. 1982. — Cataloghi III – Collezione Coleotterologica
de M. le Comte Dejean. Troisième édition, revue, corrigée et aug- di Massimiliano Spinola. Museo Regionale di Scienze Naturali,
mentée. Méquignon-Marvis Père et Fils, Paris, v-xiv + 503 p. Turin, 616 p.
[publication date for nomenclatural purposes: 30th July 1836 Gillet J. J. E. 1911. — Coleopterorum Catalogus. Pars 38: Scara-
(see Bousquet 2016)] baeidae: Coprinae I. W. Junk, Berlin, 100 p.
Deschodt C. M., Harrison J. G. & Sole C. L. 2021. — Redis- Groll E. K. 2016. — Lemoine, Victor, in Anonymous (ed.)
covery of Scarabaeus sevoistra Alluaud, 1902 (Coleoptera: Scara- Biographies of the entomologists of the world. Available online at:
baeinae): biological notes and IUCN Red Listing. Bothalia 51 (2): https://sdei.senckenberg.de/biographies/
113-119. https://hdl.handle.net/10520/ejc-bothalia_v51_n2_a10 Hamel-Leigue A. C., Mann D. J., Vaz-de-Mello F. Z. & Herzog
Dubois A., Crochet P.-A., Dickinson E. C., Nemésio A., S. K. 2006. — Hacia un inventario de los escarabajos peloteros
Aescht E., Bauer A. M., Blagoderov V., Bour R., de Car- (Coleoptera: Scarabaeidae) de Bolivia: primera compilación de
valho M. R., Desutter-Grandcolas L., Frétey T., Jäger P., los géneros y especies registrados para el país. Revista Boliviana
Koyamba V., Lavilla E. O., Löbl I., Louchart A., Malé- de Ecología y Conservación Ambiental 20: 1-18.
cot V., Schatz H. & Ohler A. 2013. — Nomenclatural and Hamel-Leigue A. C., Herzog S. K., Mann D. J., Larsen T. H.,
taxonomic problems related to the electronic publication of new Gill B. D., Edmonds W. D. & Spector S. 2009. — Dis-
nomina and nomenclatural acts in zoology, with brief comments tribución e historia natural de escarabajos coprófagos de la tribu
on optical discs and on the situation in botany. Zootaxa 3735: Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae) en Bolivia.
1-94. https://doi.org/10.11646/zootaxa.3735.1.1 Kempffiana 5 (2): 43-95.
Dubois A. 2000. — Synonymies and related lists in zoology: General Hardin G. 1960. — The competitive exclusion principle. Science
proposals, with examples in herpetology. Dumerilia 4 (2): 33-98. 131: 1292-1297. https://doi.org/10.1126/science.131.3409.1292

55
 Cupello M.

Harold E. 1859. — Beiträge zur Kenntnis einiger coprophagen world catalogue of harvestmen (Arachnidae, Opiliones). Version
Lamellicornien. Berliner Entomologische Zeitschrift 3: [193]-224. 1.0 – Checklist of all valid nomina in Opiliones with authors and
[publication date for nomenclatural purposes: 31st December dates of publication up to 2018. Adriano B. Kury, Rio de Janeiro,
1859 (based on Evenhuis 2015)] https://www.biodiversitylibrary. [ii] + 237 p.
org/page/9204087 Lacordaire J. T. 1830. — Mémoire sur les habitudes des Insectes
Harold E. 1869a. — Catalogus coleopterorum. Hucusque descriptorum, Coléoptères de l’Amérique méridionale. Annales des Sciences
synonymicus et systematicus. Tom. IV. Scarabaeidae. E. H. Gummi, naturelles 20: 185-291. https://www.biodiversitylibrary.org/
Munich: 979-1346 + [8]. [publication date for nomenclatural page/6069943
purposes: 27th May 1969 (based on Bousquet 2016)] https:// Lacordaire J. T. 1855. — Histoire naturelle des insectes. Genera
www.biodiversitylibrary.org/page/9641549 des coléoptères, ou exposé méthodique et critique de tous les genres
Harold E. 1969b. — Ueber coprophage Lamellicornien mit proposés jusqu’ici dans cet ordre d’insectes. Tome troisième. Librai-
besonderer Berücksichtigung der Pariser Sammlungen. Coleop- rie Encyclopédique de Roret, Paris, 594 p. [publication date
terologische Hefte 5: [46]-70. [publication date for nomenclatu- for nomenclatural purposes: 29th October 1855 (see Bousquet
ral purposes: 1st November 1869. This dating is based on the 2016); publication date of plate 27 depicting ‘Gromphas lacor-
November 1st record by Bates (1869) in the Proceedings of the dairei Blanch.’ is unclear and may have been different from
Entomological Society of London of the donation to the society the corresponding text. Here, for the sake of simplicity, I have
by Edgar von Harold of the fifth volume of Coleopterologische accepted that both were published together.] https://www.bio-
Hefte. This donation likely occurred at some point between July diversitylibrary.org/page/9612631
and November 1869, as the previous proceedings date from 5th Lemoine [V.] 1897. — Sur l’application des rayons Rœntgen à l’étude
July 1869 (Smith 1869). Had Harold donated the fifth volume des Articulés. Bulletin de la Société Entomologique de France [year
of Coleopterologische Hefte before July, the volume would have 1897]: 87-89. https://www.biodiversitylibrary.org/page/9210571
probably been recorded in the ‘additions to the library’ section Lewis C. T. & Short C. 1891. — Harpers’ Latin dictionary. A new
of the July proceedings rather than the one from November.] Latin dictionary founded on the translation of Freund’s Latin-German
https://www.biodiversitylibrary.org/page/16069721 lexicon edited by E. A. Andrews, LL. D. Revised, enlarged, and in
Heyne A. & Taschenberg O. 1908. — Die exotischen Käfer in great part rewritten. Harper & Brothers, Franklin Square, New
Wort und Bild. Verlag von J. F. Schreiber, Esslingen and Munich, York, and Clarendon Press, Oxford, xiv + [2019] p.
262 p. + I-L p. + 39 plates. Liddell H. G. & Scott R. 1897. — A Greek-English lexicon.
Hielkema A. 2023. — Leaf litter deposits in spiny palms: A hideout Revised and augmented throughout with the cooperation of professor
for “rare” scarabs? Scarabaeus 3: 3-6. Drisler, of Columbia College, New York. Eighth Edition. Harper &
Horn W., Kahle I., Friese G. & Gaedike R. 1990a. — Collectiones Brothers, New York, New York, xvi + 1778 p.
entomologicae. Ein Kompendium über den Verbleib entomologischer Madge R. B. 1988. — The publication dates of Dejean’s catalogues.
Sammlungen der Welt bis 1960. Teil I: A bis K. Akademie der Archives of Natural History 15 (3): 317-321.
Landwirtschaftswissenschaften der Deutschen Demokratischen Maldaner M. E., Cupello M., Ferreira D. C. & Vaz-de-
Republik, Berlin: 1-220. Mello F. Z. 2017. — Type specimens and names assigned to
Horn W., Kahle I., Friese G. & Gaedike R. 1990b. — Collectiones Coprophanaeus (Megaphanaeus) d’Olsoufieff, 1924, the largest New
entomologicae. Ein Kompendium über den Verbleib entomologischer World dung beetles (Coleoptera: Scarabaeidae: Phanaeini). Zootaxa
Sammlungen der Welt bis 1960. Teil II: L bis Z. Akademie der 4272 (1): 83-102. https://doi.org/10.11646/zootaxa.4272.1.4
Landwirtschaftswissenschaften der Deutschen Demokratischen Martínez A. 1959. — Catálogo de los Scarabaeidae Argentinos.
Republik, Berlin: 221-573. Revista del Museo Argentino de Ciencias Naturales “Bernardino
Iack-Ximenes G. E., de Vivo M., Percequillo A. R. 2005. — Rivadavia”, Ciencias Zoológicas 5: 1-126.
A new species of Echimys Cuvier, 1809 (Rodentia, Echimyidae) Mayr E. 1969. — Principles of systematic zoology. McGraw-Hill,
from Brazil. Papéis Avulsos de Zoologia 45 (5): 51-60. https://doi. New York, New York, xi + 428 p.
org/10.1590/S0031-10492005000500001 Mayr E. & Ashlock P. D. 1991. — Principles of systematic zoology.
International Commission on Zoological Nomenclature. Second edition. McGraw-Hill, New York, New York, and various
1999. — International code of zoological nomenclature. Fourth other cities, xx + 475 p.
edition adopted by the International Union of Biological Sciences. Mayr E., Linsley E. G. & Usinger R. L. 1953. — Methods and
International Trust for Zoological Nomenclature, London, 306 p. principles of systematic zoology. McGraw-Hill Book Company,
https://www.iczn.org/the-code/the-code-online/ New York, New York, Toronto and London, 328 p.
IUCN Species Survival Commission. 2012. — IUCN Red List Medina C. A. C., Medina-Uribe C. A., Quintero B. G. M.,
categories and criteria. Version 3.1. Second Edition. International Escobar-Villa A. F., Chuaire L. M. C. & Posada N. J. B.
Union for Conservation of Nature, Gland, iv + 32 p. https:// 2012. — Escarabajos coprófagos (Scarabaeinae) del eje cafetero: Guía
portals.iucn.org/library/node/10315 para el estudio ecológico. Anonymous editor, Villa María, 196 p.
Janssens A. 1940. — Contribution a l’Étude des Coléoptères lamel- Monné M. A. 2005. — Catalogue of the Cerambycidae (Coleop-
licornes coprophages. II. ‒ Remarques sur quelques Phanaeides. tera) of the Neotropical region. Part II. Subfamily Lamiinae.
Bulletin du Musée Royal d’Histoire Naturelle de Belgique 16 (34): 1-7. Zootaxa 1023: 1-760. https://doi.org/10.11646/zootaxa.1023.1.1
Judulien F. 1899. — Quelques notes sur plusieurs coprophages de Morelli E., Gonzalez-Vainer P. & Baz A. 2002. — Copropha-
Buenos Aires. Revista del Museo de La Plata 9: 371-380. https:// gous beetles (Coleoptera: Scarabaeoidea) in Uruguayan prairies:
www.biodiversitylibrary.org/page/9644378 abundance, diversity and seasonal occurrence. Studies on Neo-
Kertesz G. A. 1986. — Notes on Isis von Oken, 1817-1848. Isis tropical Fauna and Environment 37 (1): 53-57.
77 (3): 497-503. https://doi.org/10.1086/354208 Nedel L. B. 2005. — Breviário de um museu mutante. Horizontes
Kohlmann B. 1991. — Dung beetles in subtropical North America, Antropológicos 11 (2): 87-112. https://doi.org/10.1590/S0104-
in Hanski I. & Cambefort Y. (eds) Dung beetle ecology. Prince- 71832005000100006
ton University Press, Princeton: 116-132. Noriega J. A., Santos A. M. C., Calatayud J., Chozas S. &
Krell F.-T. 2015. — A mixed bag: when are early online publications Hortal J. 2021. — Short- and long-term temporal changes in
available for nomenclatural purposes? Bulletin of Zoological Nomen- the assemblage structure of Amazonian dung beetles. Oecologia
clature 72 (1): 19-32. https://doi.org/10.21805/bzn.v72i1.a14 195: 719-736. https://doi.org/10.1007/s00442-020-04831-5
Kury A. B., Mendes A. C., Cardoso L., Kury M. S. & Granado [Oken L.] 1834. — J. Th. Lacordaire, über die Lebensweise der
A. A. 2020. — World Catalogue of Opiliones. WCO-Lite: Online Käfer im mittäglichen America. Viele Käfer namentlich angeführt.

56 ZOOSYSTEMA • 2024 • 46 (2)

 Isis von Oken [volume for the year 1834] (11): 1112-1127. [The C. F. P. von Martius, Munich [‘Monachii’], iii + 44 p. + 224 p.
authorship of this review is not stated expressly anywhere, as far + 40 pls. [publication date for nomenclatural purposes: 31st
as I could find, but the context makes clear that the journal’s edi- December 1830 (cf. Sherborn 1922; Evenhuis 1997b; Bouchard
tor, Lorenz Oken (1779-1851), the famous German naturalist of et al. 2011; Bousquet 2016). The colophon on page 224 is dated
the Naturphilosophie movement, was the one who wrote it and 13th December 1833 (‘Idibus Decembris MDCCCXXXIII’), but
the other literature reviews that appeared in the journal. For this this date must refer only to the printing of the last livraison, not
reason, Oken should be credited with its authorship under Arti- being valid for the publication date of the first livraison where
cle 50.1 of the Code. In Cupello et al. (2022), we conservatively Onitis aeruginosus was given availability and which, consequently,
treated the review as published anonymously, but this seems concerns us here. The editorship of this book is also disputed. The
unjustified. Kertesz (1986) tells the details of the history of Isis several individuals and their cities listed by Evenhuis (1997b) as
von Oken and confirms that Oken was the one who prepared the editors were, in fact, purely booksellers, not editors in the sense
literature reviews] [publication date for nomenclatural purposes: of those who underwrote the costs of publication, as defined by
30th November 1834. Neal Evenhuis (personal communication, Bousquet (2016). The one cited by Bouchard et al. (2011) and
26th January 2023) confirmed that Isis von Oken appeared in Kury et al. (2020), Friedrich Fleischer, as far as I could find, is
monthly issues. Oken’s review of Lacordaire’s paper appeared in mentioned only on the 1834 title page as the bookseller in Leip-
the 11th issue of 1834, therefore, in November of that year, with zig, and is also probably incorrect as the editor. Other important
no more specific date known to me. Following Article 21.3.1, I works citing Perty’s book have simply omitted the editor alto-
take the last day of that month as the publication date]. https:// gether (e.g., Sherborn 1922; Blackwelder 1957; Scherer 1983;
www.biodiversitylibrary.org/page/27510335 Monné 2005; Bousquet 2016). In my opinion, however, there is
d’Olsoufieff G. 1924. — Les Phanaeides (Coleoptera – Lamel- no doubt that Carl Friedrich Philipp von Martius (1794-1868)
licornia), famille Scarabaeidae – tr. Coprini. Insecta 13: 4-201. was the editor of this book. The 1834 title page, besides saying
[publication date for nomenclatural purposes: 31st December that the book was produced at the ‘editor’s expenses’ (‘IMPEN-
1924 (no more precise date than the year was found)] SIS EDITORIS’), bears the information ‘praefatus est et edidit
d’Orbigny A. 1835. — Voyage dans l’Amérique Méridionale (le Brésil, Carol. Frideric. Philip. de Martius’ below the title, making it clear
la République orientale de L’Uruguay, la République Argentine, la the true identity of the person who financed the production of
Patagonie, la République du Chili, la République de Bolivia, la the book, edited it, and was thus ultimately responsible for its
République du Pérou), exécuté pendant les années 1826, 1827, 1828, publication.] https://www.biodiversitylibrary.org/page/57932830
1829, 1830, 1831, 1832 et 1833, par Alcide d’Orbigny. Tome Pfennig K. S. & Pfennig D. W. 2009. — Character displacement:
Premier. Partie Historique du voyage dans l’Amérique Méridionale. ecological and reproductive responses to a common evolutionary
Pitois-Levrault et C.e, Paris, and V.e Lerrault, Strasbourg, 672 p. problem. Quarterly Review of Biology 84 (3): 253-276. https://
[publication date for nomenclatural purposes: 31st December doi.org/10.1086/605079
1835. Taken in part from the title page, which states only the Preudhomme-de-Borre A. 1886. — Liste des Lamellicornes Lapa-
year, 1835; day and moth determined by Article 21.3.2 of the rostictiques recueillis par feu Camille van Volxem pendant son
Code.] https://www.biodiversitylibrary.org/page/49209075 voyage au Brésil et à La Plata en 1872 suivie de la description
d’Orbigny A. 1839-1843. — Voyage dans l’Amérique Méridionale (le de dix huit espèces nouvelles et un genre nouveau. Annales de la
Brésil, la République orientale de L’Uruguay, la République Argen- Société entomologique de Belgique 30: [103]-120. https://www.
tine, la Patagonie, la République du Chili, la République de Bolivia, biodiversitylibrary.org/page/12278849
la République du Pérou), exécuté pendant les années 1826, 1827, Price D. L. & May M. L. 2009. — Behavioral ecology of Pha-
1828, 1829, 1830, 1831, 1832 et 1833, par Alcide d’Orbigny. Tome naeus dung beetles (Coleoptera: Scarabaeidae): review and new
Deuxième. Partie Historique. P. Bertrand, Paris, and V.e Lerrault, observations. Acta Zoológica Mexicana (N. S.) 25 (1): 211-238.
Strasbourg, 667 p. [publication date taken from the title page; https://www.scielo.org.mx/scielo.php?script=sci_arttext&pi
no more precise or reliable dating was found in the specialised d=S0065-17372009000100022
literature] https://www.biodiversitylibrary.org/page/45472755 Ribeiro S., Moreira L. F. B., Overbeck G. E. & Maltchik L.
Pacheco T. L. & Vaz-de-Mello F. Z. 2017. — Rediscovery of 2021. — Protected areas of the Pampa biome presented land
the Neotropical genus Paracryptocanthon Howden & Cook, use incompatible with conservation purposes. Journal of Land
2002 (Coleoptera: Scarabaeidae: Scarabaeinae) in the Brazilian Use Science 16 (3): 260-272. https://doi.org/10.1080/17474
Atlantic Forest. Annales de la Société Entomologique de France 23X.2021.1934134
(N. S.) electronic-only detached version: 1-7 [publication date Roesch L. F. W., Vieira F. C. B., Pereira V. A., Schünemann A. L.,
for nomenclatural purposes: 11th April 2017 (from the jour- Teixeira I. F., Senna A. J. T. & Stefenon V. M. 2009 — The
nal’s website)] [volume-integrated version published in 2017, Brazilian Pampa: A fragile biome. Diversity 1: 182-198. https://
volume 53 (2): 99-105.] https://doi.org/10.1080/00379271. doi.org/10.3390/d1020182
2017.1308809 Sánchez M. V. & Genise J. F. 2008. — Nest and brood cham-
Papavero N. 1971. — Essays on the history of Neotropical dipterology, ber structure of two South American dung beetles: Gromphas
with special reference to collectors (1750-1905). Vol. I. Museu de lacordairei Brullé and Ontherus sulcator (Fabricius) (Coleoptera:
Zoologia, Universidade de São Paulo, São Paulo, Brazil, 216 p. Scarabaeidae: Scarabaeinae). The Coleopterists Bulletin 62 (1):
https://doi.org/10.5962/bhl.title.101715 49-61. https://doi.org/10.1649/986.1
Papavero N. 1973. — Essays on the history of Neotropical dipterology, Scherer G. 1983. — Die von J. B. v. Spix und C. F. Ph. v. Martius
with special reference to collectors (1750-1905). Vol. II. Museu de in Südamerika gesammelten Coleopteren. Spixiana 9: 295-305.
Zoologia, Universidade de São Paulo, São Paulo, i-iii + [217]- Sherborn C. D. 1922. — Index animalium sive index nominum
444 p. https://doi.org/10.5962/bhl.title.101597 quae ab A. D. MDCCLVIII generibus et speciebus animalium
Papavero N. 1994. — Rudimentos de latim, in Papavero N. (ed.). imposita sunt societatibus eruditorum adiuvantibus. Sectio secunda.
Fundamentos práticos de taxonomia zoológica (coleções, bibliografia, A kalendis ianuariis, MDCCCI usque ad finem Decembris, MDC-
nomenclatura). Editora Unesp, São Paulo:121-150. CCL. Part I. Introduction, bibliography and index A-Aff.: 1-128.
Perty J. A. M. 1830. — Insecta Brasiliensia: 1-60, pls. 1-12, in 1801-1850. The Trustees of the British Museum, London, cxxxi
Perty J. A. M. 1830-1833. Delectus animalium articulatorum, + 128 p. https://www.biodiversitylibrary.org/page/36191976
quae in itinere per Brasiliam annis MDCCCXVII-MDCCCXX da Silva P. G. 2011. — Dung beetles (Coleoptera: Scarabaeidae:
jussu et auspiciis Maximiliani Josephi I. Bavariae regis augustissimi Scarabaeinae) of two non-native habitats in Bagé, Rio Grande
peracto collegerunt Dr. J. B. de Spix et Dr. C. F. Ph. de Martius. do Sul, Brazil. Zoological Studies 50 (5): 546-559.

57
 Cupello M.

da Silva P. G. 2017. — Dung beetles (Coleoptera, Scarabaeinae) Spix J. B. von & Martius C. F. P. von. 1823. — Reise in Brasilien
from high-altitude grasslands in São Joaquim National Park, auf Befehl Sr. Majestät Maximilian Joseph I. Königs von Baiern in
Santa Catarina, southern Brazil. Check List 13 (6): 817-830. den Jahren 1817 bis 1820 gemacht und beschrieben. Erster Theil.
https://doi.org/10.15560/13.6.817 Anonymous editor, Munich, x + 412 p. https://www.biodiver-
da Silva P. G., Audino L. D., Nogueira J. M., de Moraes L. P. & sitylibrary.org/page/28880206
Vaz-de-Mello F. Z. 2012b. — Escarabeíneos (Coleoptera: Spix J. B. von & Martius C. F. P. von. 1828. — Reise in Brasilien
Scarabaeidae: Scarabaeinae) de uma área de campo nativo no auf Befehl Sr. Majestät Maximilian Joseph I. Königs von Baiern in
bioma pampa, Rio Grande do Sul, Brasil. Biota Neotropica 12 (3): den Jahren 1817 bis 1820 gemacht und beschrieben. Zweiter Theil.
[1]-8. https://doi.org/10.1590/S1676-06032012000300024 C. F. P. von Martius, Munich, viii + [413]-884. https://www.
da Silva P. G. & Bogoni J. A. 2014. — Dung beetles (Coleoptera: biodiversitylibrary.org/page/28862211
Scarabaeidae: Scarabaeinae) attracted to rotten eggs in the Atlantic Spix J. B. von & Martius C. F. P. von. 1831. — Reise in Brasilien
Forest in subtropical southern Brazil. The Coleopterists Bulletin auf Befehl Sr. Majestät Maximilian Joseph I. Königs von Baiern in
68 (2): 339-342. https://doi.org/10.1649/0010-065X-68.2.339 den Jahren 1817 bis 1820 gemacht und beschrieben. Dritter und
da Silva P. G. & Di Mare R. A. 2012. — Escarabeíneos copro- letzter Theil. C. F. P. von Martius, Munich, viii + [885]-1388 +
necrófagos (Coleoptera, Scarabaeidae, Scarabaeinae) de frag- 40 p. https://www.biodiversitylibrary.org/page/28885488
mentos de Mata Atlântica em Silveira Martins, Rio Grande do Streit B. 2012. Edmonds Phanaeini collection donated to Texas
Sul, Brasil. Iheringia, Série Zoologia 102 (2): 197-205. https:// A&M. Scarabs 70: 14-15.
doi.org/10.1590/S0073-47212012000200012 Sturm J. 1843. — Catalog der Kaefer-Sammlung von Jacob Sturm.
da Silva P. G., Garcia M. A. R. & Vidal M. B. 2008. — Besouros Privately published, Nuremberg [Nürnberg], III-XII + 386 p.
copro-necrófagos (Coleoptera: Scarabaeidae stricto sensu) coletados + pl. I-VI. [publication date for nomenclatural purposes: 27th
em ecótono natural de campo e mata em Bagé, RS. Ciência e Natura, July 1843 (see Bousquet 2016)] https://doi.org/10.5962/bhl.
UFSM 30 (2): 71-91. https://doi.org/10.5902/2179460X9839 title.37837
da Silva P. G., Garcia M. A. R. & Vidal M. B. 2009. — Besouros Tremoleras J. 1910. — Coleopterologische Skizze von Uruguay.
copro-necrófagos (Coleoptera: Scarabaeidae sensu stricto) do municí- Entomologische Blätter 6: 22-28, 39-41. https://www.biodiver-
pio de Bagé, RS (bioma campos sulinos). Biociências 17 (1): 33-43. sitylibrary.org/page/33946292
da Silva P. G., Vaz-de-Mello F. Z. & Di Mare R. A. 2011. — Uchoa M. A. & Rodrigues M. M. 2019. — Dung beetles (Coleop-
Guia de identificação das espécies de Scarabaeinae (Coleoptera: tera: Scarabaeoidea) population patterns in three environments
Scarabaeidae) do município de Santa Maria, Rio Grande do Sul, in the Midwest of Brazil. EntomoBrasilis 12 (1): 19-26. https://
Brasil. Biota Neotropica 11 (4): 329-345. https://doi.org/10.1590/ doi.org/10.12741/ebrasilis.v12i1.825
S1676-06032011000400027 Vaz-de-Mello F. Z. & Cupello M. 2018a. — The type specimens
da Silva P. G., Vaz-de-Mello F. Z. & Di Mare R. A. 2012a. — of South American dung beetles. Part I: On the species described
Attractiveness of different bait to the Scarabaeinae (Coleoptera: in the genus Canthon Hoffmannsegg, 1817 by the German ento-
Scarabaeidae) in forest fragments in extreme southern Brazil. mologist Adolf Schmidt (1856-1923). Spixiana 41 (1): 33-76.
Zoological Studies 51 (4): 429-441. Vaz-de-Mello F. Z. & Cupello M. 2018b. — The type specimens
da Silva P. G., Vaz-de-Mello F. Z. & Di Mare R. C. 2013. — of South American dung beetles, Part II: The species described
Diversity and seasonality of Scarabaeinae (Coleoptera: Scara- by the Austrian coleopterist Ludwig Redtenbacher (1814-1876)
baeidae) in forest fragments in Santa Maria, Rio Grande do Sul, in the 1868 part of the ‘Reise der österreichischen Fregatte Novara
Brazil. Anais da Academia Brasileira de Ciências 85 (2): 679-697. um die Erde’ (Insecta: Coleoptera, Scarabaeidae, Scarabaeinae).
https://doi.org/10.1590/S0001-37652013005000033 Annalen des Naturhistorischen Museums in Wien, B 120: 42-58.
Simões-Clivatti T. R. O. & Hernández M. I. M. 2022. — Ecologi- Vaz-de-Mello F., Larsen T., Silva F., Gill B., Spector S. &
cal indication metrics on dung beetles metacommunities in native Favila M. 2014. — Anisocanthon pygmaeus. The IUCN Red
forests and Pinus monocultures. Frontiers in Ecology and Evolution List of Threatened Species 2014: e.T137106A520010. Available
10:972176: 1-13. https://doi.org/10.3389/fevo.2022.972176 online at: https://doi.org/10.2305/IUCN.UK.2014-1.RLTS.
Smith A. B. T. 2006. — A review of the family-group names for the T137106A520010.en. (last accessed on 28th January 2023).
superfamily Scarabaeoidea (Coleoptera) with corrections to nomen- Waterhouse C. O. 1891. — New Scarabaeidae in the British
clature and a current classification. Coleopterists Society Monograph 5: Museum: a fifth contribution. The Annals and Magazine of Natural
144-204. https://digitalcommons.unl.edu/entomologypapers/122/ History, including Zoology, Botany and Geology, Sixth Series 8 (43):
Smith A. Z. 1986. — A history of the Hope Entomological Collec- 53-61. [publication date for nomenclatural purposes: 1st July
tions in the University Museum, Oxford, with lists of archives and 1891 (Evenhuis 2003)] https://www.biodiversitylibrary.org/
collections. Clarendon Press, Oxford, xiii + 172 p. page/19244729
Smith F. 1869. — [proceedings from July 5, 1869]. Proceedings of Wiktionary 2019a. — punctatus. Available online at: https://
the Entomological Society of London [year 1869]: xvi-xviii. https:// en.wiktionary.org/wiki/punctatus (last accessed on 23rd May
www.biodiversitylibrary.org/page/14787690 2023).
Smith H. M. & Smith R. B. 1972. — Chresonymy ex synonymy. Wiktionary 2019b. — amazonicus. Available online at: https://
Systematic Zoology 21 (4): 445. https://doi.org/10.1093/sys- en.wiktionary.org/wiki/amazonicus (last accessed on 23rd May
bio/21.4.445 2023).
Souza Jr. C. N., Shimbo J. Z., Rosa M. R., Parente L. L., Alen- Wiktionary 2022. — dichrous. Available online at: https://
car A. A., Rudorff B. F. T., Hasenack H., Matsumoto M.,
en.wiktionary.org/wiki/dichrous (last accessed on 23rd May 2023).
Wiktionary 2023a. — δι-. Available online at: https://en.wiktionary.
Ferreira L. G., Souza-Filho P. W. M., de Oliveira S. W.,
org/wiki/%CE%B4%CE%B9-#Ancient_Greek (last accessed
Rocha W. F., Fonseca A. V., Marques C. B., Diniz C. G.,
on 23rd May 2023).
Costa D., Monteiro D., Rosa E. R., Vélez-Martin E.,
Wiktionary 2023b. — -us. Available online at: https://en.wiktionary.
Weber E. J., Lenti F. E. B., Paternost F. F., Pareyn F. G. C.,
org/wiki/-us#Latin (last accessed on 23rd May 2023).
Siqueira J. V., Viera J. L., Neto L. C. F., Saraiva M. M.,
Sales M. H., Salgado M. P. G., Vasconcelos R., Galano S.,
Mesquita V. V. & Azevedo T. 2020. — Reconstructing three
decades of land use and land cover changes in Brazilian biomes Submitted on 11 April 2023;
with landsat archive and earth engine. Remote Sensing 12 (2735): accepted on 26 July 2023;
[1]-27. https://doi.org/10.3390/rs12172735 published on 9 February 2024.

58 ZOOSYSTEMA • 2024 • 46 (2)

 ENDNOTES

1. — Article 16.4.2 provides another, even clearer example in see Cupello & Vaz-de-Mello 2015). Not finding it there, we
which it is the author’s intention, not the actual materilisation deemed the holotype lost. When I was writing this paper,
of this intention, that determines availability. In this Article, the however, I recalled that Federico Ocampo had mentioned to
Code rules that, for a new species-group name published after me in an email exchange in early 2012 that he had planned
1999 to be available, authors must state their intention that during the 2000s to revise Gromphas, but that, eventually,
the name-bearing types, if extant, are or will be deposited in I took over independently the project and he aborted the
a collection. This need not actually happen, however. A name idea. Recalling this, the suspicion grew in me that Ocampo
whose holotype is a living specimen will not become unavail- might have borrowed the holotype from the MNHN during
able if its holotype eventually dies in the wild and fails to end the 2000s and that it could still be with him. I contacted him
up in a collection. Provided that this was not what the authors in June 2021 and he kindly confirmed that this is indeed the
originally intended ‒ i.e., if their original vision was that the case. The holotype, as stated by Blanchard (1846), is a female
living holotype would eventually be caught and deposited in and bears the labels shown in Fig. 10A.
a collection ‒, and that the authors stated in the original de-
scription their original intention, then the name will remain 4. — The presence in Paraguay is still speculative. The only
available regardless of what actually happens to the holotype. reason I mention the country here is Martínez’s (1959) re-
cord of Paraguay along with Uruguay and the Argentinian
2. — I say zoological names overall, and not only new names, provinces of Buenos Aires and Corrientes as composing the
because the presence of names in available works is relevant range of G. dichroa. The basis for this record was, however,
for nomenclatural aspects beyond simply establishment of not explained. Since I found no Paraguayan G. dichroa in
availability, such as defining prevailing usage and reversal of the Martínez collection material at the CMNC or elsewhere,
precedence as governed by Art. 23.9. I suspect that this was either a lapse, a guess from the other
places where the species was known to occur, or in litteris in-
3. — For our revision of Gromphas, Cupello & Vaz-de-Mello formation from Paraguayan collectors. If the record is real, it
(2013, 2015) searched without success for the holotype of most likely refers to the savannas, wetlands, or gallery forests
Gromphas dichroa in the dung beetle collection of the MNHN, of the Humid Chaco of southern Paraguay, given its proxim-
where the d’Orbigny South American material, of which the ity to the areas in southern Brazil and northeastern Argentina
specimen is part, is mostly housed (Horn et al. 1990b; but where the species was confirmedly caught.

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