ArticlePDF Available

Revision of the Neotropical tribe Alvarengiini Frey, 1975 (Coleoptera: Scarabaeidae)

March 2024
Austral Entomology 63(1):12-48

March 2024
63(1):12-48

DOI:10.1111/aen.12676

Authors:

Matheus Bento

Instituto Nacional de Pesquisas da Amazônia

Mary E. Jameson

Wichita State University

Claudio Ruy Vasconcelos Fonseca

Instituto Nacional de Pesquisas da Amazônia

The leaf chafer tribe Alvarengiini Frey, a long-neglected taxon that has been overlooked in the biodiversity literature from its inception, is revised for the first time. As a result of this research, the tribe is composed of two species in two genera distributed from Bahia, Brazil (in the north), to Parana, Brazil (in the south), and Paraguay. Two synonyms are proposed: Alvarengius silphoides Frey, syn. nov. for Ottokelleria dispar (Burmeister) and Alvarengius Frey, syn. nov. for Ottokelleria D’Andretta & Martínez. A new genus is described, Carinochilus, gen. nov., and includes one species: Carinochilus marginatus (Burmeister), comb. nov. The lectotype for O. dispar and a neotype for C. marginatus are designated. We provide an identification key to the tribes of Rutelinae (in English and Portuguese) and incorporate classification changes in the subfamily. A key for the identification of Alvarengiini genera and species is also provided. All taxa are circumscribed and illustrated, and distributional data are synthesised in maps.

Content uploaded by Claudio Ruy Vasconcelos Fonseca

Content may be subject to copyright.

TAXONOMIC ARTICLE

Revision of the Neotropical tribe Alvarengiini Frey, 1975

(Coleoptera: Scarabaeidae)

Matheus Bento

| Mary Liz Jameson

| Paschoal Grossi

Claudio Ruy Vasconcelos da Fonseca

Coordenaç˜

ao de Biodiversidade, Laborat

orio de

Sistem

atica e Ecologia de Coleoptera (LASEC),

Instituto Nacional de Pesquisas da Amazônia

(INPA), Manaus, Brazil

Wichita State University, Wichita, Kansas, USA

Programa de P

os-Graduaç˜

ao em Entomologia,

Universidade Federal Rural de Pernambuco

(UFRPE), Recife, Brazil

Correspondence

Matheus Bento, Coordenaç˜

ao de Biodiversidade,

Laborat

orio de Sistem

atica e Ecologia de

Coleoptera (LASEC), Instituto Nacional de

Pesquisas da Amazônia (INPA), Manaus, Brazil.

Email: mabento2008@gmail.com

Funding information

Conselho Nacional de Desenvolvimento

Científico e Tecnol

ogico, Grant/Award Numbers:

309786/2019-3, 424048/2018-3, 449366/2014-6;

Coordenaç˜

ao de Aperfeiçoamento de Pessoal de

Nível Superior, Grant/Award Numbers:

88881.690179/2022-01, 001

Abstract

The leaf chafer tribe Alvarengiini Frey, a long-neglected taxon that has been over-

looked in the biodiversity literature from its inception, is revised for the first time.

As a result of this research, the tribe is composed of two species in two genera dis-

tributed from Bahia, Brazil (in the north), to Paran

a, Brazil (in the south), and

Paraguay. Two synonyms are proposed: Alvarengius silphoides Frey, syn. nov. for

Ottokelleria dispar (Burmeister) and Alvarengius Frey, syn. nov. for Ottokelleria

D’Andretta & Martínez. A new genus is described, Carinochilus,gen. nov., and

includes one species: Carinochilus marginatus (Burmeister), comb. nov. The lecto-

type for O. dispar and a neotype for C. marginatus are designated. We provide an

identification key to the tribes of Rutelinae (in English and Portuguese) and incor-

porate classification changes in the subfamily. A key for the identification of Alvar-

engiini genera and species is also provided. All taxa are circumscribed and

illustrated, and distributional data are synthesised in maps.

KEYWORDS

chafers, identification, morphology, Scarabaeoidea, taxonomy

INTRODUCTION

The tribal and subtribal classification of ruteline scarabs

(Scarabaeidae: Rutelinae) has accumulated problematic

issues over time (Jameson 1998; Smith et al. 2006) includ-

ing classification and relationships of the poorly known

Neotropical tribe Alvarengiini Frey, 1975, whose species

have been overlooked since their discovery due to a con-

fusing classification and nomenclatural history. Over

190 years of nomenclatural complications (homonymy,

synonymy and transfers between higher-level categories)

have hindered the rediscovery of these species, obscured

the taxonomy and relationships of the Rutelinae and

blurred our understanding of Neotropical biodiversity.

Alvarengiini is a small tribe of uncommon and sexually

dimorphic ruteline scarabs that habit the Atlantic rainfor-

est from Brazil to eastern Paraguay. Specimens of this

tribe are infrequently found in collections and rarely iden-

tified, partly because of nomenclatural and taxonomic

confusion, as well as their similarity with closely related

subfamilies. Adults are distinctive among ruteline scarabs,

with relatively small to medium size (10–20 mm in

length), body light to dark brown, rounded oval shape, a

large clypeal expansion, and males with finely to coarsely

crenulate margins and carinae. Because of their overall

gestalt and tarsal morphology, they can be confused with

members of the Dynastinae or Melolonthinae. Very little

is known of the natural history of the group, and the

immature stages for all the species remain unknown.

Taxa in this tribe have had complicated taxonomic

and nomenclatural histories, having been transferred

Matheus Bento: https://zoobank.org/urn:lsid:zoobank.org:author:185B257D-F0E6-

4921-BD3E-4346031B7F28

Mary Liz Jameson: https://zoobank.org/urn:lsid:zoobank.org:author:1D7E1826-

B328-4CD3-B8DF-5FADA76EE625

Paschoal Grossi: https://zoobank.org/urn:lsid:zoobank.org:author:5347A9A1-F9DD-

44E8-ABB8-2542E8EDF1FB

Claudio Ruy Vasconcelos da Fonseca: https://zoobank.org/urn:lsid:zoobank.org:

author:6D15DBCA-E796-4954-883E-424A990EF669

Zoobank number for the paper: https://zoobank.org/urn:lsid:zoobank.org:pub:

F39EBBB5-0E38-4144-B8CF-704082313428

Version of record published on 14 March 2024

Received: 31 July 2023 Revised: 21 November 2023 Accepted: 29 November 2023

DOI: 10.1111/aen.12676

between the subfamilies Melolonthinae, Dynastinae and

Rutelinae (Burmeister 1847;Frey1975;Lacordaire1855;

Ohaus 1928;Smith&Evans2005). Important works

(e.g., Blackwelder 1944; Blanchard 1850; Machatschke 1965,

1972;Ohaus1934) completely overlooked the names that

are now included in the group. Hence, to our knowledge,

this tribe has not been included in any identification keys,

diagnoses and comparative notes (e.g., Jameson 1990;

Soula 1998). Even after transfer to the Rutelinae (Smith &

Evans 2005) and inclusion in important nomenclatural

works (Bouchard et al. 2011;Krajcik2007;Smith2006), this

group continued to be omitted from discussion of ruteline

classification (e.g., Jameson & Ratcliffe 2011;Soula2011). In

part, these omissions occurred due to a scarcity of informa-

tion, further complicating our knowledge of the group’s

biodiversity and natural history.

Due to the long confusion concerning the tribe and

lack of comprehensive study, we aim to provide a founda-

tion for the Alvarengiini. To address this broad gap in our

understanding of leaf chafer biodiversity, we recover his-

torical information, synthesise the taxonomy and classifi-

cation of the tribe based on morphological traits, provide

reliable means of identification for the genera and species

and integrate distributional data. Taxa at all taxonomic

levels are redescribed, illustrated, circumscribed and com-

pared. Generic and specific synonyms are provided, and a

lectotype for Ottokelleria dispar (Burmeister 1847) and

a neotype for Carinochilus marginatus (Burmeister 1847)

are designated. A new key to tribes of Rutelinae in English

and Portuguese is also provided, in which the tribe Alvar-

engiini is included for the first time. Finally, a distribution

map for both Alvarengiini species is included.

MATERIALS AND METHODS

We examined type and non-type material (161 specimens)

from the following institutions (curators’names

parenthesised):

CEIOC—Coleç˜

ao Entomol

ogica do Instituto Oswaldo

Cruz, Rio de Janeiro, Brazil (M

arcio Felix and Claudia

Leal Rodrigues).

CEMT—Seç˜

ao de Entomologia da Coleç˜

ao Zool

ogica da

Universidade Federal do Mato Grosso, Cuiab

a, Brazil

(Fernando Zagury Vaz-de-Melo).

CERPE—Coleç˜

ao Entomol

ogica da Universidade

Federal Rural de Pernambuco, Recife, Brazil (Paschoal

Coelho Grossi and Daniele Regina Parizotto).

EPGC—Everardo and Paschoal Grossi Collection, Nova

Friburgo, Brazil (Everardo Grossi).

CMN—Canadian Museum of Nature, Ottawa, Canada

(François Génier).

DZUP—Coleç˜

ao Entomol

ogica Pe. Jesus Santiago Moure,

Universidade Federal do Paran

a, Curitiba, Brazil (Lúcia

Massutti de Almeida).

ISNB—Institut Royal des Sciences Naturelles de Belgique,

Brussels, Belgium (Wouter Dekoninck).

MACN—Museo Argentino de Ciencias Naturales

‘Bernardino Rivadavia’, Buenos Aires, Argentina

(Arturo Roig Alcina).

MLUH—Martin-Luther-Universität,ZentralmagazinNaturwis-

senschaftlicher Sammlungen, Zoologische Sammlung,

Halle, Germany (Hendrik Müller and Karla Schneider).

MNRJ—Museu Nacional do Rio de Janeiro, Universidade

Federal do Rio de Janeiro, Rio de Janeiro, Brazil

(Marcela Monné).

MZUSP—Museu de Zoologia da Universidade de S˜

Paulo, S˜

ao Paulo, Brazil (Sonia Casari and Carlos

Campaner).

NHM—Natural History Museum, London,

United Kingdom (Max Barclay).

NMB—Naturhistorisches Museum Basel, Basel,

Switzerland (Eva Sprecher and Isabelle Zürcher).

USNM—National Museum of Natural History,

Washington, DC, USA (Brett C. Ratcliffe).

ZMHB—Museum für Naturkunde der Humboldt-

Universität, Berlin, Germany (Johannes Frisch and

Bernard Jäger).

Terminology

Morphological terms follow Lawrence and Britton (1994)

for general terminology including female and male geni-

talia, with the adoption of the term ‘tectum’for the distal

portion of the male phallobasis (non-apodeme sensu

Krell 1996), and Kukalov

a-Peck and Lawrence (1993) and

Browne and Scholtz (1994) for hindwings.

Herein, the modifying term ‘sub-’was used as a prefix

in Harris’s(

1979) sense to mean ‘slightly less than’to a

given structure. The term ‘glabrous’is assumed in a strict

sense from Harris (1979) and Torre-Bueno (1985), mean-

ing only a pubescence-free surface, lacking setae. The

term ‘inerm’is used for the cuticular surfaces without any

acute process or projection (Harris 1979).

Definition of characters and measurement

standards

The body length was measured from the apex of the clyp-

eus to the apex of the elytra, and the body width was

measured between humeri. Burmeister’s(

1847) lines were

converted to millimetres using the French and German

systems (C. Zorn, personal communication, 24 July 2023;

http://preussische-masse.de/alte_masse/alte_masse_

preussisch.html). The interocular width was measured in

transverse eye diameters, expressed as the ratio of dis-

tance between eyes by the width of an eye in dorsal view.

Measurements relative to the crenulation of margins and

carinae were modified from Bento and Grossi (2019)asfollows:

•Crenulation size: Crenulations were considered thick

when larger than 0.02 mm in diameter (usually associated

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 13

20521758, 2024, 1, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/aen.12676 by INPA - Instituto Nacional de Pesquisas da Amazonia, Wiley Online Library on [20/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License

with the presence of setae) and fine when very small

(viewed under higher magnification), equal to or less than

0.02 mm in diameter (lacking associated setae).

Considering that finely crenulate surfaces of males are

always dense, the following crenulation density patterns

were defined referring only to the thick crenulations

(>0.02 mm in diameter):

•Densely crenulate margin: crenulations distributed

throughout the margin, distance between crenulations

equal or less than 0.3 mm and setae longer than the

distance between crenulations.

•Moderately crenulate margin: crenulations distributed

throughout the margin, distance between crenulations

greater than 0.3 mm and setae shorter than the dis-

tance between crenulations.

•Sparsely crenulate margin: margin mostly smooth, with

a few crenulations.

•Smooth margin: lacking crenulations.

For punctures and setae, some characters convention-

ally used for Rutelinae (e.g., Jameson & Ratcliffe 2011;

Jameson & Smith 2002) were redefined as follows:

•Puncture density: Surfaces were considered densely

punctate when the punctures were almost confluent or

separated by less than 2 puncture diameters, moder-

ately punctate when the punctures were 2–6 puncture

diameters apart or sparsely punctate when the punc-

tures were separated by more than 6 puncture

diameters.

•Puncture size: Punctures were considered large if equal

or greater than 0.17 mm in diameter, moderately large

if 0.08–0.169 mm, moderate if 0.03–0.079 mm and

small when less than 0.03 mm.

•Setae density: Surfaces were considered densely setose

when with many setae covering completely the surface,

moderately setose when with many setae, but with the

surface visible, and sparsely setose when with only a

few setae.

•Setae length: defined as minute setae when less than

0.05 mm in length, short if between 0.05 and 0.19 mm,

moderately long if 0.2–0.5 mm or long if greater than

0.5 mm.

For taxonomic purposes, male genitalia are diagnostic at

the generic level. However, because of lack of intraspecific

constancy (i.e., variation) in paramera form, male genitalia

are not reliable for species circumscription and diagnosis.

Features of the labium are important for identification.

The labium is subdivided in three regions: prementum

(apex), mentum (middle), and submentum or gulamen-

tum (base). ‘Labium’is used when more than one region

is considered. For instance, the shape of the labium in

Alverengiini is subtrapezoidal (labium here refers to the

prementum plus the mentum).

Characters of the protarsomeres, mesotarsomeres and

metatarsomeres are sexually dimorphic and essential for

the identification of species and genera. We use Arabic

numerals to refer to specific tarsomeres, with ‘1’referring

to the basal tarsomere and ‘5’referring to the apical

(most distal) tarsomere.

Illustrations

Photographs were taken by a Leica DFC295 camera

coupled to the stereomicroscope Leica M165C with 0.73–

12magnification. The photographs were processed in

the LAS Version 4.2 and Helicon Focus (Helicon Soft) pro-

grams. Light source for illumination was LED, and the illu-

mination system follows Kawada and Buffington (2016).

Locality data

The distribution maps were generated by SimpleMappr

(Shorthouse 2010). The geographical coordinates of the

localities were extracted from Google Maps, n.d, and the

georeferenced points were plotted on the distribution

maps. The verbatim label data from type specimens are

transcribed in quotation marks, with ‘/’separating lines

on the same label and ‘//’separating labels. Label data

from non-type specimens are provided as follows: ‘coun-

try, state or province, locality, date, collector or old collec-

tion (quantity, sex symbol and collection acronym)’.

Specimens with no or minimal data were added at the

end of the examined material section, and existent data

were given.

Rules of zoological nomenclature

As part of this research, stabilisation of the taxonomy and

classification required designation of a lectotype and neo-

type. Nomenclatural complications including synonymy,

homonymy and transfers between higher taxa (genera,

tribes and subfamilies) have confused the species in the

group for over 190 years. We follow the International

Code of Zoological Nomenclature (ICZN 1999) and pro-

vide express statements of purpose when needed, speci-

men accessibility and specimen deposition. These

specimens are the name-bearing type specimens. We pro-

vide express statements regarding the need for these

specimens (ICZN 1999, Articles 74 and 75.3), as well as

information regarding accessibility in public institutions

(ICZN 1999, Article 75.3.7).

Species concept

We adopted the Evolutionary Species Concept (ESC)

developed by Simpson (1961) and more recently defined

14 BENTO ET AL.

by Wiley and Mayden (2000). Based on this view, an evo-

lutionary species is ‘an entity comprised of organisms

that maintains its identity from other such entities

through time and over space and which has its own inde-

pendent evolutionary fate and historical tendencies’.We

therefore agree with the arguments provided by Wiley

and Lieberman (2011) and tend to prefer a process-based

species concept rather than an ‘operational’one. Accord-

ing to Wiley and Lieberman (2011), the ESC is a general

concept that provides ontological basis for the applica-

tion of other ‘operational’concepts like the Morphologi-

cal Species Concept, which is used in this work as one

criterion for discovery and delimitation of species.

TAXONOMY

Taxonomic history of Alvarengiini

Ohaus (1918) classified the Rutelinae into two informal

groups based on the position of the labrum with respect to

the clypeus, namely, ‘homalochilidae’with the labrum par-

allel to the clypeus (Figure 1a)and‘orthochilidae’with the

labrum vertically positioned with respect to the clypeal

apex (Figure 1b). In ‘homalochilidae’, the representative

tribes included Rutelini MacLeay, 1819 and Anomalini

Streubel, 1839, while the ‘orthochilidae’included the tribes

Anoplognathini MacLeay, 1819, Geniatini Burmeister, 1844,

Adoretini Burmeister, 1844, and Anatistini Lacordaire, 1855

(previously Spodochlamyini Ohaus, 1918 [see Jameson &

Ratcliffe 2011;Smith2006]). Years later, Frey (1975)pro-

posed the melolonthine tribe Alvarengiini based on the

monotypic genus and species Alvarengius silphoides

Frey, 1975. The tribe and subordinate taxa took a circuitous

and complicated path, and after having been classified in

related subfamilies, they are currently considered members

of the orthochilous group of the subfamily Rutelinae

(Burmeister 1847;Frey1975; Smith & Evans 2005).

The historical background of some Alvarengiini taxa

preceded the formal naming of the tribe (Frey 1975).

Dejean (1833,1837) proposed the genus Pachylus and

placed it among the genera that we refer to today as

members of the scarab subfamily Dynastinae

(Dejean 1833), as well as Melolonthinae (both Scarabaei-

dae) (Dejean 1837). Dejean (1833) did not include any

known species in this genus, whereas in the third cata-

logue, Dejean (1837) listed the genus Pachylus and

included three species from Brazil: Pachylus euryalus,

Pachylus marginatus and Pachylus serratulus, for which he

did not provide a description, thus making the name

Pachylus Dejean a nomen nudum (ICZN 1999, Article 12.1).

Later, Burmeister (1847) described one of the species

listed by Dejean, namely, P. marginatus, and a new spe-

cies, Pachylus dispar, thus validating the genus. Burmeister

(1847) attributed Dejean as the author (although Burmeis-

ter is the appropriate author), and he placed the genus in

the ‘Cyclocephalidae spurii’(Dynastinae), a group that

included aberrant (or ‘spurious’) dynastines such as Hexo-

don Olivier at that time. The rationale for Burmeister to

group these two genera apart from the ‘Oryctomorphi-

dae’and the ‘Cyclocephalidae genuini’was the widened

and dorsoventrally flattened body, the expanded elytral

margins and small eyes.

The unusual characteristics of Pachylus species led to

the creation of higher-level group names. Lacordaire

(1855) contested Burmeister’splacementofPachylus along

with Hexodon and stated that members of the former

genus lack expanded elytral margins. He redescribed

Pachylus and created a subgroup within ‘Cyclocéphalides’

(Dynastinae) to place it, namely, ‘Pachylides’, a family-level

group taxon that could be distinguished from the true

Cyclocephalids in his second subgroup ‘Cycloceph

alides

vrais’based on the toothed tarsal claws. Lacordaire was

somewhat conflicted in placing the genus Pachylus within

the Cyclocephalini. Even though he failed in finding other

characters to distinguish the genus, he stated that it was

‘true that by their facies these insects are very distant from

the Cyclocephalides, among which appears to me to be sit-

uated’(Lacordaire 1855, p. 395; translated from French).

This higher-level group name is not treated as valid and

available (see Smith 2006), but it is an indication of the

unusual character states in the genus Pachylus.Harold

(1869)citedPachylus and placed the genus within Dynasti-

nae and, interestingly, placed it between Hexodon and

Polymoechus LeConte (=Parastasia Westwood; now con-

sidered a member of the Rutelinae). In the same work

(Harold 1869), P. serratulus Dejean (nomen nudum)was

placed as a synonym of P. dispar. To our knowledge, the

name P. euryalus Dejean (nomen nudum) was never

applied to any taxon and is not available.

More than half a century later, Ohaus (1928) trans-

ferred Pachylus from Dynastinae (Cyclocephalini) to Rute-

linae ‘orthochilidae’based on the labrum visible and

vertical, as well as on the unequal tarsal claws. Ohaus

(1928) redescribed the genus including mouthparts and

male genitalia of P. dispar and P. marginatus. After com-

paring species with Platycoelia Dejean (as Callichloris

Curtis), he believed it is necessary to create a distinct sub-

tribe for Pachylus, which, according to Ohaus, should be

classified ‘between Platycoeliina and Brachysternina’

(Anoplognathini). Ohaus, however, neglected to create

such a subtribe, leaving Pachylus with no tribal place-

ment. That may be the reason why Krajcik (2007) consid-

ered the genus Rutelinae incertae sedis.

In addition to the unstable classification of the genus

Pachylus and the tribe Alvarengiini is the problem of hom-

onymy. Apparently unknown to Burmeister (1847), an opi-

lionid (Arachnida) genus named Pachylus was described

by Kollar (1839) (in Koch 1839), thus making the scarab

genus Pachylus Burmeister a junior homonym. More than

100 years passed before the problem of homonymy was

noticed and Ottokelleria D’Andretta & Martínez, 1957

was proposed as a replacement name for Pachylus.In

addition, D’Andretta and Martínez (1957) fixed O. dispar

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 15

FIGURE 1 (a) Head ‘homalochilidae’in lateral view, Macraspis festiva Burmeister; (b) head ‘orthochilidae’in lateral view, Trizogeniates sp.;

(c) Chlorota sp. in caudal view, arrow indicating elytral margin lacking membranous border; (d) Anomala sp. in caudal view, arrow indicating elytral

margin with membranous border; and (e, f) labrum in dorsal view, arrow indicating the medioapical projection. Scales: (a, b) 1 mm, (c, d) 3 mm and (e,

f) 0.5 mm.

16 BENTO ET AL.

(Burmeister) as the type species of the genus. They pro-

vided some distributional data for the species, which

were known to occur in Rio de Janeiro, S˜

ao Paulo, Minas

Gerais and Espírito Santo states in Brazil. Although

D’Andretta and Martínez (1957) agreed with Ohaus’s deci-

sion to place the genus within the subfamily Rutelinae,

they provided no formal tribal placement of Ottokelleria;

thus, species in the genus were largely inaccessible.

Classification of the genus Ottokelleria (=Pachylus)in

both the Rutelinae and Dynastinae meant that information

about species in the genus was not available, and this led

to taxonomic obscurity of the genus for 190 years. Because

of this, when Frey (1975)namedthegenusAlvarengius

Frey, he did not associate his new genus with the pre-

existing ruteline genus Pachylus. In fact, Frey (1975) consid-

ered the new taxon neither ruteline nor dynastine: He

placed the genus in its own tribe (the Alvarengiini) within

‘Melolonthiden’(the historical equivalent for the subfamily

Melolonthinae). He inadequately described A. silphoides

and barely delimited the taxa, providing a combined

description for both genus and tribe, and founded them

on sparsely informative characters. Frey did not discuss the

relationships of the genus to other groups or his rationale

for placing the new tribe within the Melolonthinae. He

compared Alvarengius only with the tribe Geniatini

(Rutelinae), ‘The labrum is fused, similar to the Geniatini

(Ruteliden)’(translated from German). Finally, Smith and

Evans (2005), in their checklist of New World Melolonthi-

nae (Scarabaeidae), transferred A. silphoides (as a member

of the Alvarengiini) to Rutelinae based on a specimen

examined by the latter author. They provided no further

justification (e.g., characters or comparisons). This transfer,

coupled with descriptive insufficiency in Frey (1975), has

contributed to a serious lack of taxonomic foundation,

diagnosis and circumscription of the tribe Alvarengiini.

Smith and Evans (2005) explicitly transferred

A. silphoides Frey and the Alvarengiini from the Melo-

lonthinae to the Rutelinae, but they did not discuss

Ottokelleria. Instead, formal placement of the genus Otto-

kelleria remained neglected until Bouchard et al. (2011),

who were the first to consider this genus as a member of

the tribe Alvarengiini and addressed nomenclatural prob-

lems concerning the family-group name ’Pachylides’

(Lacordaire, 1855) (Bouchard et al. 2011; Smith 2006),

which is unavailable due to the homonymy (ICZN 1999,

Article 11.7.2).

Thus, for the taxa in the Alvarengiini, there is a dearth of

information and a wholesale lack of a foundation with

which to address the biodiversity of this important and

unusual tribe. In this work, we clarify the tribe and the sub-

ordinate taxa and provide comparative characters that form

the basis for understanding subfamilial relationships

(Rutelinae, Dynastinae or Melolothinae) and classification of

the genera and species in the tribe, place Ottokelleria within

the nomenclatural framework and propose a new taxon.

Key to tribes of Rutelinae (Scarabaeoidea,

Scarabaeidae) (after Jameson 1990,2000a;

Janssens 1949; Ohaus 1934)

1—Labrum horizontally positioned with respect to the

longitudinal axis of clypeus, with the base in a poste-

rior plane to apex, forming a distinct and deep cly-

peolabral suture (Figure 1a) ................................................ 2

10—Labrum vertically positioned with respect to the lon-

gitudinal axis of clypeus, with the base in the same

plane as apex and somewhat fused to the clypeal

apex, forming a shallow clypeolabral suture

(Figure 1b) .................................................................................. 3

2—Lateral margin of elytra without membranous border

(Figure 1c); antenna with 10 antennomeres (excep-

tion: Eremophygus Ohaus; Lasiocala Blanchard; Micro-

ogenius Gutiérrez; Parachrysina Bates; Peruquime

Mondaca & Valencia; Pseudogeniates richterianus

Ohaus); protibia usually tridentate, with an inner api-

cal spur ........................................................................... Rutelini

20—Lateral margin of elytra with membranous border

(Figure 1d); antenna with nine antennomeres; proti-

bia usually bidentate, with an inner subapical spur

...................................................................................... Anomalini

3—Labrum and prementum both simple, without med-

ioapical projection ................................................................... 4

30—Labrum with medioapical projection (Figure 1e,f);

prementum with or without medioapical projection

(Figure 3a) ................................................................................... 5

4—Labium subtrapezoidal (Figure 6a,b); mentum fused

at base to gulamentum (Figure 4a); ventral expan-

sion of the clypeus (longitudinally, at middle) equal

to or longer than twice the labrum length

(Figures 2a and 4d,e); antenna with nine antenno-

meres; inner protibial spur present (Figure 6e,f)

................................................................................... Alvarengiini

40—Labium subquadrate (Figure 2d); mentum and gula-

mentum separated by a suture (Figure 2d); ventral

expansion of the clypeus (longitudinally, at middle)

shorter than labrum (exception: male of Spodochla-

mys caesarea) (Figure 2b); antenna with 10 antenno-

meres; inner protibial spur absent (Figure 2c)

....................................................................................... Anatistini

5—Labrum with medioapical projection superimposing

the apex of prementum; prementum without med-

ioapical projection ................................................. Adoretini

50—Labrum and prementum both with medioapical

projection (Figure 3a) ............................................................ 6

6—Protarsomeres cylindrical, not densely setose ven-

trally (Figure 3d) ....................................... Anoplognathini

60—Protarsomeres dorsoventrally flattened and densely

setose ventrally in males and females or only in

males (exception: Geniatosoma Costa Lima and

some members of Geniates Kirby) (Figure 3b,c)

........................................................................................ Geniatini

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 17

Chave para as tribos de Rutelinae

(Scarabaeoidea, Scarabaeidae) (Depois de

Jameson 1990,2000a; Janssens 1949;

Ohaus 1934)

1—Labro horizontalmente posicionado com relaç˜

ao ao

eixo longitudinal do clípeo, com a base num plano

posterior ao 

apice, formando uma sutura clipeolabral

distinta e profunda (Figure 1a) .......................................... 2

10—Labro verticalmente posicionado com relaç ˜

ao ao

eixo longitudinal do clípeo, com a base no mesmo

plano do 

apice e mais ou menos fundida ao 

apice

clypeal, formando uma sutura clipeolabral rasa

.......................................................................................................... 3

2—Margem lateral dos élitros sem borda membranosa

(Figure 1c); antena com 10 antenômeros (exceç˜

ao:

Eremophygus Ohaus; Lasiocala Blanchard; Microogen-

ius Gutiérrez; Parachrysina Bates; Peruquime

FIGURE 2 (a) Head in frontal view, Ottokelleria dispar (Burmeister) (male; mouthparts removed). Spodochlamys iheringi Ohaus: (b) head in lateral

view; (c) left anterior leg in ventral view, arrow indicating the lacking tibial spur; and (d) labium and maxillae. Scales: (a) 0.5 mm and (b–d) 1 mm.

18 BENTO ET AL.

Mondaca & Valencia; Pseudogeniates richterianus

Ohaus); protíbia geralmente tridentada, com um

espor˜

ao apical interno ............................................ Rutelini

20—Margem lateral dos élitros com borda membranosa

(Figure 1d); antena com 9 antenômeros; protíbia

geralmente bidentada, com um espor˜

ao subapical

interno ..................................................................... Anomalini

3—Labro e premento ambos simples, sem projeç˜

medioapical (Figure 2b) ........................................................ 4

30—Labro com projeç˜

ao medioapical (Figure 1e,f); pre-

mento com ou sem projeç˜

ao medioapical

(Figure 3a) ................................................................................... 5

4—L

abio subtrapezoidal (Figure 6a,b); mento basalmente

fundidoaogulamento(Figure4a); expans ˜

ao ventral

do clípeo medialmente igual ou mais comprida que

duas vezes o comprimento do labro (Figures 2a and

4d,e); antena com 9 antenômeros; espor˜

ao protibial

interno presente (Figure 6e,f) .......................... Alvarengiini

40—L

abio subquadrado (Figure 2d); mento e gulamento

separados por uma sutura (Figure 2d); expans˜

ventral do clípeo medialmente mais curta que o

labro (exceç˜

ao: macho de Spodochlamys caesarea)

(Figure 2b); antena com 10 antenômeros; espor˜

protibial interno ausente (Figure 2c) .......... Anatistini

5—Labro com projeç˜

ao medioapical sobrepondo o



apice do premento; premento sem projeç˜

ao medioa-

pical .............................................................................. Adoretini

50—Labro e premento ambos com projeç˜

ao medioapical

(Figure 3a) ................................................................................... 6

6—Protarsômeros cilíndricos, n˜

ao densamente cerdosos

ventralmente (Figure 3d) ..................... Anoplognathini

60—Protarsômeros dorsoventralmente achatados e

densamente cerdosos ventralmente em machos e

fêmeas ou somente em machos (exceç˜

ao: Geniato-

soma Costa Lima e alguns membros de Geniates

Kirby) (Figure 3b,c) ............................................... Geniatini

FIGURE 3 Lobogeniates sp.: (a) head in ventral view showing labrum and labium; (b) protarsus in dorsal view; (c) protarsus in lateral view; and

(d) Hylamorpha elegans Burmeister, protarsus in lateral view. Scale: 1 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 19

Tribe Alvarengiini Frey, 1975

‘Pachylides’Lacordaire, 1855, 394 (unavailable; perma-

nently invalid).

Alvarengiini Frey, 1975, 84 (original description; Melo-

lonthinae); Smith & Evans, 2005, 54 (transfer to Rutelinae);

Smith, 2006, 173 (catalogue); Krajcik, 2007, 20 (checklist);

Bouchard et al., 2011, 256 (catalogue); Ratcliffe et al., 2015,192.

Type genus: Alvarengius Frey, 1975, 85.

Diagnosis

The following character combination distinguishes adults

of Alvarengiini from other Rutelinae. We consider tribal

characters to be consistent with orthochilous Rutelinae:

1. clypeal ventral expansion well developed and medially

equal to or longer than twice the labrum length

(Figures 2a and 4a);

2. labrum vertically positioned with respect to dorsal face

of clypeus and with apical margin rounded or slightly

angled medially, without a medioapical projection

(Figures 2a and 4d,e);

3. labium subtrapezoidal and strongly convex

(Figures 4d,e and 6a,b), with mentum basally fused to

gulamentum (without a suture) (Figure 4a);

4. maxilla with stipital lobes fused; galea cuneiform and

without conspicuous teeth (Figure 5a);

5. mandible reduced and inerm, hidden by clypeus in

dorsal view; molar lobe indistinct, unsulcate and

untoothed (Figure 5b,c) (vs. molar lobe strongly devel-

oped, sulcate and toothed in the tribe Anatistini [see

Jameson & Ratcliffe 2011, fig. 29]);

6. Ventrites 1–5 with a wide laterolongitudinal carina

(Figure 5d), almost as wide as elytral border; and

7. New World distribution.

Description

Male

Clypeal ventral expansion obliquely directed with respect

to the longitudinal axis of frons (Figure 4d). Antennal club

oblong and longer than antennomeres 2–6 combined

(Figure 4b). Lateral margins of elytra moderately to densely

crenulate (Figure 6g,h). Trochanter, femur and tibia with

margins finely crenulate. Inner metatibial spur apically

acute. Protarsus, mesotarsus and metatarsus with a finely

crenulate ventrolongitudinal keel. Protarsomere 5 enlarged

and oval, with a ventromedial projection laterally or dorso-

ventrally flattened (Figure 6e,f) and an internoapical pro-

jection digitiform and with an apical seta (Figure 5g);

anterior protarsal claw enlarged and strongly curved with

apex rounded and unequally bifid (Figures 5e,g and 6e).

Female

Clypeal ventral expansion directed perpendicularly with

respect to the longitudinal axis of frons (Figure 4e).

Antennal club subglobose and shorter than antenno-

meres 2–6 combined (Figure 4c). Margins of trochanter,

femur and tibia without fine crenulation. Inner metatibial

spur apically rounded. Protarsus, mesotarsus and metatar-

sus without ventrolongitudinal keel. Protarsomere 5 sim-

ple, with anterior claw toothed near base (Figure 5f).

Remarks

Nomenclature

The type genus of the tribe Alvarengiini Frey, 1975 is

Alvarengius Frey, 1975. In this work, we consider the

genus Alvarengius to be a junior synonym of the genus

Ottokelleria D’Andretta & Martínez, 1957. Although the

type genus of the tribe is synonymised, the family-group

name is conserved (ICZN 1999, Article 40.1).

Sister group relationships

The phylogenetic relationship of Alvarengiini with the other

tribes within Rutelinae has not been examined. Based on

mouthpart features, the tribe shares characters with the

orthochilus Rutelinae (Anatistini, Anoplognathini, Geniatini

and Adoretini), and it appears most closely allied with the

tribe Anatistini. Based on molecular phylogenetic analysis

(McKenna et al. 2015), the Anatistini were recovered in a clade

that included the Anoplognathini and Rutelini (non-mono-

phyletic) (McKenna et al. 2015;seeMooreetal.2018,fig.2).

Tribal-level relationships of the Rutelinae should be

approached with caution. Phylogenetic analyses suggest

that Rutelinae is a paraphyletic grade of tribes (Ahrens

et al. 2011; Ahrens et al. 2014; Gunter et al. 2016). Some

analyses provide support for Dynastinae +Rutelinae with

some ruteline tribes included in the Dynastinae (Jameson

1998; McKenna et al. 2015); other analyses provide sup-

port for a monophyletic Dynastinae (with or without the

inclusion of exemplar ruteline tribes) (Ahrens et al. 2011,

2014; Gunter et al. 2016). In part, this debate stems from a

lack of a basic foundation of biodiversity science and a

lack of adequate sampling of unusual dynastine and rute-

line taxa that have long posed classification problems

including the ‘Cyclocephalidae spurii’of Burmeister

(1847), Bates’s(1888)‘unindicated group of genera’that

was considered intermediate between Dynastinae and

Rutelinae, and the ‘Rutelisca lineage’of Jameson (1998,

2000b). As mentioned by Jameson (2000b), ‘inclusion of

additional taxa such as Oryctomorphus Guérin-Méneville

(Dynastinae: Pentodontini),

Alvarengius Frey (Melolonthi-

nae), Ottokelleria D’Andretta & Martínez (Rutelinae: incer-

tae sedis), Desmonyx Arrow (Rutelinae: Rutelini),

Mesystoechus Waterhouse (Rutelinae: Anoplognathini)

and Pseudogeniates Ohaus (Rutelinae: Rutelini) may help

to resolve the problem of relationships’(Jameson 2000b).

20 BENTO ET AL.

Phylogenetic analysis of the Alvarengiini requires broad

sampling of Rutelinae, Dynastinae and Melolonthinae in

order to reveal broader biological patterns.

Classification

Ohaus (1928) compared Ottokelleria (as Pachylus) to the

genera Hexodon and Oryctomorphus (both at the time in

Dynastinae). He used the form of mouthparts and tarsal

claws to support the transfer of this genus from Dynasti-

nae (where it was classified by Burmeister [1847]) to Rute-

linae. In order to classify the genus within the subfamily,

Ohaus (1928) first compared it to the genera Desmonyx

Arrow, Rutelisca Bates and Metapachylus Bates (Rutelini)

and noted, ‘It can be clearly seen that the upper lip

[labrum] is vertical, firmly attached with its upper end to

the lower frontal edge of clypeus’(translated from

German). This character state distinguishes the genus as a

member of the large section ‘orthochilidae’. Without jus-

tification, however, Ohaus (1928) eliminated any possibil-

ity that Ottokelleria may be a member of the tribes

Anatistini (as Spodochlamyini), Adoretini or Geniatini.

Then, based particularly on morphology of the legs, he

FIGURE 4 (a) Head in ventral view, Ottokelleria dispar (male). Antenna of O. dispar, (b) male and (c) female; (d) head in lateral view, Carinochilus

marginatus (Burmeister) comb. nov. (male); and (e) head in lateral view, O. dispar (female, right maxilla removed). Scale: 0.5 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 21

FIGURE 5 (a) Right maxilla, Carinochilus marginatus (Burmeister) comb. nov. (male); (b) left mandible in dorsal view, Ottokelleria dispar (male);

(c) left mandible in lateral view, C. marginatus (male); (d) abdomen of C. marginatus (male) in anterolateral view, arrows indicating laterolongitudinal

carinae of the ventrites; protarsus of O. dispar, (e) male and (f) female; and (g) protarsomere 5 of O. dispar (male), arrow indicating the inner apical

projection. Scales: (a–c) 0.25 mm, (d) 1 mm, (e, f) 0.5 mm and (g) 0.2 mm.

22 BENTO ET AL.

FIGURE 6 Comparison of the Alvarengiini genera. Carinochilus gen. nov.: (a) labium, arrows indicating the laterolongitudinal carinae; (c) hindwing;

(e) left anterior leg in ventral view, arrows indicating two ventromedian, longitudinal carinae on tibia; and (g) left elytron in dorsal view, arrow

indicating the moderately crenulate lateral margin. Ottokelleria: (b) labium; (d) hindwing; (f) left anterior leg in ventral view, arrow indicating a

ventromedian, longitudinal carinae on tibia; and (h) left elytron in dorsal view, arrow indicating the densely crenulate lateral margin. Scales: (a,b)

0.25 mm, (c, d) 1 mm, (e, f) 0.5 mm and (g, h) 2 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 23

discussed the similarity of Ottokelleria to the Neotropical

genus Platycoelia Dejean (as Callichloris; Anoplognathini)

and stated: ‘There are so many peculiarities that we must

place the genus [Ottokelleria,asPachylus] within a special

subtribe which has its position between Platycoeliina and

Brachysternina’(both Anoplognathini) (translated from

German).

The tribe Anoplognathini is traditionally defined by

presence of a medioapical projection on both labrum and

prementum (characters shared with Geniatini) and protar-

someres cylindrical rather than dorsoventrally flattened

and not densely setose ventrally as opposed to Geniatini,

which possess protarsomeres densely setose ventrally

and dorsoventrally flattened in only males or both sexes

(Jameson & Hawkins 2005; Lacordaire 1855). Ohaus’s

(1928) decision to place Ottokelleria (as Pachylus) in the

tribe Anoplognathini was misleading, especially when

diagnostic characters are contrasted: Anoplognathini pos-

sesses a medioapical projection on both labrum and pre-

mentum, whereas Alvarengiini lacks the projection on

both labrum and prementum. One of the defining charac-

ter states of Alvarengiini is precisely the absence of ven-

troapical projection on the labrum and prementum

(characters shared with Anatistini). The apical margin of

labrum is rounded or slightly acute medially, thus a

reduction compared to other ‘orthochilidae’possibly due

to a ‘change in lifestyle’according to Machatschke

(1965). In our view, this is not homologous with the pro-

jection in the Adoretini, Anoplognathini and Geniatini

(Figures 1e,f and 3a). Furthermore, the apical margin of

prementum is subtruncated or slightly rounded, without

median projection, sinuosity or notch. These characters

distinguish Alvarengiini from the aforementioned ortho-

chilous tribes (Anoplognathini, Adoretini and Geniatini)

and indicate affinities with the Anatistini.

Jameson and Ratcliffe (2011) hypothesised that the

Anatistini is probably monophyletic. However, the tribal

character once thought to be unique for the Anatistini

among the ‘orthochilidae’is similar in Alvarengiini. Both

tribes possess labrum and prementum with no medioapi-

cal projection, but the Alvarengiini can be easily distin-

guished from the Anatistini by the following characters

(Anatistini characters in parentheses): (1) relatively small

size (9.6–17.3 mm from apex of clypeus to the elytral apex)

(relatively large size [20–46 mm from apex of clypeus to

the apex of pygidium]; Jameson & Ratcliffe 2011); (2) cly-

peal ventral expansion medially equals or longer than

twice the labrum length (clypeal ventral expansion much

shorter than labrum except in male Spodochlamys caesarea

Burmeister, which has the apex of clypeus narrowly raised

to form a small, median projection); (3) labium subtrape-

zoidal and strongly convex, with mentum basally fused to

gulamentum; prementum apically subtruncated or slightly

rounded, without any median notch or emargination

(labium subquadrate and discally furrowed, flat or slightly

convex, with mentum and gulamentum separated by a

conspicuous suture; prementum apically truncated and

usually possessing a median notch); (4) galea cuneiform

and without developed teeth (maxillary galea with teeth

strongly developed and directed at approximately a 90

angle with respect to the longitudinal axis of maxilla);

(5) mandibles inerm and hidden by clypeus, with a molar

lobe weakly developed and without ridges, valleys or teeth

(mandibles well developed and frontally visible, with a

strongly developed molar lobe); (6) antennae with 9 anten-

nomeres (10 antennomeres); (7) protibial spur present

(absent); and (8) abdominal ventrites with a laterolongitu-

dinal carina (without laterolongitudinal carina).

Based on this uncommon character combination,

members of the Alvarengiini can be reliably distinguished.

Some characters appear to be unique in comparison to the

other tribes, such as (1) mentum fused to the gulamentum;

(2) maxillae with stipital lobes fused to each other; (3) male

protarsus, mesotarsus and metatarsus with a finely crenu-

late ventrolongitudinal keel; and (4) protarsomere 5 with

an internoapical projection slender and digitiform. The

laterolongitudinal carina of abdominal sternum was

observed in some species of Bolax Fischer von Waldheim,

1829, Geniates Kirby, 1819, Leucothyreus MacLeay, 1819,

Lobogeniates Ohaus, 1917 (all Geniatini) and Anomala dor-

salis (Fabricius, 1775) (Anomalini, cf. Ratcliffe et al. 2018,fig.

13). However, in comparison with members of the

Alvarengiini, the carina is narrower or completely disap-

pears on terminal ventrites in these genera.

Pachylus Burmeister was originally described within

‘Cyclocephalidae spurii’(historical equivalent of Cycloce-

phalini [Dynastinae]) and subsequently considered within

Cyclocephalini by Lacordaire (1855) and Harold (1869).

Based on the absence of cephalic tubercles (including sex-

ual differences on head and pronotum), presence of fili-

form tarsi and males with enlarged protarsi, Lacordaire

(1855)keptPachylus within ‘Cyclocéphalides’

(=Cyclocephalini), creating the subgroup ‘Pachylides’

(=Alvarengiini). Lacordaire (1855)stated,‘because of its

facies, these insects are very far from cyclocephalides’,yet

‘[in Pachylus] the lateral dilatation of the horizontal fold

that forms the epipleura at the base is neither wider nor

longer than in most of them [dynastine beetles]’.Thenhe

added: ‘…in fact, characters that distinguish them are

reduced to the two above; at least I could not find others’

(translated from French).

Lacordaire (1855), Endrödi (1966) and Moore et al.

(2018) discussed characters that diagnose the Cycloce-

phalini (or the historical equivalent), some of which are

shared with Alvarengiini: body rounded oval; head and

pronotum evenly convex, without horns, keels or depres-

sions; mandibles with outer margin without teeth; and

protarsus and anterior protarsal claw enlarged in males

(with exceptions in a few Cyclocephalini genera). For tax-

onomic purposes, members of the Alvarengiini can easily

be distinguished from the Cyclocephalini by the following

characters (Cyclocephalini characters in parentheses):

(1) ocular canthus cariniform (ocular canthus lacking

carina or ridge); (2) maxilla with stipital lobes fused to

24 BENTO ET AL.

each other (stipital lobes separated by sutures); (3) male

elytra with lateral margins densely to moderately crenu-

late (lateral margins of elytra lacking crenulations);

(4) prosternal process flattened, triangle-shaped, with

apex acute or rounded (prosternal process thickened and

apically truncated); (5) anterior mesotarsal and metatarsal

claw toothed and/or finely crenulate (outer claws simple

and smooth); and (6) ventrites with laterolongitudinal

carina (ventrites lacking laterolongitudinal carina).

Ohaus (1928) stated that ‘because of the corneous

labrum visible under clypeus and tarsal claws indepen-

dently movable’(translated from German), Pachylus

belonged to Rutelinae rather than Dynastinae (Table 1).

Indeed, opposing characteristics traditionally diagnose

both subfamilies (Dynastinae characters in parentheses):

(1) metatarsomeres slightly or strongly ventrally produced

at apex (metatarsomeres truncated, not ventrally pro-

duced) and (2) mesotarsomere and metatarsomere 5 with

a ventroapical split that allows the flexion of outer claw

(apex of mesotarsomere and metatarsomere 5 uncleft).

Based on these characters, members of the Alvarengiini

belong to the Rutelinae. However, exceptions are often

found to some of these characters (e.g., labrum distinct in

Peltonotus [Dynastinae]), thus indicating a poor under-

standing of phylogenetic relationships in the higher

scarabs (including the Rutelinae and Dynastinae).

Finally, Alvarengiini was first described by Frey (1975)

in Melolonthinae based on A. silphoides (=O. dispar),

which was later transferred to Rutelinae by Smith and

Evans (2005). In fact, claws independently moveable,

mandibles externally exposed, tarsomeres short and api-

cally imbricated and the shape of mesotibia and

metatibia not typical for melolonthines strongly suggest

that Alvarengiini does not belong to Melolonthinae

(Table 1).

However, the placement of Alvarengiini is difficult due

to similarities with related subfamilies (Table 1) and highly

modified character systems, which are important and use-

ful at the subfamilial level. For resolution, the enigmatic

position of Alvarengiini must await molecular data and

broad sampling from the Melolonthinae, Dynastinae

and Rutelinae.

Etymology

The name Alvarengiini has the stem based on the type

genus Alvarengius Frey, 1975 (new junior synonym of

Ottokelleria D’Andretta & Martínez, 1957). This name was

in honour of the Brazilian entomologist Moacyr Alvarenga

(1915–2010) who collected the type series.

Biology

As stated by Lacordaire (1855), ‘These insects are not

common in collections’(translated from French), and

Ohaus (1928), ‘Nothing is known about the lifestyle of the

beetle’(translated from German). However, reduced

mouthparts (sensu Nel & Scholtz 1990; Scholtz 2000)may

inform the natural history of members of the Alvarengiini.

The reduction of labrum, mandibles and maxillary galea

along with the complete fusion of the labial and stipital

lobes, absence of ligular lobes, inerm surface of the man-

dibles (including molar lobe not ridged) and the maxillary

galea (at least the absence of developed teeth on both

structures), and the presence of labial palps with two pal-

pomeres in Carinochilus gen. nov. suggest that adults of

Alvarengiini do not feed or feed very little. It is possible

that adults of Alvarengiini have a short life span because

they do not feed, and this may reduce opportunities for

collection and observation.

Collection of some male and female specimens of

Ottokelleria (as P. dispar)‘by electric light’(by Friedrich

Ohaus and Joseph Zik

an) indicates a possible nocturnal

habit in this genus (Ohaus 1928). Based on label data,

flight-intercept traps (FITs) are also efficient for collecting

Alvarengiini individuals. Adult males of O. dispar are good

fliers and were collected during the day at Macaé de Cima

region, Brazil.

Distribution

Alvarengiini is composed of two genera and two species

that are found in the Atlantic Forest in Brazil, from Bahia

state in the northeast to Paran

a state in the south and

east to Paraguay (Figures 7and 8).

Key to genera and species of Alvarengiini

1—Antennal club oblong and longer than antenno-

meres 2–6 combined (Figure 4b). Clypeal ventral

expansion directed obliquely to the longitudinal

axis of frons (Figure 4d). Anterior protarsal claw

enlarged and strongly curved, apically cleft

(Figure 5e,g). All tarsi with a finely crenulate ventro-

longitudinal keel (Figure 5e). Inner metatibial spur

apically acute …Male ....................................................... 2

10—Antennal club subglobose and shorter than anten-

nomeres 2–6 combined (Figure 4c). Clypeal ventral

expansion directed perpendicularly to the longitudi-

nal axis of frons (Figure 4e). Anterior protarsal claw

not enlarged, slightly curved. Tarsi without ventro-

longitudinal keel (Figure 5f). Inner metatibial spur

apically rounded …Female ............................................... 3

2—Labium with finely crenulate laterolongitudinal carinae,

labial palp with two palpomeres (Figure 6a). Ocular can-

thus glabrous (Figure 4d). Protibia with two ventrolon-

gitudinal carinae (Figure 6e). Protibial spur shorter than

protarsomeres 1 and 2 combined (Figure 6e)

............................. Carinochilus marginatus gen. & comb. nov.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 25

TABLE 1 Comparative characters/character states among Alvarengiini and related subfamilies—Melolonthinae characters based on Evans and

Smith (2020).

Characters/taxa Alvarengiini Rutelinae Dynastinae Melolonthinae

Metatarsomeres with

ventro-posterior apices

produced (yes/no)

Yes Yes No No

Apex of metatibia with (1

spur, 2 spurs or 0 spurs)

2 2 2 1, 2 or 0

Protarsal claw of male at

apex (split/entire)

Split Split/entire

Split/entire Split/entire

Protarsal claw

independently moveable

(yes/no)

Yes Yes No/yes No

Mesotarsal and metatarsal

claws independently

moveable (yes/no)

Yes Yes No No

Male with one protarsal claw

(enlarged/not enlarged)

Enlarged Enlarged Enlarged/not enlarged Not enlarged

Ventrites (with/without)

laterolongitudinal carina

With With/without Without With/without

Mesotarsal and metatarsal

claws unequal in length

(yes/no)

Yes Yes

No No

Labrum form ‘orthochilus’

(yes/no)

Yes Yes/no No Yes/no

Mesotarsal and metatarsal

claws deeply cleft

(yes/no)

Yes/no Yes/no

No Yes/no

Abdominal spiracle position

(e.g., I–VI located in

pleural membrane VII

between sternite and

tergite)

e,f

II–IV in pleura; V–VII on

the limit of the

sternites

Vary in position (from

Beutel &

Leschen 2016)

I and II, I–III or I–IV lie in

pleural membrane, III–VI,

IV–VI, or V and VI located

on sternites and VII

usually between tergite

and sternite (from Beutel

& Leschen 2016)

I–IV, I–VorI–VI located in

pleural membrane and VII

on the line of fusion

between sternite and

tergite or on tergite (from

Beutel & Leschen 2016)

Procoxae (transverse/

conical)

Transverse Transverse Transverse Transverse/conical

Empodium of mesotarsus

and metatarsus laterally

flattened (yes/no)

No Yes No No

Hindwing with sclerite 1AX

form

: (1) tail distal arch

strongly reduced and

apex broadly rounded or

(2) head and

anterodorsal margin very

convex and anterodistal

surface noticeably

narrow medially

211 2

Some Anomalini [Rutelinae]. See tab. 2 in Ratcliffe et al. (2018).

Yes (exception: Leptohoplia [Rutelinae] with only one claw or one claw greatly reduced).

No (exception: Hoplia [Melolonthinae] with only one claw).

Yes (‘Lasiocalina’[Rutelinae], but this group is homalochilous).

Lacordaire says, ‘The situation of their last abdominal spiracles proves that it is not of Melolonthids; there cannot be some discussion about putting them among

Cetoniinae; as for Rutelinae and in Dynastinae, they are placed so much on the limit of these two groups, that I do not see other reasons to classify them in the second only

for the brevity (shortness) of their labrum.’

The only character identified by Beutel and Leschen (2016) as separating Melolonthinae from Dynastinae +Rutelinae.

Beutel and Leschen (2016).

26 BENTO ET AL.

FIGURE 7 Distribution map of Carinochilus marginatus (Burmeister) (open star symbol represents state record only).

FIGURE 8 Distribution map of Ottokelleria dispar (Burmeister) (orange symbol represents state record only).

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 27

20—Labium without carinae, labial palp with three pal-

pomeres (Figure 6b). Ocular canthus apically setose

(Figures 2a and 4e). Protibia with a ventrolongitudi-

nal carina (Figure 6f). Protibial spur subequal or lon-

ger than protarsomeres 1 and 2 combined

(Figure 6f) ............................................. Ottokelleria dispar

3—Labium with smooth laterolongitudinal carinae,

labial palp with two palpomeres. Elytral apex entire

(not subtruncate; e.g., Figure 19b,e) and bordered

(Figure 14a,b) .................................................................................

................. Carinochilus marginatus gen. & comb. nov.

30—Labium without carinae, labial palp with three palpo-

meres. Elytral apex subtruncate (Figure 19b,e)and

unbordered (Figure 19e) ....................... Ottokelleria dispar

Chave para gêneros e espécies de

Alvarengiini

1—Clava antennal oblonga e mais comprida que os

antenômeros 2–6 combinados (Figure 4b).

Expans˜

ao ventral do clípeo direcionada obliqua-

mente ao eixo longitudinal da fronte (Figure 4d).

Garra protarsal anterior alargada e fortemente curvada,

com uma fenda apical (Figure 5e,g). Todos os tarsos

com uma quilha ventrolongitudinal finamente crenu-

lada (Figure 5e). Espor˜

ao metatibial interno com 

apice

agudo …Macho ................................................................................... 2

10—Clava antennal subglobosa e mais curta que os

antenômeros 2–6 combinados (Figure 4c). Expans˜

ventral do clípeo direcionada perpendicularmente

ao eixo longitudinal da fronte (Figure 4e). Garra pro-

tarsal anterior n˜

ao alargada, levemente curvada.

Tarsos sem quilha ventrolongitudinal (Figure 5f).

Espor˜

ao metatibial interno com 

apice arredondado

…Fêmea .................................................................................... 2

2—L

abio com carenas laterolongitudinais finamente crenula-

das, palpo labial com 2 palpômeros (Figure 6a). Canthus

ocular glabro (Figure 4d). Protíbia com 2 carenas ventro-

longitudinais (Figure 6e). Espor˜

ao protibial mais curto

que os protarsômeros 1 e 2 combinados (Figure 6e)

...............................Carinochilus marginatus gen. & comb. nov.

20—L

abio sem carenas, palpo labial com 3 palpômeros

(Figure 6b). Canthus ocular apicalmente cerdoso

(Figures 2a and 4e). Protíbia com uma carena ventro-

longitudinal (Figure 6f). Espor˜

ao protibial subigual ou

mais comprido que os protarsômeros 1 e 2 combina-

dos (Figure 6f) .............................................. Ottokelleria dispar

3—L

abio com carenas laterolongitudinais lisas, palpo labial

com 2 palpômeros. ´

Apice elitral inteiro (n ˜

ao subtrun-

cado; e.g., Figure 19b,e)ecombordas(Figure14a,b)

............................. Carinochilus marginatus gen. & comb. nov.

30—L

abio sem carenas, palpo labial com 3 palpômeros.

Apice elitral subtruncado (Figure 19b,e)esembordas

(Figure 19e) .................................................... Ottokelleria dispar

Carinochilus gen. nov.

(Figures 4d,5a,c,d and 6a,c,e,g)

https://zoobank.org/urn:lsid:zoobank.org:act:878CDD01-

1C7B-4018-88FC-F497B0CC1735

Type species: Pachylus marginatus Burmeister, 1847, pre-

sent designation.

Diagnosis

Adult specimens in the genus Carinochilus are easily distin-

guished from Ottokelleria by the following characters: ocular

canthus glabrous (Figure 4d) (setose in Ottokelleria). Labium

with laterolongitudinal carinae and with two palpomeres

(Figure 6a) (labium without lateral carinae and palps with

three palpomeres in Ottokelleria). Posterior margin of pygid-

ium without setae (setose in Ottokelleria). Protibial spur

shorter than protarsomeres 1 and 2 combined (Figure 6e)

(protibial spur subequal or longer than protarsomeres 1

and 2 combined in Ottokelleria). Additional characters for

diagnosis of males and females include the following:

Description

Male

Frons convex (Figure 4d). Lateral margin of elytra moder-

ately crenulate, with setae shorter than distance between

the crenulations (Figure 6g). Protibia with two ventrolon-

gitudinal carinae (Figure 6e). Protarsomeres 2 and 3 dis-

tinctly longer than wide (Figure 6e). Protarsomeres 3 and

4 with a ventroapical projection dorsoventrally flattened

(Figure 11a,d). Protarsomere 5 with a ventromedial pro-

jection dorsoventrally flattened (Figure 6e). Paramera

unarticulated to the tectum, with prolongations lamelli-

form and apically rounded (Figure 12a–h).

Female

Elytra apically entire (not subtruncated; e.g., Figure 19,e),

with lateral margins sparsely crenulate and bordered

(Figure 14a,b). Protarsomeres lacking keels, projections

and striations. Anterior protarsal claw unenlarged.

Remarks

Based on several morphological characters, we propose the

genus Carinochilus for Ottokelleria marginata and differenti-

ate it from Ottokelleria (for O. dispar). Additionally, Carinochi-

lus differs from monotypic genus Ottokelleria by (opposing

characters of Ottokelleria given in parentheses): (1) mandible

lacking prostheca (mandible with membranous prostheca);

28 BENTO ET AL.

(2) third axillary sclerite with enlarged head (narrowed

head); (3) hindwing with MP3 curved (Figure 6c)(MP3

almost straight); (4) union of the AA +AA3+4formingan

almost 90angle (Figure 6c) (obtuse angled); (5) posterior

margin of pygidium without setae (with setae); (6) male pro-

tarsomere 5 somewhat slender and longer than wide (large

andaswideaslong);(7)marginalsetaeofmaleelytra

shorter than the distance between the crenulations

(Figure 6g) (setae longer than the distance between the

crenulations); (8) paramera fused to the tectum and with

lamelliform prolongations (paramera articulated to the tec-

tum and with prolongations ensiform); and (9) female elytra

apically entire and bordered (female elytra apically subtrun-

cated and unbordered).

Etymology

The composite generic name is masculine in gender and

derives from the Latin carina (=carina) and the Greek

cheilos (=labium) in reference to the labium bearing

laterolongitudinal carinae.

Distribution

(Figure 7)

BRAZIL. Bahia: Iguaí. Espírito Santo: Venda Nova do Imi-

grante, Santa Maria de Jetib

a, Santa Leopoldina. Minas

Gerais: Ipatinga. Paran

a: Campina Grande do Sul, Pinhais.

FIGURE 9 Carinochilus marginatus (Burmeister) comb. nov., male: (a, b) dorsal habitus; (c, d) head and pronotum in dorsal view, arrows indicating

sparsely to moderately crenulate lateral margins of the pronotum. Scales: (a, b) 3 mm and (c, d) 2 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 29

FIGURE 10 Carinochilus marginatus (Burmeister) comb. nov.: (a, b) frontal, (c, d) lateral and (e, f) laterofrontal views of the male head; (g, h)

ventral view of the female head. Scale: 0.5 mm.

30 BENTO ET AL.

Carinochilus marginatus (Burmeister, 1847),

comb. nov.

(Figures 9–12,13a,b and 14–16)

Pachylus marginatus Dejean, 1833, 152 (catalogue) (nomen

nudum); Dejean, 1837, 180 (catalogue) (nomen nudum).

Pachylus marginatus Burmeister, 1847, 24 (original

description; Dynastinae); Lacordaire, 1855, 395 (comment);

Harold, 1869, 1240 (catalogue); Ohaus, 1928, 401 (rede-

scription; transfer to Rutelinae).

Ottokelleria marginata D’Andretta & Martínez, 1957,

269 (combination change); Krajcik, 2007, 90 (checklist).

Material examined

Neotype

Male deposited at ZMHB, labelled: ‘Espirito Santo/S.

Leopoldina/Michaelis’//‘Pachylus marginatus Burm.’

(handwritten) (mouthparts cardmounted). Invalid syntype,

one female of P. dispar incorrectly identified as

P. marginatus (at ISNB), labelled: ‘Gynden’(green, hand-

written)//‘Marginatus/B./Br. C.D.’(white, handwritten with

green border)//‘Pachylus/marginatus/Burm./det. Burmeis-

ter’(white, handwritten)//‘Pachylus marginatus’(white,

handwritten)//‘Collection/E. Candèze’(white with black

border, printed)//‘Type’(white with black border, red

printed letters). See Remarks section for C. marginatus.

Other material

(18 males, 13 females). Paraguay, Paraguari, 26.2 km SE

Ybycui/III.1986, M. Pogue & M. Solis (legs.) (m#, USNM);

Paraguay, Sapucay, 2.I.1905, W. Foster (leg.) (f#, NHM);

Paraguay, Sapucay, 1918, Nevinson Coll. (4 f#, NHM); Brazil,

Espírito Santo, Linhares, III.1953, Pedro Almeida Teles (m#,

CMN); Brazil, Espírito Santo, Venda Nova do Imigrante,

ii.1998, FZVM [leg.] (2m#, CEMT, CERPE); Brazil, Espírito

Santo, Santa Maria de Jetib

a, ‘Rio Bonito’, no date, no col-

lector (2 m#, DZUP); Brazil, Espirito Santo, Santa Leopoldina,

Michaelis (leg.) (m#, ZMHB); Brazil, Espírito Santo, Baixo

Timbuí, 21.XII.1898, L. Ohaus (leg.) (f#, ZMHB); Brazil, S˜

Paulo, Rio Paran

a, Porto Cabral, III–IV.1944, L. Travassos

(leg.) (f#, CMN); Brazil, Minas Gerais, Ipatinga, xi.1989, E. &

P. Grossi (2 f#, CERPE); Brazil, Paran

a, Lucena (f#, ZMHB);

Brazil, Paran

a, Campina Grande do Sul, Estrada da Manda-

çaia, 16.xii.2008–31.i.2009, F.W.C. Leivas (leg) (10 m#, CERPE,

CEMT); Brazil, Paran

a, Pinhais, 12.III.2006, V.C.P. Silva (leg)

(f#, CERPE); Brazil, Bahia, Fry Coll. (f#, NHM); Brazil, Iguaí

(m#, CEMT); Brazil, Mato Grosso do Sul, Dourados, Itaum,

III.1974, M. Alvarenga (leg.) (f#, CMN).

Diagnosis

See Diagnosis section for Carinochilus.

Redescription

Male

Shape, size and colour.Rounded oval. Length 14–16 mm,

width 6–8 mm. Head and pronotum dark brown to reddish

brown (Figure 9a–d). Elytra mostly light brown to yellowish

brown with humeral and lateral margins darker.

Head.(Figure 4d). Frons slightly convex. Interocular width

3.4–4.3 times wider than transverse eye diameter. Clypeus

anteriorly rounded, with lateral angulations near base.

Ocular canthus cariniform, apically rounded and glabrous

(Figure 10c,d). Labrum reduced, at least twice wider than

long and with apical margin rounded or slightly angu-

lated medially (Figure 10a,b). Clypeolabral suture barely

or well defined (Figure 10e,f). Labium (Figure 6a) subtra-

pezoidal and strongly convex, with laterolongitudinal cari-

nae finely crenulate and convergent to the apex of

prementum creating a truncated or slightly rounded mar-

gin; base of mentum with lateral tufts of setae convergent

or not; mentum basally fused to gulamentum and apically

fused to prementum (lacking sutures). Labial palps with

two palpomeres; basal palpomere inserted in a deep

depression. Maxilla (Figure 5a) rugous and moderately

setose, with setae moderately long; galea cuneiform and

short, not reaching the apex of prementum. Mandible

(Figure 5c) reduced and hidden by clypeus, with molar lobe

undeveloped and inerm, incisive margin glabrous and

inerm. Antenna with nine antennomeres, with club trilamel-

late and oblong, longer than antennomeres 2–6combined.

Pronotum.(Figure 9c,d). With surface slightly rugous to

rugostriate. Lateral margins sparsely to moderately crenu-

late anteriorly. Anterior bead complete. Anterior angles

acute and posterior angles rounded. Posterior margin

bisinuous and unbeaded. Scutellum as wide as long and

apically rounded. Surface moderately punctate, punctures

moderate.

Elytra.(Figure 6g). Humeral region moderately setose lat-

erally. Dorsal surface with irregularly aligned punctures

giving an aspect rugous or slightly rugous, with intervals

irregular and convex. Lateral margins bordered and mod-

erately crenulate, with setae shorter than distance

between crenulations. Epipleura gradually narrowed and

incomplete. Hindwing (Figure 6c). 3Ax strongly con-

stricted at middle, with enlarged head. PC basally forming

a membranous flap and distally with minute, serial

branchlets creating a margin serrate and sclerotised.

Anterior margin with setae short and spiniform. MP3

curved; AA +AA3+4 forming a sub-right angle.

Pygidium.Posterior margin bordered, medially thickened

and glabrous.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 31

Thoracic venter.Posterior prosternal process flattened

and triangular, narrowly to broadly rounded apically,

sparsely to moderately setose posteriorly. Mesometaste-

rnum, mesepisternum, mesepimeron, metepisternum

moderately setose. Mesometasternum not projected.

Mesendosternite pyramidal. Mesophragma medially

bilobed and not bordered. Metendosternite in caudal

view T-shaped (Figure 13a), with stem at least three

times longer than wide. Metendosternal arms apically

bifurcate, as long as stem, distinctly longer than wide.

Metendosternite in dorsal view abruptly narrowed ante-

riorly (Figure 13b).

Legs.Mesocoxal and metacoxal plate short and obliquely

truncated with respect to inner margin. Protibia externally

tridentate, with proximal tooth reduced; inner margin

finely crenulate; posterior face with two mediolongitudinal

carinae, inner carina not reaching the base of protibia

(Figure 6e). Protibial spur shorter than protarsomeres 1

and 2 combined, with base in dorsal view narrower than

apex of protarsomere 1. Protarsus shortened and enlarged,

with a finely crenulate ventrolongitudinal keel. Protarsome-

res 3 and 4 with a ventroapical projection dorsoventrally

flattened. Protarsomere 5 enlarged and oval, with a ventro-

medial projection dorsoventrally flattened and apically

truncated. Protarsomere 5 with an internoapical projection

bearing one apical seta. Protarsal, mesotarsal and metatar-

sal empodium bisetose. Anterior protarsal claw enlarged

and strongly curved, with apex unequally cleft. Posterior

protarsal claw simple and apically pointed. Mesofemur and

metafemur, mesotrochanter and metatrochanter with pos-

terior margin finely crenulate. Mesofemur and metafemur

with an anterior row of unequally spaced setae. Mesofe-

mur moderately setose dorsally. Mesotibia and metatibia

with inner and outer margins finely crenulate; inner margin

with long setae, outer margin with spines. Mesotibia and

metatibia with two ventrolongitudinal carinae: inner carina

smooth and outer carina finely crenulate. Mesotibial and

metatibial spurs pointed and finely crenulate ventrolater-

ally. Mesotarsus slightly narrower and longer than

FIGURE 11 Variation of male tarsal claws in Carinochilus marginatus (Burmeister) comb. nov.: (a, d) protarsal, (b, e) mesotarsal and (c, f) metatarsal

claws. Scale: 5 mm.

32 BENTO ET AL.

protarsus and metatarsus. Mesotarsus and metatarsus with

a finely crenulate ventrolongitudinal keel. Mesotarsal and

metatarsal claws finely crenulate ventrally and with ventral

tooth inconspicuous to strongly developed.

Abdomen. Ventrally glabrous. Ventrites 2–4 subequal in

lengths. Ventrite 5 slightly longer than preceding ones.

Ventrites 1–5 with a laterolongitudinal carina well devel-

oped and as wide as elytral border (Figure 5d). Ventrite

FIGURE 12 Variation in the male genitalia, Carinochilus marginatus (Burmeister) comb. nov.: (a, c, e, g) dorsal view and (b, d, f, h) lateral view.

Scale: 1 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 33

6 with posterior margin slightly emarginated medially

and without setae.

Genitalia.Aedeagus (Figure 12a–h). Paramera basally

fused and apically free to each other; shorter than

tectum and unarticulated to it; prolongations lamelliform,

apically rounded and setose, with a median membrane

that allows overlapping horizontal movement.

Female

Shape and size.Body shape rounded oval. Length 16.5–

17.3 mm. Width 7.5–8.3 mm. Females are more robust

than males and differ from themas follows:

Coloration.Dorsum and venter dark brown to light

brown with black reflections. Legs dark brown to black.

FIGURE 13 Comparison of metendosternites between genera of Alvarengiini Frey. Carinochilus marginatus (Burmeister) comb. nov., male:

(a) caudal and (b) dorsal views. Ottokelleria dispar, male: (c) caudal and (d) dorsal views. O. dispar, female: (e) caudal and (f) dorsal views. Scales: (a, b)

1 mm, (c, d) 0.5 mm and (e, f ) 1 mm.

34 BENTO ET AL.

Head.Densely punctate to rugopunctate. Interocular

width 4.2–4.7 times wider than transverse eye diameter.

Frontoclypeal suture absent. Anterior margin of clypeus

lacking lateral angulations near the base in dorsal view.

Labium smooth, with laterolongitudinal carinae smooth

and thick (Figure 10g,h). Maxilla externally rounded, sub-

quadrate or with a broadly rounded lobe (Figure 10g,h).

Antennal club subglobose, shorter than antennomeres 2–

6 combined.

Pronotum. Densely punctate, with lateral margins bor-

dered and smooth.

Elytra. With intervals regular and convex. Lateral

margins sparsely crenulate. Apex rounded (Figure 14b),

not subtruncate (Figure 19b,e). Epipleuron shelf-like,

gradually tapering from base to metacoxa and

from ventrites 2–7 to apex (not subparallel, e.g.,

Figure 19b).

Thoracic venter.Sparsely setose and densely punctate,

with crateriform punctures.

Legs.Anterior, median and posterior legs with margins

and carinae completely smooth. Protibial spur robust,

with base in dorsal view as wide as apex of protarsomere

1. Protarsomeres simple, ventrally smooth and inerm.

Anterior protarsal claw unenlarged and with a strong ven-

tromedial tooth. Mesotrochanter and metatrochanter and

mesofemur and metafemur with margins smooth.

FIGURE 14 Carinochilus marginatus (Burmeister) comb. nov., female: (a, b) dorsal habitus and (c, d) head and pronotum in dorsal view. Scales: (a,

b) 3 mm and (c, d) 1 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 35

Metafemur ventrally glabrous. Inner metatibial spur api-

cally rounded. Mesotarsomeres and metatarsomeres with-

out ventrolongitudinal keel. Mesotarsomeres and/or

metatarsomeres 1–4 each with a ventroapical projection.

Metatarsal claws ventrally smooth, toothed or untoothed.

Abdomen.Sparsely setose ventrally.

External genitalia.(Figure 15a–f). Gonocoxites with pos-

terior margin moderately setose, setae long. Proximal

gonocoxites large, subtriangular to subquadrate in dor-

sal view, with posterior margin subtruncated to some-

what rounded. Distal gonocoxites small and almost

hidden by proximal gonocoxites. Paraprocts short to

long, almost two to three times longer than wide at mid-

dle, with apex narrowly to broadly rounded. Proctiger

medially divided into two small to large, subrectangular

to rounded pieces.

Remarks

Burmeister (1847) validly described this species in Pachy-

lus (=Ottokelleria) from Dejean (1833,1837) and stated

that the species was ‘Brasilien’. His description was

based on female specimens as ‘reported by Dupont’

(i.e., Dupont provided Burmeister with the characteristics

of the species, but Burmeister did not study any speci-

mens directly). The length range in the description (‘7–8

lines’) indicated that at least two specimens were part of

the original type series. According to the description, the

species is ‘somewhat larger’than O. dispar, which was

described as ‘3.5–5 lines’. A single female specimen

bearing a type label was located at ISNB. It is identified

as P. marginatus and bears a Burmeister identification

label (Figure 16). The handwriting of the labels can be

assigned clearly to Dupont and Candèze. Indeed,

because Burmeister described this species based only on

characteristics ‘reported’by Dupont, the specimen is not

expected to bear Burmeister labels (because Burmeister

did not have specimens for examination). The size range

was originally reported in the ‘line’standard of measure-

ment, which has several variations depending on the

country system adopted. Because the characteristics of

the specimens were reported by Dupont and described

by Burmeister, the conversion from the line standard to

millimetres was made considering both the French sys-

tem (2.23 mm per line) and the German system

FIGURE 15 Variation of the female external genitalia of Carinochilus marginatus (Burmeister) comb. nov.: (a, d) ventral, (b, e) lateral and (c, f )

dorsal views of the female external genitalia. pct, proctiger; pgx, proximal gonocoxite; ppt, paraproct. Scale: 0.5 mm.

36 BENTO ET AL.

(2.18 mm per line), thus providing a length range of

15.61–17.84 mm (French) and 15.26–17.44 mm (German)

for the type specimens and the species description. This

size range is consistent with the length of female

specimens of C. marginatus that we examined (16.5–

17.3 mm). The specimen in Figure 16 is labelled as the

‘type’specimen, but its characters are incompatible with

the original description of the species in terms of length

and elytral sculpturing. In addition, the original descrip-

tion of C. marginatus includes the following: ‘…the

elytra with three distinct ribs and less clearly raised wrin-

kles’(translated from German), referring to three convex

intervals of rugous elytra (e.g., Figures 9a,b and 14a,b).

However, these characteristics are not observed in the

ISNB specimen (Figure 16). Instead, the specimen is con-

specific with O. dispar based on body length (12.9–

15.4 mm) and form of the elytra (punctostriate and non-

rugous, with numerous convex intervals). We concluded

that this specimen cannot be the valid type of

C. marginatus because it is incompatible with the original

description and redescriptions. Additionally, based on

determination labels of authoritative experts including

Ohaus, Lacordaire and Arrow, the species concept for

C. marginatus doesnotconcurwiththesyntype

specimen in Figure 16 (which is, in fact, conspecific with

O. dispar). Ohaus (1928) subsequently redescribed both

sexes from Espírito Santo, and his description is consis-

tent with Burmeister’s(

1847) original description. Ohaus

(1928) described the male as follows: ‘Reddish brown

on the obverse, with the disc of the wings [elytron] light

reddish brown …the lateral border of the pronotum is

densely [not in the same sense of this work] notched …

on median and posterior legs the outer claw has a

barely visible [vestigial], basal tooth’(translated from

German). Dupont’s collection was split between ISNB,

NHM and the Museum National d’Histoire Naturelle

(Paris, France). The specimen was not located in these

collections. In fact, because Burmeister did not examine

the specimen directly (Dupont provided characters for

the description), locating the type is a tricky task. Thus,

weconsiderthetypespecimen(s)tobelostandwe

designateaneotypebasedonamalespecimenthat

was used by Ohaus (1928) to redescribe C. marginatus.

The specimen is from Santa Leopoldina, Espírito Santo

state, Brazil (deposited at ZMHB), and is conspecific with

the females originally (Burmeister 1847)andsubse-

quently described (Lacordaire 1855;Ohaus1928)based

on body size, punctation of head and elytra, and elytral

FIGURE 16 Pachylus marginatus Burmeister (=Ottokelleria marginata). Invalid female syntype at ISNB, incorrectly identified as P. marginatus

(specimen is, in fact, conspecific with Ottokelleria dispar): (a) head and pronotum in dorsal view, (b) head in ventral view, (c) elytral apex in dorsal view,

(d) metatarsus and (e) labels.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 37

sculpturing. The taxon has had a complicated taxonomic

history; designation of a neotype male will help to stabilise

nomenclature and allow for future comparisons. As per the

ICZN (1999), this specimen is publicly accessible and depos-

ited in a well-recognised scientific institution (ZMHB).

Etymology

The specific epithet marginatus is Latin for margino and

means ‘enclosed with a border’.

Distribution

(Figure 7)

PARAGUAY. Paraguari, Sapucay. BRAZIL. Bahia: Iguaí.

Espírito Santo: Santa Maria de Jetib

a, Venda Nova do

Imigrante, Linhares, Santa Leopoldina. Minas Gerais:

Ipatinga. Paran

a: Campina Grande do Sul, Pinhais. Mato

Grosso do Sul: Dourados. S˜

ao Paulo: Porto Cabral.

Ottokelleria D’Andretta & Martínez, 1957

(Figures 2a,4a–c,e,5b,f,g and 6b,d,f,h)

Pachylus Dejean, 1833, 152 (catalogue) (nomen nudum);

Dejean, 1837, 180 (catalogue) (nomen nudum).

Pachylus Burmeister, 1847, 23 (original description; Dynasti-

nae); Lacordaire, 1855, 395 (redescription); Harold, 1869, 1240

(catalogue); Ohaus, 1900, 212; Ohaus, 1928, 396 (redescription;

transfer to Rutelinae); Jameson, 2000b, 332 (synonym).

Type species: Pachylus dispar Burmeister, 1847 (subse-

quent designation).

Ottokelleria D’Andretta & Martínez, 1957, 268 (nomen

novum for Pachylus Burmeister [preoccupied, nec Kollar in

Koch 1839]; type species designation); Jameson, 2000b,

332 (priority, valid name).

Type species: Pachylus dispar Burmeister, 1847 (original

designation).

Alvarengius Frey, 1975, 84 (original description; Melo-

lonthinae); Jameson, 2000b, 335 (relationships); Smith &

Evans, 2005, 54 (transfer to Rutelinae); Smith, 2006,

173 (catalogue); Krajcik, 2007, 20 (checklist); Bouchard

et al., 2011 (catalogue); Moore et al., 2018, 5, tab. 1 (inad-

vertent error in spelling). Syn. nov.

Type species: Alvarengius silphoides Frey, 1975 (designa-

tion by monotypy).

Diagnosis

Adult specimens in the genus Ottokelleria are easily

distinguished from Carinochilus by the following

characters: ocular canthus setose (Figure 2a) (glabrous in

Carinochilus). Labium without laterolongitudinal carinae, with

three palpomeres (Figure 6b) (labium with laterolongitudinal

carinae, with two palpomeres in Carinochilus). Elytra punc-

tostriate (Figure 6h) (with rugous surface in Carinochilus).

Protibial spur subequal or longer than protarsomeres 1 and

2 combined (Figure 6f) (shorter than protarsomeres 1 and 2

combined in Carinochilus). Additional characters for diagno-

sis of males and females include the following:

Description

Male

Frons flat (Figure 2a). Lateral margin of elytra densely crenu-

late, with setae longer than distance between the crenula-

tions (Figure 6h). Protibia with one ventrolongitudinal carina

(Figure 6f). Protarsomeres 2 and 3 apically wider than long

(Figure 6f). Protarsomeres 4 and 5 each with a ventroapical

projection striated and laterally flattened (Figure 17f). Para-

mera articulated to the tectum, with prolongations ensiform

(Figure 18a–d).

Female

Elytra apically subtruncated (Figure 19b), with lateral mar-

gins smooth and not bordered (Figure 19e). Tarsi without

keels, projections and striations. Protarsomere 5 and ante-

rior protarsal claw unenlarged (Figure 5f).

Remarks

See Remarks section of Carinochilus gen. nov. for a con-

trast between Carinochilus and Ottokelleria and for sexual

differences in the genera.

Based on the new synonymy herein proposed

(ICZN 1999, Article 61.3.1) between the type species

A. silphoides Frey, 1975 and O. dispar (Burmeister, 1847),

the monotypic genus Alvarengius Frey, 1975 is considered

to be a junior subjective synonym of Ottokelleria

D’Andretta & Martínez, 1957. Therefore, if future studies

consider Alvarengius not to be congeneric with Ottokel-

leria, the name remains available to designate the origi-

nally proposed taxon (Article 10.6).

In a synopsis of the cyclocephaline beetles

(Dynastinae), Moore et al. (2018, tab. 1) placed Alvarengius

in synonymy under Pachylus. In the context of this work,

Moore et al. (2018) do not propose any nomenclatural act

and were concerned mostly with providing historical back-

ground for cyclocephaline scarabs. Furthermore, assuming

that the names Ottokelleria and Pachylus are objective syn-

onyms, we interpret this act as an inadvertent error in spell-

ing the name Alvarengius in place of Ottokelleria.

Another taxon name was entered into the literature,

‘Pachylus euryalus’. This name appeared in Dejean (1833,

38 BENTO ET AL.

1837),butitwasnotdescribed.Toourknowledge,itisnot

applied to any taxon. We consider the name a nomen

nudum.

Etymology

The generic epithet is a replacement name created in

honour of Don Otto Paulo Keller, a Brazilian entomolo-

gist dedicated to the study of the systematics and

biology of Cicadidae (Hemiptera) (see D’Andretta &

Martínez 1957).

Distribution

(Figure 8)

BRAZIL. Minas Gerais: Passa Quatro, Virgínia. Espírito Santo:

no locality data. Rio de Janeiro: Nova Friburgo, Rio de

Janeiro, Itatiaia, Petr

opolis. S˜

ao Paulo:Sales



opolis, S˜

ao José

do Barreiro, Mogi das Cruzes, S˜

ao Paulo. Paran

a: Reserva

Biol

ogica Bom Jesus, Piraquara, S˜

ao José dos Pinhais.

Ottokelleria dispar (Burmeister, 1847)

(Figs 13c–f, 17–22)

FIGURE 17 Ottokelleria dispar (Burmeister), male, showing colour variation: (a) dorsal habitus, (b) ventral habitus, (c, d) dorsal habitus, (e) head in

ventral view and (f) protarsus. Scales: (a, b) 3 mm, (c, d) 5 mm, (e) 0.3 mm and (f) 0.2 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 39

Pachylus serratulus Dejean, 1833, 152 (catalogue) (nomem

nudum); Dejean, 1837, 180 (catalogue) (nomen nudum);

Harold, 1869, 1240 (synonymy).

Pachylus dispar Burmeister, 1847, 24 (original description;

Dynastinae); Lacordaire, 1855,395(comment);

Lacordaire, 1869, pr. 35 n. 4 (atlas); Harold, 1869, 1240 (cata-

logue); Ohaus, 1928, 400 (redescription; transfer to

Rutelinae).

Ottokelleria dispar D’Andretta & Martínez, 1957,

269 (new combination); Krajcik, 2007, 90 (checklist).

Alvarengius silphoides Frey, 1975, 84 (original

description; Melolonthinae); Smith & Evans, 2005,

54 (transfer to Rutelinae); Krajcik, 2007,20(checklist).

Syn. nov.

Material examined

Lectotype

Male (here designated). (Figure 22)ofP. dispar

at MLUH, labelled: ‘dispar */♀marginatus/Dej./

Bras. Imr.’(green with black borders,

handwritten).

Paralectotypes

Same data as lectotype (2 f#, MLUH); ‘Pachylus/Dej.

Brm.’(white, handwritten)//‘dispar Brm/Bras. Bs.’

(white, handwritten)//‘Syntipus’(red, handwritten)//

‘MACN—En/10773’(white, printed) (f#, MACN)

(Figure 20a–c). Holotype male (Figure 21)ofA. silphoides at

NMB labelled: ‘Rio de Janeiro—Gb/(Floresta da Tijuca)/

IV.1966/M. Alvarenga col. [leg]’(white with black margins,

printed)//‘Type/Alvarengius silphoides/G. Frey 1974’(red,

printed and handwritten) (NMB). Paratype male of

A. silphoides at NMB labelled with same data as holotype

and with male genitalia dissected and mounted.

Other material

(84 males, 39 females). Brazil, Paran

a, Piraquara, 05.ii. 2007,

Grossi & Rocha (leg.) (2 m#, CERPE); idem, but 13.iii.2012,

F. Leivas & P. Grossi [leg.] (10 m#, CERPE, CEMT); Brazil,

Paran

a, S˜

ao José dos Pinhais, Serra do Mar, 02.iii.1987

(1 m#, DZUP); Brazil, S˜

ao Paulo, Jabaquara, 20.xii.1943,

H. Zellibor Collection (1 m#, CMN); Brazil, S˜

ao Paulo, Sales-



opolis, Estaç˜

ao Biol

ogica de Boracéia, 9–15.i.2003, Malaise,

A.C.C. Macedo & J. S. Freitas (leg.) (4 m#, MZUSP); idem,but

FIGURE 18 Ottokelleria dispar (Burmeister), (a–d) variation of the male paramera in caudal view. Scale: 0.5 mm.

40 BENTO ET AL.

FIGURE 19 Ottokelleria dispar (Burmeister), female, showing colour variation: (a, b) dorsal habitus, (c) ventral habitus, (d) head in ventral view and

(e) abdomen in caudal view. Scales: (a–c) 3 mm, (d) 0.5 mm and (e) 1 mm.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 41

22.iii–26.iv.2001, S. A. Casari & I. B. Francini (leg.) (18 m#,

MZUSP); Brazil, S˜

ao Paulo, Mogi das Cruzes, Parque das

Neblinas, 24–28.ii.2015, RVNunes (leg.) (10 m#, CERPE,

CEMT); idem, but iii.2006, M. Uehara (leg.) (1 f#, CERPE);

Brazil, S˜

ao Paulo, S˜

ao José do Barreiro, Serra da Bocaina,

xi.1968, Alvarenga & Seabra (leg.) (1 m#, DZUP); Brazil, Rio

de Janeiro, Corcovado, 1956, Martínez Collection (1 m#,

1 f# CMN); Brazil, Rio de Janeiro, Parque Nacional do Ita-

tiaia, ii.1957, Martínez Collection (1 f#, CMN); Brazil, Rio de

Janeiro, Nova Friburgo, 3–11.iii.1990, S. A. Marshall (1 m#,

CMN); Brazil, Rio de Janeiro, Nova Friburgo, Macaé de Cima,

i.2000, P. Grossi (Leg.) (2 m#, CERPE); idem, but i. 2005,

P. Grossi (leg.) (1 f#, CERPE); idem, but i.2006, B. Miller (leg.)

(2m#, CERPE); idem, but xi.2004 (9 m#, CERPE, CEMT); idem,

but xii.2003 (1 m#, CERPE); idem, but xi.2008 (1 m#, CERPE);

idem, but ii.2000, Geniér, Lopes & Vaz-de-Mello (Legs.) (5 f#,

CERPE, CEMT); idem, but xi.1998 (8 m#, CERPE); Brazil, Rio

de Janeiro, Teres

opolis, ii.1991, A. Bello (Leg.) (1 m#, CERPE);

Brazil, Rio de Janeiro, Itatiaia, xii.1964, A. Bello (Leg.) (1 f#,

CERPE); idem, but 3.iv.1932, J. F. Zik

an Collection (1 f#,

CEIOC); Brazil, Rio de Janeiro, Floresta da Tijuca, Pico da

Tijuca, 8.ii.1953 (1 f#, CEIOC); idem, but Corcovado, iii.1961,

M. Alvarenga (Leg.) (1 f#, DZUP); Brazil, Rio de Janeiro, Insti-

tuto Zoobotânico de Morro Azul, xi–xii. 2002, Paulo de

Frontin (leg.) (1 f#, CERPE); Brazil, Rio de Janeiro, Itatiaia, 5.

iii.1932, I. Sonne Flieg (leg.), Coleç ˜

ao J. F. Zik

an (1 m#,

CEIOC); idem, but 15.iii.1941(1 m#, CEIOC); idem, but Mar-

omba, 10.ii.1936 (1 m#, CEIOC); idem, but 13.ii.1928 (1 f#,

CEIOC); idem, but 13.ii.1926 (1 m#, ZMHB); idem,but3.

iii.1929 (1 f#, CEIOC); Brazil, Rio de Janeiro, Itatiaia, 11.

iii.1914, Coleç˜

ao J. F. Zik

an (1 f#, CEIOC); idem, but 1.ii.1914

(1 f#, CEIOC); idem, but 16.ii.1916 (1 f#, ZMHB); idem,but

1926 (1 f#, ZMHB); Brazil, Rio de Janeiro, Itatiaia, 5.ii.1959,

Hebert, Coleç˜

ao Fiocruz (1 f#, CEIOC); Brazil, Rio de Janeiro,

Corcovado, 9.i.1899, F. Ohaus (leg.) (1 m#, ZMHB); Brazil, Rio

de Janeiro, Petr

opolis, 15.i.1899, F. Ohaus (leg.) (1 m#,

ZMHB); idem, but 19.i.1899 (1 f#, ZMHB); idem,nodate

(1 m#, 1 f#, ZMHB); Brazil, Rio de Janeiro, G

avea, 21.xi.1904,

Vasquez (leg.) (1 f#, ZMHB); Brazil, Rio de Janeiro, Serra dos

Órg˜

aos, 1902, S. R. Wagner (leg.) (1 f#, BMNH); Brazil, Rio de

Janeiro, 1905, Frey Collection (2 f#, BMNH); Brazil, Espírito

Santo, no data, J. Moser Collection (1 m#, ZMHB); Brazil,

Minas Gerais, Passa Quatro, 26.i.1923, J. Zik

an (leg.) (1 m#,

ZMHB); Brazil, Minas Gerais, Passa Quatro, Serra dos Coel-

hos, 26.i.1923, Coleç˜

ao J. F. Zik

an (1 m#, CEIOC); Brazil,

Minas Gerais, Virgínia, Fazenda Campos, 25.i.1916, Coleç˜

J. F. Zik

an (1 f#, CEIOC); Brazil, no data (1 m#, 1 f#, BMNH);

Brazil, no data (5 f#, ZMHB; 1 f#, ISNB); Brazil, no data, Ger-

mar Collection (1 f#, ZMHB); Brazil, no data, Nevinson Col-

lection (1 f#, BMNH); No data (1 f#, ISNB); Brazil, no data, E.

Candèze Collection (1 f#, ISNB).

Diagnosis

See Diagnosis section for Ottokelleria D’Andretta &

Martínez, 1957.

Redescription

Male

Shape, size and colour.Body shape rounded oval.

Length 9.6–12.1 mm. Width 4.3–5.9 mm. Mostly brown.

Head black, with clypeus ventrally yellowish. Pronotum

black, with lateral margins and hypomera yellow.

Elytra light to dark brown, with sutural and lateral

margins darker. Coxae, anterior and median femura yel-

lowish.3pt?>

Head.(Figures 2a and 4a). Rugopunctate. Interocular

width 4.3 times wider than transverse eye diameter. Frons

flattened. Frontoclypeal suture absent. Anterior margin of

clypeus raised and slightly angled medially in dorsal view.

Ocular canthus cariniform, apically rounded and setose,

setae moderately long. Labrum reduced, with apical mar-

gin rounded to slightly angled medially, wider than long.

Labium (Figure 6b) subtrapezoidal and strongly convex,

moderately setose, with setae long. Labial palp with three

palpomeres, with apical palpomere slightly flattened,

longer than palpomeres 1 and 2. Maxilla rugous, with a

rounded, external lobe. Stipes moderately setose, with

moderately long setae. Maxillary palp with four palpo-

meres, apical palpomere longer than palpomeres 2 and 3

combined. Antenna (Figure 4b) with nine antennomeres,

with club trilamellate and oblong, longer than antenno-

meres 2–6 combined.

Pronotum.Densely punctate, punctures moderate. Ante-

rior margin beaded. Lateral margins moderately

to densely crenulate anteriorly, each crenulation with a

minute seta. Anterior angles acute. Posterior

margin bisinuous and not beaded, with lateral angles

rounded.

Scutellum.As long as wide and apically rounded. Moder-

ately punctate, with punctures moderate.

Elytra.Humeral region densely setose, with moderately

long setae. Disc punctostriate, with intervals convex and

regular. Lateral margins bordered and densely crenulate,

with setae longer than distance between the crenulations.

Epipleura gradually narrowed and incomplete. Sutural

stria apically deep.

Hindwing.(Figure 6d). 3Ax with narrow head. PC basally

forming a narrow, membranous flap and distally with minute,

serial branchlets creating a margin serrate and sclerotised.

Anterior margin with moderately long setae. MP3 almost

straight. AA +AA3+4 forming obtuse, rounded angle.

Pygidium.Densely punctate, posterior margin medially

thickened and setose.

42 BENTO ET AL.

Thoracic venter.(Figure 17b). Posterior prosternal process

triangular, flattened and moderately setose posteriorly.

Mesometasternum, mesepisternum, mesepimeron and

metepisternum moderately setose, with punctures mod-

erate and setae long. Mesometasternum not projected.

Mesendosternite pyramidal. Mesophragma medially

bilobed. Metendosternite in caudal view (Figure 13c)

T-shaped, with stem at least three times longer than wide.

Metendosternal arms as long as stem and distinctly lon-

ger than greater width, apically bifurcate. Metendoster-

nite in dorsal view gradually narrowed anteriorly

(Figure 13d).

Legs.Procoxae and mesocoxae prominent. Protibia exter-

nally tridentate, with inner margin finely crenulate and

ventral surface with one longitudinal carina. Protibial spur

equal or longer than protarsomere 1, with ventrolateral

margins finely crenulate. Protarsus shorter and wider than

mesotarsus and metatarsus. Protarsomeres with a ventro-

longitudinal keel finely crenulate. Protarsomeres 4 and

5 each with a ventroapical projection laterally flattened

and internally striated (Figures 5e,g and 17f). Protar-

somere 5 enlarged oval, with a slender, internoapical pro-

jection bearing an apical seta. Empodia bisetose. Anterior

protarsal claw enlarged and strongly curved, with apex

rounded and unequally cleft. Posterior protarsal claw sim-

ple. Mesotrochanter and metatrochanter, mesofemur and

metafemur with posterior margin finely crenulate. Mesoti-

bia and metatibia with margins finely crenulate: inner

margin with long setae, outer margins with spines. Meso-

tibia and metatibia with two ventrolongitudinal carinae:

inner carina smooth and outer carina finely crenulate.

Mesotibial and metatibial spurs pointed and finely crenu-

late ventrolaterally. Mesotarsomeres and metatarsomeres

with a finely crenulate ventrolongitudinal keel. Anterior

mesotarsal and metatarsal claw with a ventromedial tooth

distinctly developed; posterior claw simple. Metacoxa

moderately setose. Metatarsomeres with spine-like ven-

tral setae.

Abdomen.Sparsely setose. Ventrites slightly shortened

medially. Ventrites 1–5 with laterolongitudinal carina well

developed and as wide as elytral border, somewhat inter-

rupted by peritreme of Ventrite 5. Ventrite 6 with poste-

rior margin moderately emarginated medially and with

mediolateral setae moderately long to long.

Genitalia.Aedeagus (Figure 18a–d). Paramera articulated

to tectum. Prolongations basally fused and apically free to

each other, with basal margin deeply to slightly emargi-

nated medially. Prolongations ensiform and longer than

half of tectum, with apex pointed and densely setose ven-

trally; slightly curved in lateral view.

Variation.Males vary from light brown to black (rarely),

with and without yellow lateral maculae on the pronotum

(Figure 17a–d).

Female

Size.Length 12.9–15.4 mm. Width 6–7.4 mm.

Females differ from males as follows:

Colour.Dark to light brown.

Head.(Figures 4e and 19a,b,d). Moderately punctate, with

punctures moderate. Interocular width 4.7 times wider

than transverse eye diameter. Anterior margin of clypeus

rounded to subrectangular, without median angulation.

Antennal club subglobose, shorter than antennomeres

2–6 combined (Figure 4c).

Pronotum.(Figure 19a,b). Moderately punctate, punctures

moderate. Lateral margins smooth.

Elytra.(Figure 19a,b,e). Lateral margins bordered and

smooth. Apex subtruncated and unbordered (Figure 19b,e).

Epipleuron shelf-like, gradually tapering from base to meta-

coxa, subparallel from ventrites 2–7tosubapex.

Hindwing.Anterior margin with setae short.

Thoracic venter.(Figures 19c and 20b). Mesometaste-

rnum, mesepisternum, mesepimeron and metepisternum

glabrous and sparsely to moderately punctuate. Meten-

dosternite with stem twice longer than wide. Metendos-

ternal arms pointed and slightly curved (Figure 13e,f).

Legs.Protibia internally smooth. Protibial spur basally

equal or wider than apex of protarsomere 1. Tarsi inerm.

Protarsomere 5 unenlarged. Anterior protarsal claw unen-

larged, with a strong ventromedial tooth (Figure 5f).

Mesotrochanter and metatrochanter, mesofemur and

metafemur with posterior margin smooth. Mesotibia

and metatibia with margins and carinae smooth. Mesotar-

someres and metatarsomeres 1–4 chalice-shaped. Inner

metatibial spur apically rounded.

Abdomen.Posterior margin of ventrite 6 unemarginated.

Variation.Females vary from light brown to dark brown

(Figure 19a,b). Basal margin of paramera exhibits different

degrees of emargination (Figure 18a–d).

Remarks

For over 190 years, this species has remained a mystery,

its classification has been fraught with complications and

its identity has been obscured. Burmeister (1847)

described P. dispar based on at least one male and one

female specimen, both of which are deposited in his col-

lection in Halle, Germany. Specimens were sent by Carl

Heinrich Bescke, a dipterist who collected in Brazil in the

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 43

FIGURE 20 Pachylus dispar Burmeister (=Ottokelleria dispar), paralectotype female (MACN): (a) dorsal habitus, (b) ventral habitus and (c) labels.

Scale: 2 mm.

44 BENTO ET AL.

early 1800s. We located one male and three female syn-

types deposited at MACN and MLUH (Figures 20 and 22).

For purposes of nomenclatural stability, we designate the

male specimen from MLUH as the lectotype (ICZN 1999,

Article 74). Morphological analysis of the type series of

A. silphoides and O. dispar demonstrated that these spe-

cies are conspecific. As such, A. silphoides is a new syno-

nym of O. dispar.

FIGURE 21 Alvarengius silphoides Frey (=Ottokelleria dispar (Burmeister) syn. nov.), holotype male: (a) dorsal habitus; aedeagus in (b) caudal view,

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 45

FIGURE 22 Pachylus dispar Burmeister (=Ottokelleria dispar), lectotype male (here designated, MLUH): (a) dorsal habitus, (b) ventral habitus and

(c, d) labels.

46 BENTO ET AL.

Etymology

The specific epithet is Latin for ‘dispar’, which means

‘different’.

Distribution

(Figure 8)

BRAZIL. Minas Gerais: Passa Quatro, Virgínia. Espírito Santo:

no locality data. Rio de Janeiro: Nova Friburgo, Rio de

Janeiro, Itatiaia, Petr

opolis. S˜

ao Paulo:Sales



opolis, S˜

ao José

do Barreiro, Mogi das Cruzes, S˜

ao Paulo. Paran

a:Reserva

Biol

ogica Bom Jesus, Piraquara, S˜

ao José dos Pinhais.

ACKNOWLEDGEMENTS

All the curators and collections mentioned in this

paper are thanked for providing material and facilitations

to study type and non-type material, especially Hans-

Joachim Händel (MLUH), Bernard Jäger (ZMHB) and

Matthias Seidel (Natural History Museum Wien, Vienna,

Austria) for kindly hosting MB in their institutions during

his sandwich PhD period. For their collection and type

specimen expertise, we thank Karla Schneider (MLUH), Pol

Limbourg and Alain Drumont (ISNB), Carsten Zorn

(Gnoien, Germany) and Matthias Seidel (Natural History

Museum Wien, Vienna, Austria). For expertise with melo-

lonthine characters, we are grateful to Art Evans (Virginia

Museum of Natural History, Martinsville, VA, USA). This

work was developed as part of MB’s master’s degree and

supported by the Brazilian agencies Conselho Nacional de

Desenvolvimento Científico e Tecnol

ogico (CNPq) and

Coordenaç˜

ao de Aperfeiçoamento de Pessoal de Nível

Superior (CAPES—Finance Code 001). MB was also

financed by the CAPES (Brazil) through doctoral (Finance

Code 001) and sandwich doctorate abroad (PDSE; Code

88881.690179/2022-01) scholarships. PG was funded by

CNPq (Process Numbers 449366/2014-6, 424048/2018-3

and 309786/2019-3). We are grateful to Andrew Smith

and José Mondaca for their careful review and rich sug-

gestions for this manuscript.

CONFLICT OF INTEREST STATEMENT

The authors declare no conflicts of interest.

ORCID

Matheus Bento https://orcid.org/0000-0001-9846-9728

Mary Liz Jameson https://orcid.org/0000-0002-3076-

2028

Paschoal Grossi https://orcid.org/0000-0001-6601-5967

Claudio Ruy Vasconcelos da Fonseca https://orcid.org/

0000-0002-1955-288X

ENDNOTE

This quote referenced Endrödi (1985) who considered Oryctomorphus

to be a member of the tribe Pentodontini (Dynastinae).

REFERENCES

Ahrens, D., Schwarzer, J. & Vogler, A.P. (2014) The evolution of scarab

beetles tracks the sequential rise of angiosperms and mammals.

Proceedings of the Royal Society, 281(1791), 20141470. Available

from: https://doi.org/10.1098/rspb.2014.1470

Ahrens, D., Scott, M. & Vogler, A.P. (2011) The phylogeny of monkey bee-

tles based on mitochondrial and ribosomal RNA genes (Coleoptera:

Scarabaeidae: Hopliini). Molecular Phylogenetics and Evolution,

60(3), 408–415. Available from: https://doi.org/10.1016/j.ympev.

2011.04.011

Bates, H.W. (1888) Insecta. Coleoptera. Rutelidae. In: Salvin, F.D. &

Godwin, O. (Eds.) Biologia Centrali Americana, Vol. 2, pp. 216–296.

Bento, M. & Grossi, P.C. (2019) Description of the female of Spodochlamys

marahuaca Jameson & Ratcliffe, 2011 (Melolonthidae, Rutelinae,

Anatistini) and additional records for the Brazilian species of the

genus. Acta Amazonica, 49(3), 193–196. Available from: https://doi.

org/10.1590/1809-4392201803571

Beutel, R.G. & Leschen, R.A. (Eds). (2016) Coleoptera, beetles. Morphology

and systematics. Berlin: Walter de Gruyter, p. 684.

Blackwelder, R.E. (1944) Checklist of the Coleopterous insects of Mexico,

Central America, the West Indies, and South America. Parts 1–6, Bulle-

tin, Vol. 185. United States National Museum, pp. 1–1492.

Blanchard, C.E. (1850 [1851]) Ordre des Coleoptera. In: Milne-Edwards, H.,

Blanchard, C.E. & Lucas, H. (Eds.) Muséum d’Histoire Naturelle de

Paris. Catalogue de la Collection Entomologique. Classe des Insectes,

Vol. 1. Paris: Gide and Baudry, pp. 129–240.

Bouchard, P., Bousquet, Y., Davies, A.E., Alonso-Zarazaga, M.A.,

Lawrence, J.F., Lyal, C.H.C., et al. (2011) Family-group names in Cole-

optera (Insecta). ZooKeys, 2(88), 1–972.

Browne, D.J. & Scholtz, C.H. (1994) The morphology and terminology of

the hindwing articulation and wing base of the Coleoptera, with

specific reference to the Scarabaeoidea. Systematic Entomology, 19,

122–143.

Burmeister, H. (1847) Handbuch der Entomologie, Vol. 5, pp. 23–24.

D’Andretta, M.A.V. & Martínez, A. (1957) Rutelidos nuevos o poco conoci-

dos (Col. Scarabaeoid. Rutelidae). Revista Brasileira de Entomologia,

7, 263–272.

Dejean, P.F.M.A. (1833) Catalogue des Coleoptères de la Collection de

M. de le Comte Dejan. Troisième édition, revue, corrigée et

augmentée. [Livraison 5]. Paris: Méquignon-Marvis Père et Fils,

pp. 1–443.

Dejean, P.F.M.A. (1837) Catalogue des Coleoptères de la Collection de

M. de le Comte Dejan. [Livraisons 1–2]. Paris: Méquignon-Marvis,

pp. 150–195.

Endrödi, S. (1966) Monographie der Dynastinae (Coleoptera, Lamellicor-

nia). Entomologische Abhandlungen Staatlichen Museum für Tier-

kunde, 33, 1–457.

Endrödi, S. (1985) The Dynastinae of the world. Dordrecht: Dr. W. Junk,

pp. 1–848.

Evans, A.V. & Smith, A.B.T. (2020) On the tribal classification of the Nearc-

tic Melolonthinae (Coleoptera: Scarabaeidae), with descriptions of

new species of Acoma Casey, 1889. Zootaxa, 4748, 51–77.

Frey, G. (1975) Eine neue Sudamerikanische gattung und tribus der

Melolonthiden. Entomologische Arbeiten aus dem Museum, 26,

84–86.

Google Maps. https://www.google.com.br/maps [Accessed January–

December 2018].

Gunter, N.L., Weir, T.A., Slipinski, A., Bocak, L. & Cameron, S.L. (2016) If

dung beetles (Scarabaeidae: Scarabaeinae) arose in association

with dinosaurs, did they also suffer a mass co-extinction at the

K-Pg boundary? PLoS ONE, 11(5), e0153570. Available from: https://

doi.org/10.1371/journal.pone.0153570

Harold, B. (1869) Catalogus Coleopterum: Scarabaeidae, Vol. 4. Monachii,

p. 1240.

Harris, R.A. (1979) A glossary of surface sculpturing. Occasional Papers in

Entomology, 28, 1–31.

International Commission on Zoological Nomenclature. (1999) Interna-

tional Code of Zoological Nomenclature, 4th edition. London: 106pp.

REVISION OF THE LEAF CHAFER TRIBE ALVARENGIINI FREY 47

Jameson, M.L. (1990) Revision, phylogeny and biogeography of the gen-

era Parabyrsopolis Ohaus and Viridimicus, new genus (Coleoptera:

Scarabaeidae: Rutelinae). Coleopterists Bulletin, 44, 377–422.

Jameson, M.L. (1998) Phylogenetic analysis of the subtribe Rutelina and

revision of the Rutela generic groups (Coleoptera: Scarabaeidae:

Rutelinae: Rutelini). Bulletin of the University of Nebraska State

Museum, 14, 183 pp.

Jameson, M.L. (2000a) Key to tribes of Rutelinae (Scarabaeidae). In:

Ratcliffe, B.C. & Jameson, M.L. (Eds.) Generic guide to New World

scarab beetles.http://unsm-ento.unl.edu/Guide/Guide-introduction/

Guideintro.html [Accessed 6th July 2020].

Jameson, M.L. (2000b) Synopsis of the Mexican and Guatemalan genera

Rutelisca Bates and Metapachylus Bates (Coleoptera: Scarabaeidae:

Rutelinae) with comments on classification of the subtribe Rutelina.

Proceedings of the Entomological Society of Washington, 102, 332–349.

Jameson, M.L. & Hawkins, S.J. (2005) Synopsis of the genera of Geniatini

(Coleoptera: Scarabaeidae: Rutelinae) with an annotated catalog of

species. Zootaxa, 874(1), 1–58. Available from: https://doi.org/10.

11646/zootaxa.874.1.1

Jameson, M.L. & Ratcliffe, B.C. (2011) The Neotropical scarab beetle tribe

Anatistini (Coleoptera: Scarabaeidae: Rutelinae). Bulletin of the Uni-

versity of Nebraska State Museum, 26, 1–100.

Jameson, M.L. & Smith, A.B.T. (2002) Revision of the South American

genus Brachysternus Guérin-Méneville (Coleoptera: Scarabaeidae:

Rutelinae: Anoplognathini: Brachysternina). The Coleopterists Bulle-

tin, 56(3), 321–366. Available from: https://doi.org/10.1649/0010-

065X(2002)056[0321:ROTSAG]2.0.CO;2

Janssens, A. (1949) Table synoptique et essai de classification pratique

des Coléoptères Scarabaeidae. Institut Royal Des Sciences Naturelles

de Belgique, 25, 1–30.

Kawada, R. & Buffington, M.L. (2016) A scalable and modular dome illu-

mination system for scientific microphotography on a budget. PLoS

ONE, 11(5), e0153426. Available from: https://doi.org/10.1371/

journal.pone.0153426

Koch, C.L. (1839) Die Arachniden getreu nach der Natur abgebildet und

deschrieben. C. H. Zehschen Buchhandlung,7,1–30.

Krajcik, M. (2007) Checklist of Scarabaeoidea of the world 2. Rutelinae.

Animma.x,4,x1–273.

Krell, F.T. (1996) Die Kopulationsorgane des Maikäfers Melolontha melo-

lontha (Insecta: Coleoptera: Scarabaeidae). Ein Beitrag zur vergle-

ichenden und funktionellen Anatomie der ektodermalen Genitalien

der Coleoptera. Stuttgarter Beiträge zur Naturkunde. Serie A

(Biologie), 537, 1–101.

Kukalov

a-Peck, J. & Lawrence, J.F. (1993) Evolution of the hind wing in

Coleoptera. The Canadian Entomologist, 125(2), 181–258. Available

from: https://doi.org/10.4039/Ent125181-2

Lacordaire, J.T. (1855) Histoire Naturelle des Insectes: Genera des Coléop-

tères, Vol. 3. Paris: Librairie Encyclopédique de Roret, pp. 1–594.

Lacordaire, J.T. (1869) Genera des Coléoptères. Histoire Naturelle des

Insectes, Atlas, Vol. 3. Paris: 318 pp.

Lawrence, J.F. & Britton, E.B. (1994) Australian beetles. Melbourne: Mel-

bourne University Press 192 pp.

Machatschke, J.W. (1965) Coleoptera Lamellicornia. Fam. Scarabaeidae,

subfam. Rutelinae section Orthochilidae. Genera Insectorum, 199C,

1–145.

Machatschke, J.W. (1972) Scarabaeoidea: Melolonthidae, Rutelinae.

Coleopterorum Catalogus Supplementa, 66, 1–361.

McKenna, D.D., Farrell, B.D., Caterino, M.S., Farnum, C.W., Hawks, D.C.,

et al. (2015) Phylogeny and evolutionof Staphyliniformia and Scara-

baeiformia: forest litter as a stepping stone for diversification

ofnonphytophagous beetles. Systematic Entomology, 40(1), 35–60.

Moore, M.R., Cave, R.D. & Branham, M.A. (2018) Synopsis of the cycloce-

phaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). Zoo-

Keys, 745(745), 1–99. Available from: https://doi.org/10.3897/

zookeys.745.23683

Nel, A. & Scholtz, C.H. (1990) Comparative morphology of the mouth-

parts of adult Scarabaeoidea (Coleoptera). Entomology Memoir

Department of Agricultural Development, 80, 1–84.

Ohaus, F. (1900) Bericht über eine entomologische Reise nach Central-

brasilien. Stettiner Entomologische Zeitung, 61, 193–274.

Ohaus, F. (1918) Scarabaeidae: Euchirinae, Phaenomerinae, Rutelinae.

Coleopterum Catalogus, 10, 9–217.

Ohaus, F. (1928) Beitrag zur Kenntnis der Ruteliden (Col. Lamell.).

Deutsche Entomologische Zeitschrift, Berlin, 25, 385–406.

Ohaus, F. (1934) Coleoptera Lamellicornia: fam. Scarabaeidae: subfam

Rutelinae. Genera Insectorum, 199, 1–172.

Ratcliffe, B.C., Jameson, M.L., Figueroa, L., Cave, R.D., Paulsen, M.J.,

Cano, E.B., et al. (2015) Beetles (Coleoptera) of Peru: a survey of the

families. Scarabaeoidea. Journal of the Kansas Entomological Society,

88(2), 186–207. Available from: https://doi.org/10.2317/kent-88-02-

186-207.1

Ratcliffe, B.C., Jameson, M.L. & Zorn, C. (2018) Ganganomala saltini Rat-

cliffe, Jameson, and Zorn, a new genus and species of Anomalini

(Coleoptera: Scarabaeidae: Rutelinae) from Bangladesh and Nepal,

with a revised circumscription of the tribe. The Coleopterists Bulletin,

72(4), 717–735. Available from: https://doi.org/10.1649/0010-065X-

72.4.717

Scholtz, C.H. (2000) Evolution of flightlessness in Scarabaeoidea (Insecta,

Coleoptera). Deutsche Entomologische Zeitschrift, 47(1), 5–28.

Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce

publication-quality point maps. Available from: http://www.

simplemappr.net

Simpson, G.G. (1961) Principles of animal taxonomy. New York: Columbia

University Press.

Smith, A.B.T. (2006) A review of the family-group names for the super-

family Scarabaeoidea (Coleoptera) with corrections to nomencla-

ture and a current classification. Coleopterists Society Monograph,5,

144–204.

Smith, A.B.T. & Evans, A.V. (2005) A supplement to the checklist of the

New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) with

notes on their tribal classification. Zootaxa, 1032(1), 29–60. Avail-

able from: https://doi.org/10.11646/zootaxa.1032.1.2

Smith, A.B.T., Hawks, D.C. & Heraty, J.M. (2006) An overview of the classi-

fication and evolution of the major scarab beetle clades

(Coleoptera: Scarabaeoidea) based on preliminary molecular ana-

lyses. Coleopterists Society Monograph,5,35–46.

Soula, M. (1998) Les Coléoptères du Monde: Rutelini 2. Canterbury: Hillside

Books 116 pp.

Soula, M. (2011) Les Coléoptères du Nouveau Monde: Révision du genre

Bolax (Coleoptera: Scarabaeidae: Rutelinae: Geniatini). Association

Entomologique pour la Connaissance de la Faune Tropicale—AECFT,

85 pp.

Torre-Bueno, J.R. (1985) A glossary of entomology: Smith’s“an explanation

of terms used in entomology”. New York: New York Entomological

Society, p. 323.

Wiley, E.O. & Lieberman, B.S. (2011) Phylogenetics: theory and practice of

phylogenetic systematics, 2nd edition. Hoboken, New Jersey: John

Wiley & Sons 406 pp.

Wiley, E.O. & Mayden, R.L. (2000) The evolutionary species concept. In:

Species concepts and phylogenetic theory: a debate. New York, New

York: Columbia University Press, pp. 70–89.

How to cite this article: Bento, M., Jameson, M.L.,

Grossi, P. & da Fonseca, C.R.V. (2024) Revision of the

Neotropical tribe Alvarengiini Frey, 1975

(Coleoptera: Scarabaeidae). Austral Entomology,

63(1), 12–48. Available from: https://doi.org/10.

1111/aen.12676

48 BENTO ET AL.

All genera of the world: Subfamilies Dynastinae, Rutelinae and Cetoniinae (Animalia: Arthropoda: Insecta: Coleoptera: Scarabaeidae)

Article

Full-text available

May 2024

There are about 1500 genus-group names available in the scarab subfamilies Dynastinae, Rutelinae and Cetoniinae by the end of 2023. Of these, 997 refer to currently accepted genera, 227 in Dynastinae, 261 in Rutelinae and 509 in Cetoniinae. A further >650 genus-group names are given subgeneric rank or placed in synonymy. The chronology of descriptions of currently valid genera shows similar patterns in each of the three subfamilies with many described in the 1840s, a large group in the late 1800s (especially the Cetoniinae) and early 1900s (Dynastinae and Rutelinae) and a steady increase in numbers following World War 2. We predict that discovery of new genera, as well as the potential for further splitting of mega-genera with elevation of subgenera to full generic level, will continue the upward trend, albeit somewhat balanced by combining the numerous monobasic genera.

On the tribal classification of the Nearctic Melolonthinae (Coleoptera: Scarabaeidae), with descriptions of new species of Acoma Casey, 1889

Article

Full-text available

Mar 2020

The subfamily Melolonthinae (Coleoptera: Scarabaeidae) is defined and characterized, and a brief summary of the world melolonthine tribes and their distributions are provided. Nearctic genera previously considered incertae sedis (Acoma Casey, 1889, Chaunocolus Saylor, 1937 and Chnaunanthus Burmeister, 1844, Phobetus LeConte, 1856, and Warwickia Smith & Evans, 2005) are each placed in the proposed new tribes Acomini, Chnaunanthini, Phobetusini, and Warwickiini, respectively. Tribal assignments for all Nearctic melolonthine genera are presented. Acoma chihuahuaensis, A. eusexfoliata, A. nonglabrata, and A. pararobusta are all new species described from Mexico. The only known example of a female Acoma, represented by a specimen of A. knulli Howden, 1958, is figured and characterized. The generic composition of the Nearctic Melolonthini and Rhizotrogini is examined. Madiniella Chalumeau & Gruner, 1976, previously placed in Tanyproctini, is transferred to Rhizotrogini. The subfamilies Oncerinae and Podolasiinae are each removed as tribes from the Melolonthinae and elevated to the subfamily level within Scarabaeidae. The subtribe Triodonina is placed in synonymy with the tribe Rhizotrogini. An updated generic checklist and tribal key of the Nearctic Melolonthinae are provided.

Description of the female of Spodochlamys marahuaca Jameson & Ratcliffe, 2011 (Melolonthidae, Rutelinae, Anatistini) and additional records for the Brazilian species of the genus

Article

Full-text available

Aug 2019

The previously unknown female of Spodochlamys marahuaca Jameson & Ratcliffe, 2011 is described and illustrated for the first time based on two specimens collected in the region of the Juruá River, Amazonas state, Brazil. The specimens constitute a new record for Brazil. The identification key for species of Spodochlamys is updated, as well as the geographic distribution of the other two Brazilian species, S. caesarea Burmeister and S. iheringi Ohaus.

Ganganomala Saltini Ratcliffe, Jameson, and Zorn, a New Genus and Species of Anomalini (Coleoptera: Scarabaeidae: Rutelinae) from Bangladesh and Nepal, with a Revised Circumscription of the Tribe

Article

Full-text available

Dec 2018

Ganganomala saltini Ratcliffe, Jameson, and Zorn, a new genus and new species of Anomalini (subtribe Anomalina), is described fromBangladesh and Nepal. To place the new genus within the context of the tribe and subtribe, we provide a key to the four subtribes of world Anomalini, a list of comparative characters and character states for diagnosis and classification of Old World Anomalina, and comparative diagnostic characters for genera of Old World Anomalina. Circumscription, description, diagnosis, and illustrations for the new genus and species are provided. As a result of this research, the Anomalina includes 50 genera and subgenera, and advances our understanding of global anomaline biodiversity.

Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae)

Article

Full-text available

Mar 2018

The cyclocephaline scarabs (Scarabaeidae: Dynastinae: Cyclocephalini) are a speciose tribe of beetles that include species that are ecologically and economically important as pollinators and pests of agriculture and turf. We provide an overview and synopsis of the 14 genera of Cyclocephalini that includes information on: 1) the taxonomic and nomenclatural history of the group; 2) diagnosis and identification of immature life-stages; 3) economic importance in agroecosystems; 4) natural enemies of these beetles; 5) use as food by humans; 6) the importance of adults as pollination mutualists; 7) fossil cyclocephalines and the evolution of the group; 8) generic-level identification of adults. We provide an expanded identification key to genera of world Cyclocephalini and diagnoses for each genus. Character illustrations and generic-level distribution maps are provided along with discussions on the relationships of the tribe’s genera.

Beitrag zur Kenntnis der Ruteliden. (Col. lamell.)

Article

Apr 2008

F. Ohaus

International code of zoological nomenclature = Code international de nomenclature zoologique

Book

Jan 1999

W. D. L. Ride

Principles of Animal Taxonomy

Article

Jun 1962

Principles of Animal Taxonomy

Book

Dec 1961

George Gaylord Simpson

Australian Beetles

Book

Jan 1994

There are possibly 30 000 species of beetle in Australia - no one knows for sure. But professional entomologists, students and keen amateurs alike will find what is known about this striking, diverse and intriguing order of insects in this comprehensive guide by two of the world's recognised experts. Australian Beetles is a detailed account of the beetle fauna of Australia; it includes general discussions of adult and larval anatomy, general biology and economic significance, keys to families based on larvae and adults, and a brief coverage of each of the 117 families occurring on the continent.

A glossary of surface sculpturing.

Article

Jan 1979

Harris, R. A.

Revision of the Neotropical tribe Alvarengiini Frey, 1975 (Coleoptera: Scarabaeidae)

Abstract

Recommended publications

On the tribal classification of the Nearctic Melolonthinae (Coleoptera: Scarabaeidae), with descript...

Liomenochilus ongi (Coleoptera: Scarabaeidae: Melolonthinae: Systellopini), a new genus and new spec...

Phylogenetic relationships within Diplotaxini Kirby (Coleoptera: Melolonthidae: Melolonthinae) with...

Taxonomic and Nomenclatural Changes in the Pelidnotine Scarabs (Coleoptera: Scarabaeidae: Rutelinae:...