Beetles (Coleoptera) of Peru A Survey of The Families. Part I.

Beetles (Coleoptera) of Peru: A Survey of the Families. Part I.
Overview
Author(s): Caroline S. Chaboo
Source: Journal of the Kansas Entomological Society, 88(2):135-139.
Published By: Kansas Entomological Society
DOI: http://dx.doi.org/10.2317/0022-8567-88.2.135
URL: http://www.bioone.org/doi/full/10.2317/0022-8567-88.2.135
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JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
88(2), 2015, pp. 135139
Beetles (Coleoptera) of Peru: A Survey of the Families. Part I. Overview

CAROLINE S. CHABOO
Division of Entomology, Biodiversity Institute, 1501 Crestline Drive, Suite 140,
University of Kansas, Lawrence, Kansas, USA, 66045. E-mail: cschaboo@ku.edu
ABSTRACT: The beetle fauna of Peru has not been enumerated since Blackwelder (19441957).
This contribution is the first modern synopsis of beetle families, based on ongoing fieldwork
and a literature review. Ninety nine families of beetles are now documented from Peru and
nine families are reported in Peru for the first timeBiphyllidae, Bothrideridae, Eucinetidae,
Monotomidae, Rhipiceridae, Scirtidae, Scraptiidae, Silvanidae, and Throscidae. A species list
for each family, updated higher taxonomy, and relevant literature citations are presented.
Most families are so poorly known in the tropics and so poorly represented in museum
collections, that their rarity and/or conservation status cannot be determined. Altogether the
project represents the first modern catalogue of the entire beetle fauna of a hyperdiverse
Neotropical country.
KEY WORDS: Checklist, biodiversity, new records, distribution, insects
RESUMEN: La fauna de escarabajos de Peru no se ha enumerado desde Blackwelder (1944

1957). Esta contribucion es la primera sinopsis moderna de las familias de escarabajos, basada
en el trabajo de campo en curso y una revision de la literatura. Noventa y nueve familias de
escarabajos de Peru estan documentadas,y ocho familias se registran en Peru por primera
vezBiphyllidae, Bothrideridae, Eucinetidae, Monotomidae, Rhipiceridae, Scirtidae, Scrap-
tiidae, Silvanidae, y Throscidae. Se presenta una lista de especies para cada familia, taxonoma
superior actualizada, y citas bibliograficas pertinente. La mayora de las familias son tan poco
conocidas en los tropicos y tan mal representados en las colecciones del museo, que su rareza
y / o estado de conservacion no se puede determinar. En total, el proyecto representa el primer
catalogo moderno de toda la fauna de escarabajos de un pas Neotropical hiperdiverso.
PALABRAS CLAVE: Lista de verificacion, biodiversidad, nuevos registros, distribucion, insectos
Peru is known for its exceptionally high species richness, endemism, and habitat
complexity, yet little is known about the beetle fauna of the country (Larsen et al.,
2011). The beetle fauna of Peru has not been enumerated since Blackwelder (1944
1957). Erichson (1847) and the supplements by Kirsch (1873, 1876) were historically
the only treatments focused on the Peruvian fauna as a whole.
This series, Beetles of Peru, is an attempt to present the first modern synopsis of
beetle families, based on a literature review, extraction of institutional records, and
our fieldwork in Peru since 2007. Checklists are presented for each of 99 families we
now know to occur in Peru.
Why Checklists?
Lists of species in a place, from a park to the entire globe, are a critical step in opening
sites for biodiversity study, management, and conservation. When taxonomic lists are
done by a trained taxonomist, we can validate names after reviewing the scattered
primary literature to trace all taxonomic changes related to that species. This can be
a challenge for diverse groups like insects where both the number of species and the
scattering of literature pose special issues. Species lists can open research by providing
presence/absence information on species, their validated names with accurate spelling,
Accepted 31 April 2014; Revised August 20 2015

E 2015 Kansas Entomological Society
136 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
original authorship, and year. Such lists provide an entree to the literature (for original
authorship) and are essential tools for biogeography and conservation. When
combined with museum records of voucher specimens, checklists provide more
complete information about species distributions and help identify sites of particular
interestfor their species richness, special communities, rarity, gaps of information
and spur species inventories. Checklists can also serve as reference points for
determining shifting species distributions, due to anthropogenic, landscape, and
climate change impacts. They are crucial to monitoring native, alien, and invasive
species. When combined with photographs of species and/or maps, checklists become
valuable educational tools, for the lay public and scientists.
A natural but recent development in the development of accurate species lists has
been the growth of online databases and the global-networking of such databases.
The critical and growing need for accurate checklists has led to the development of
a guide to best practices in assembling national checklists (Hamer et al., 2012),
supported by the Global Biodiversity Information Facility (GBIF), to aid efficiency
and consistency in standards of such works. Today, there is even a journal,
CheckList (CheckList.com), devoted to such species lists. There are interesting
discussions about the difficulty and challenges of validating scientific names (Vaidya,
2013) and even some tools that might help cross-reference names (see Page, 2012).
The Catalog of Life (http://www.catalogueoflife.org/) seeks to compile and reconcile
names for all species names available, and provide information on their distributions
and relationships.
This Project
Herein we report a total of 99 families from Peru. We also report ,10,000 species
names, with a species list for each family, updated higher taxonomy, and relevant
literature citations. Perus beetle fauna is exceedingly rich at the genus and species
levels. Nine families are reported in Peru for the first timeBiphyllidae,
Bothrideridae, Eucinetidae, Monotomidae, Rhipiceridae, Scirtidae, Scraptiidae,
Silvanidae, and Throscidae. Most of these families are so poorly known in the
tropics, and so poorly represented in museum collections, that their rarity and/or
conservation status cannot be determined. Based on occurrence in adjoining
countries, more families are anticipated to occur in Peru.
This series represents the first modern catalogue of the entire beetle fauna of
a hyperdiverse Neotropical country. It is preliminary in the sense that we can now
target fieldwork and study of institutional collections to assemble an accurate list of
species occurring in Peru. The participating family authors welcome any notification
of errors, so we can continue to improve knowledge of Perus fauna. From the
present base, we can determine localities, distributions, habitat, and host records to
date. Ultimately, this work will open the way for revisions, new species descriptions,
identification keys, new biological research, and the conservation status of species.
A great deal of research is warranted. Beyond this effort, I hope that other
entomologists will take interest in the Peruvian fauna. Ideally, Peruvian students will
be recruited and trained for this effort.
Materials and Methods

Data for this series was compiled from three primary sources: literature survey
(starting with Blackwelder, 19441957 and Titschack, 19511954), institutional
VOLUME 88, ISSUE 2 137
collections, and specimens collected in my field work in Peru. The compilation by

Titschack (Volume 2: 1951, Volume 3: 1952, Volume 4: 1954) partially treated the
beetle fauna of Peru. Volume 2 is a compilation of articles published in 1941, Volume
3 is a compilation of articles published in 1942, while Volume 4 was to be published
in 1943 but, because of the bombing of Hamburg during that year, publication of its
articles was delayed until 1954.
How to Use This Series

Families are presented in this and upcoming issues of the Journal of the Kansas
Entomological Society; they do not follow a strict phylogenetic sequence as they are
published after a review process. The classification followed here follows the higher-
level classification of Bouchard et al. (2011). Group taxonomic names follow the
latest accepted taxonomies within the particular family treatment. We provide
citations of fundamental sources for Peru species data; citations of original species
description sources will be provided in future revisions of the family treatments, as
new species are described and more entomology collections are examined.
New Records
Species that are recorded for Peru for the first time are indicated within each family
section by a * or as new record, with their locality data and depository. Some new
records are based my field surveys or on information extracted from specimen label
data in several institutional collections: Brigham Young University Collection, USA
(BYU); California Academy of Sciences, USA (CAS); Collection of Histeridae of
Nicolas Degallier, France (CHND); Louisiana State Arthropod Museum, USA,
(LSAM); Lund Museum of Zoology - Insect collections, Sweden (MZLU); Museo de
Entomologa de la Universidad Nacional de Tumbes, Peru; Museum of Comparative
Zoology, USA (MCZ); Museum de Universidade de San Marcos, Peru (MUSM);
Snow Entomology Museum Collection, USA (SEMC); Texas A & M University, USA
(TAMU); U.S. National Museum, USA (USNM); Palacky University, Olomouc,
Czech Republic (UPOL); Museo de Entomologa Klaus Raven Buller, Universidad
Nacional Agraria La Molina, Peru (UNAD); Universidad Nacional Santiago Antunez
de Mayolo, Peru; and University of Georgia Collection of Arthropods, USA (UGCA).
In each family section, these institutions are indicated by their standardized acronyms
following Evenhuis (2007; http://hbs.bishopmuseum.org/codens/codens-r-us.html).
Fieldwork
I have been conducting fieldwork in Peru since 1997, in the Madre de Dios region,
southern Peru. My field sites are at three field stations managed by the Amazon
Conservation Association (http://www.amazonconservation.org/). Since 2011, I have
been using a standardized protocol for insect study which includes canopy and
terrestrial Malaise traps, flight intercept traps, colored pan traps, UV/MV light traps,
pitfall traps, and beating/sweeping of vegetation. The by-catch includes many other
specimens that are being sent to an expanding network of researchers for further study.
New Species
No new species are described within these family presentations. My collections
include many new genera and new species (e.g., Ptiliidae: more than 50 new species,
M. Darby, pers. comm.). Within each section, putative new genera and new species
are indicated as sp. 1, sp. 2, etc. (e.g., Buprestidae) or sp. A, sp. B, etc. (e.g.,
Carabidae). Please contact me for details and specimen loans.
Acknowledgements
Funding for my leaf beetle research in Peru began in 2008 with a grant from the
Amazon Conservation Association (PI: CS Chaboo). A NSF-PASI fellowship
supported a visit to Tambopata (PI: J. Vivanco). The Beetles of Peru project is
supported by NSF-EPSCoR #66928 (PI: CS Chaboo) and by the University of
Kansas Department of Ecology and Evolutionary Biology-General Research Fund
(PI: CS Chaboo). Fieldwork in Peru is conducted with Peru research permits No.
506-2011-AG-DGFFS-DGEFFS and No. 0159-2010-AG-DGFFS-DGEFFS to CS
Chaboo. Grants from the KU Caribbean and Latin American Studies program and
the Ernst Mayr Travel Fund, Harvard University, funded collections study and
specimen identifications in Manchester, UK and Berlin, Germany. I thank the
Amazon Conservation Association and staff over the yearsNigel and Renate
Pitman, Alessandro Catenazzi, Gabriela Nunez, Robinson Palomino, Adrian
Tejedor, and Andrew Wilcoxfor their support in facilitating my research and
annual field courses. I thank Jorge Vivanco and Tiffany Weir for their support with
a PASI Fellowship. I am deeply grateful to former collaborators in my lab, Daniel
Bennett and Matthew Gimmel, without whom both the fieldwork and the beetle
synopsis would be impossible. I am also grateful to many KU undergraduate
students who participated in my annual fieldwork since 2010. Other fieldwork
participants made this project enjoyableCheryl Barr, Pedro Cedeno, Maria Jose
Endarra, R. Wills Flowers, Timo Forster, Steve Goddard, Malena Vilchez, Ernesto
Razuri, and Diana Silva. I thank the many collaborating authors who generously
participated in this project so far and continue to examine specimens collected to
identify and describe the many new species collected. I also thank the many reviewers
whose time and expertise improved each family contribution; some became co-
authors in the process. I thank the JKES Editorial team, Roy Beckemeyer, Alan
Burke, Larry Kipp, and Ai-Ping Liang for overseeing the review process. I thank
Sarah Carbonel, Ana Huamantinco, Luis Figueroa, and Gerardo Lamas at MUSM
for their support. I am grateful to Diana Silva, the Museum of the University of San
Marcos, Lima, Peru, and to Cheryl Barr, Wills Flowers, Fernando Merino, and Bill
Shepard for their unwavering support and help.
Literature Cited
Blackwelder, R. E. 19441957. Checklist of the coleopterous insects of Mexico, Central America, The
West Indies, and South America. Part 4. U.S. National Museum, Bulletin 185. Smithsonian
Institution, Washington, D.C. 763 pp.
Bouchard, P., Y. Bousquet, A. E. Davis, M. A. Alonso-Zarazaga, J. F. Lawrence, C. H. C. Lyall, A. F.
Newton, C. A. M. Reid, M. Schmitt, S. A. Slipinski, and A. B. T. Smith. 2011. Family-group names
in Coleoptera (Insecta). ZooKeys 88:1972.
The Catalog of Life. http://www.catalogueoflife.org/ [accessed July 7 2015].
Erichson, W. F. 1847. Conspectus Coleopterorum, quae in Republica Peruana observata sunt. Archiv fur
Naturgeschichte 1847:67185.
Evenhuis, N. L. 2007. The insect and spider collections of the world website, http://hbs.bishopmuseum.
org/codens/codens-r-us.html [Accessed July 31 2015].
Gentry, A. H. 1977. Endangered plant species and habitats of Ecuador and Amazonian Peru. In Prance,
G. T., and T. S. Elias (eds.), Extinction Is Forever. New York Botanical Garden, N.Y.
Hamer, M., J. Victor, and Smith, G. F. 2012. Best Practice Guide for Compiling, Maintaining and
Disseminating National Species Checklists, version 1.0, released in October 2012. Copenhagen:
Global Biodiversity Information Facility, 40 pp. ISBN: 87-92020-48-8, Accessible at http://www.
gbif.org/orc/?doc_id54752.
Kirsch, T. 1873. Neue sudamerikanische Kafer. Deutsche entomologische Zeitschrift 27:187213.
Kirsch, T. 1876. Beitrage zur Kenntnis der Peruanischen Kaferfauna auf Dr. Abendroths Sammlunggen
basirt. Deutsche entomologische Zeitschrift 20:81133.
Larsen, T. H., F. Escobar, and I. Armbrecht. 2011. Insects of the Tropical Andes: Diversity patterns,
processes and global change. In Herzog, S. K., R. Martnez, and P. M. Jrgensen, H. Tiessen (eds.).
Climate Change and Biodiversity in the Tropical Andes 17:228244. Sao Jose dos Campos, Brazil
and Paris, France: Inter-American Institute of Global Change Research (IAI) and Scientific
Committee on Problems of the Environment (SCOPE), xii+348 pp.
Manley, G. V. 1986. Mastogenius Sol. (Coleoptera: Buprestidae) of Ecuador and Peru west of the Andean
Mountains with descriptions of five new species. The Coleopterists Bulletin 40(3):223231.
Page, R. D. M. 2012. Using Google Refine and taxonomic databases (EOL, NCBI, uBio, WORMS) to
clean messy data. iPhylo Blog [accessed July 22 2014].
Titschack, E. 19511954. Beitrage zur Fauna Perus: nach der Ausbeute der Hamburger Sudperu-
Expedition 1936, anderer Sammlungen, wie auch auf Grund von Literaturangaben. Fischer, Jena.
Vaidya, G. 2013. Validating scientific names with the GBIF Checklist Bank. VertNet Blog [accessed July
22 2014].
Beetles of Peru: Biogeography
Author(s): Caroline S. Chaboo and Alessandro Catenazzi
DOI: http://dx.doi.org/10.2317/kent-88-02-140-143.1
URL: http://www.bioone.org/doi/full/10.2317/kent-88-02-140-143.1

terms_of_use.
critical research.
88(2), 2015, pp. 140143
Beetles of Peru: Biogeography

CAROLINE S. CHABOO1,3 AND ALESSANDRO CATENAZZI2
Peru is the third largest country in South America, after Brazil and Argentina.
Administratively, Peru is divided into 24 Regiones (formerly Departamentos) each
with a regional government and capital. The two largest cities are Lima and
Arequipa. Below is a broad outline of Perus geography, vegetation, and habitats.
Perus 1.285 sq. km. lie entirely within the tropics from the equator to ,18uS
latitude. This latitudinal position of Peru, along with changes in the strength of the
coastal oceanic currents, fosters a zonation of climate from north to south.
Perus topography is complex, due to the age and orogeny of the Andes
Mountains (Antonelli et al., 2009; Hoorn et al., 2010) and the evolution of
Amazonia (Hoorn and Wesselingh, 2010). The Peruvian Andes are relatively young
mountains dating to the Cretaceous, with periodic uplifting events over the past 50
ma years (Hoorn et al. 2010) above the subduction zone where the Nazca plate slides
beneath the South American plate. The Peruvian landscape has three broad
geographic zones running longitudinally in a north to south axiscoastal desert (or
Costa), central Andes (or Sierras), and eastern wet forest zone (or Selva) with the
headwaters of the Amazon River and the Amazon basin. These broad zones roughly
correspond to Perus climate regions. Brack (1986) recognized nine terrestrial
ecoregions. Additionally, 83 of 104 life zones defined in the Holdridge system are
found in Peru (Oficina Nacional de Evaluacion de Recursos Naturales, 1976). See
Morrone (2001, 2006) for recent classification of life zones.
The reader is encouraged to access maps and descriptions in other works that treat
Perus biogeography and biodiversity (e.g., Gentry, 1990; Kalliola et al., 1993).
Perus vertebrates are better studied animals (e.g., Amphibians: Duellman and Lehr,
2009; Catenazzi and von May, 2014; Birds: Fjeldsa and Krabbe, 1990; Stotz et al.,
1996; Schulenberg et al., 2010); some texts are site-specific, e.g. Reserva Cusco
Amazonico (Duellman, 2005). Reports of rapid inventories conducted by
Conservation International and the Field Museum (Schulenberg and Awbrey,
1997, 2001; Pitman et al., 2004, 2011, 2012; Gilmore et al., 2010) provide information
on geology, vegetation and vertebrate faunas in montane and lowland Amazon
rainforests. The geographic and ecological scales of those sources concern generally
larger-bodied vertebrates, but entomologists will find them valuable resources as we
develop our own arthropod-relevant distinctions in ecoregions and habitats.
Early arthropod-focused inventories provide some closer descriptions of particular
sites but these habitats are not altered today; for example, Manu (Wilson and
Sandoval, 1996) and Tambopata (Erwin, 1985). More recent insect surveys give
1
Division of Entomology, Biodiversity Institute and Department of Ecology and Evolutionary Biology,
1501 Crestline Drive, Suite 140, University of Kansas, Lawrence, Kansas 66045, U.S.A.; E-mail:
cschaboo@ku.edu
2
Department of Zoology, Southern Illinois University Carbondale, Carbondale Illinois 62901, E-mail:
acatenazzi@siu.edu
3
Corresponding author
Accepted 1 August 2015; Revised 1 September 2015
insights into certain habitats: dung beetles in Megantoni National Park (Larsen,
2005); beetle communities of Perus southwestern bamboo forests (Jacobs, 2009);
and carabid communities in Andean forests (Maveety et al., 2014).
Climate: Elevation, latitude, rain shadow and other local effects contribute to
dramatic differences in temperature and precipitation from north to south and up
the Andean slopes. Year-round average temperatures may vary little in Peru, but the
diurnal temperature variation can be great. One particularity of the Peruvian climate
is the phenomena known as El Nino (the child because of its general coincidence
with Christmas) and La Nina. These phenomena are associated with the Southern
Oscillation, changes in surface air pressure between the waters of the tropical western
and eastern Pacific Ocean. The periodic oscillation produces a warming of coastal
waters in Peru during El Nino, with warm water replacing cold water in the
Humboldt current, heavy rains often with floods along the coast, and local drought
in the south. La Nina is the other extreme of the oscillation, and usually manifests in
lower than average temperatures in Peru.
Coast: This narrow strip below 1000 m elevation and ,3000 km long (,11% of
Perus territory) is mostly desert because of three factors: the cold Humboldt current
transfers little moisture to on-shore winds, Pacific on-shore winds yield little rainfall,
and the Andes blocks rain-bearing Amazon winds. There is basically one type of arid
climate north of Piura (with seasonal rains) and the hyper-arid climate typical of
Lima that extends all the way into the Atacama desert of Chile. This coast has three
broad ecological areas: tropical forest, equatorial dry forest, and desert. The coast is
widest in the north, where it borders Ecuador, and the vegetation is equatorial forest
or savanna with particular tree species like sapote (Capparis scabrida) and mesquite
(Prosopis pallida). Peru even has mangrove swamps in the Tumbes area and at the
estuary of the Piura River (Echevarra and Sarabia, 1993). The coast to the south is
desert with sand dunes interrupted with coastal oases of scrub and dry forest around
seasonal rivers coming off the Andes. Major rivers are Chicama, Santa, Rimac (at
Lima), Canete, Ro Grande, and Majes-Colca. Heavy coastal fogs foster a peculiar
vegetation type called lomas with a mix of grasses and herbs.
Andes (Sierras) mountains: These form the jagged spine of Peru, reaching up to
6000 m; the highest peak, Mt. Huascaran, is 6768 m. Eight major mountain ranges
are recognized in Perus Sierras: Cordillera Occidental, Cordillera Central,
Cordillera Oriental, Cordillera de Huancabamba, Cordillera de Colan, Cordillera
del Condor, Cordillera Azul, and Cordillera de Vilcanota (Duellman and Lehr,
2009). Peru has more than 20 extinct and active volcanoes (Siebert and Simkin, 2002)
due to the countrys location on a zone of active plate subduction. Deep inter-
Andean valleys and canyons, some with walls over 1000 m high, form major
biogeographic barriers. Canon del Colca is one of the deepest in the world. Snow
falls in the highest Sierras and Peru has several glaciers. The southern Andes is more
flattened and is referred to as altiplano. Amazon winds are blocked by the Andes and
deposit their rains on Perus eastern slopes, making this region some of the wettest
and most biodiverse in the world. The cloud forests here are probably the longest
intact continuous belt of cloud forests in the world.
Amazon basin: The complex terrain of eastern Peru underlies an extraordinary
drainage network with many rivers flowing east and north to connect eventually with
the Amazon, or southwards into the Madre de Dios (a tributary of the Madeira
river). Major rivers are the Amazon, Maranon, Ucayali, Putumayo, Napo,
Huallaga, Yavar in the Amazon/Solimoes basin, and Madre de Dios in the Madeira
basin. These large rivers flow swiftly down the steep slopes, around ridges and hills at
the base of the Andes, and form large floodplains with numerous oxbow lakes and
shifting channels. The major lowland relief is around the Sierra del Divisor, central
Peru near Brazil. Perus lowland zone (selva baja) is widest in the north (Loreto
Province) with the large tributaries of the Amazon River.
Perus eastern slopes are among the wettest in the world and are cloaked in
vegetation that changes from alpine vegetation to grassland to cloud forests around
30003500 m (tree-line). Mid-elevation montane forests merge downslope into
lowland tropical rainforests, palms, bamboos, and savannas.
Acknowledgements
We acknowledge NSF-EPSCoR # 66928 (PI: CS Chaboo) for supporting the
Beetles of Peru project and the University of Kansas Department of Ecology and
Evolutionary Biology-General Research Fund (PI: CS Chaboo). We also thank
Wills Flowers and Bill Shepard for reviewing the manuscript and suggesting
additional citations.
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Explicativa. Oficina Nacional de Evaluacion de Recursos Naturales, Peru.
Pitman, N., E. Ruelas Inzunza, D. Alvira, C. Vriesendorp, D. K. Moskovits, A. del Campo, T. Watcher,
D. F. Stotz, S. Noningo Sesen, E. Tuesta Cerron, and R. C. Smith. 2012. Peru: Cerros de
Kampankis. Rapid Biological Inventories: 24. The Field Museum, Chicago, Illinois.
Pitman, N., R. C. Smith, C. Vriesendorp, and D. Moskovits. 2004. Peru: Ampiyacu, Apayacu, Yaguas,
Medio Putumayo. Rapid Biological Inventories: 12. The Field Museum, Chicago, Illinois.
Pitman, N., C. Vriesendorp, D. K. Moskovits, R. von May, D. Alvira, T. Watcher, D. F. Stotz, and A. del
Campo. 2011. Peru: Yaguas-Cothue. Rapid Biological Inventories: 23. The Field Museum,
Chicago, Illinois.
Schulenberg, T. S., and K. Awbrey (eds). 1997. The Cordillera del Condor Region of Ecuador and Peru: A
Biological Assessment. RAP Working Papers 7:231 pp. Conservation International, Washington
DC.
Schulenberg, T. S., and K. Awbrey (eds). 2001. Biological and Social Assessments of the Cordillera de
Vilcabamba, Peru, 2001, RAP Working Papers 12:396 pp. Conservation International, Washington
DC.
Schulenberg, T. S., D. F. Stotz, D. F. Lane, J. P. ONeill, and T. A. Parker III. 2010. Birds of Peru.
Revised and updated edition. Princeton University Press, Princeton and Oxford, 664 pp.
Siebert, L., T. Simkin. 2002. Volcanoes of the World: An illustrated catalog of Holocene volcanoes and
their eruptions. Smithsonian Institution, Global Volcanism Program Digital Information Series,
GVP-3, http://www.volcano.si.edu [Accessed July 20 2015].
Stotz, D. F., J. S. Fitzpatrick, T. A. Parker III, and D. K. Moskovits. 1996. Neotropical Birds: Ecology
and Conservation. Chicago: University of Chicago Press.
Wilson, D. E., and A. Sandoval. 1996. Manu: The Biodiversity of Southeastern Peru. Lima: Editorial
Horizonte, 579 pp.
Beetles (Coleoptera) of Peru: A Survey of the Families.
Hydroscaphidae and Sphaeriusidae (Myxophaga)
Author(s): Caroline S. ChabooWilliam D. Shepard and

terms_of_use.
critical research.
88(2), 2015, pp. 144145
Beetles (Coleoptera) of Peru: A Survey of the Families. Hydroscaphidae

and Sphaeriusidae (Myxophaga)
CAROLINE S. CHABOO1,2 AND WILLIAM D. SHEPARD3
The beetle suborder Myxophaga comprises four families, two of which occur in
Peru. The remaining two families have been collected close to Peru (Lepiceridae from
Ecuador; Torridincolidae from Brazil and Paraguay) and may yet be found in Peru
with more intensive collecting in the correct microhabitat. Myxophagan adults are
minute and may have a ventral plastron; all known larvae have spiracular gills.
Hydroscaphidae Leconte, 1874

Diversity in Peru: 1 genus, 1 species.
Recognition: Adults are easily recognized by the tiny, teardrop-shaped body (12
mm long), truncate elytra exposing several abdominal tergites, and clubbed
antennae.
Habitat: Both adults and larvae feed on and live in algae, in shallow margins of
streams. Many are associated with hot springs.
Notes: Within the Neotropics, the genus Hydroscapha was mentioned as ranging
from Mexico to Panama and Peru by Reichardt (1973), who was citing an earlier
paper by Hinton. No species name was associated with these distributional records
and presumably they represent undescribed species. Blackwelder (19441957) did not
record this family for Peru. Adults and larvae can be easily collected by removing
rocks and wood from the water and picking up individuals with fine forceps.
Checklist:
Hydroscapha undetermined species (Reichardt, 1973)
Sphaeriusidae Erichson, 1845

Recognition: Adults are easily recognized by their tiny (about 1mm long),
glabrous, round bodies, in combination with only three abdominal ventrites; the
second ventrite is much shorter than the others. Larvae are onisciform in shape,
wide-bodied, with a hypognathous head and having all thoracic segments and
abdominal segments 18 dorsally with an apical fringe of flattened setae (Britton,
1966).
1
Division of Entomology, Biodiversity Institute, University of Kansas, 1501 Crestline Drive, Suite 140,
Lawrence, Kansas 66045.
2
Essig Museum of Entomology, University of California, 1101 Valley Life Sciences Bldg., #4780,
Berkeley, California 94720.
3
Corresponding author. E-mail: cschaboo@ku.edu
Accepted 29 April 2014; Revised 09 April 2015
Habitat: Adult and larval sphaeriusids are usually semiaquatic, inhabiting sandy
or gravelly margins of rivers and streams. Some Asian species are found in leaf litter
and moss (Lobl, 1995).
Notes: The family Sphaeriusidae contains a single genus, Sphaerius, which is
cosmopolitan but spotty in distribution. It was recorded from Peru by Beutel and
Arce-Perez (2005), who noted that the South American specimens are unstudied.
Blackwelder (19441957) did not record the family from Peru. These beetles are
collected commonly when light-trapping near bodies of water. They can also be
collected by sieving the beach sands of streams and lakes.
Checklist:
Sphaerius undetermined species (Beutel & Arce-Perez, 2005)
Acknowledgements
We acknowledge NSF-EPSCoR #66928 (PI: CS Chaboo) for supporting the
Evolutionary Biology-General Research Fund (PI: CS Chaboo).
Literature Cited
Beutel, R. G., and R. Arce-Perez. 2005. Sphaeriusidae Erichson, 1845 (Jach 1999) (5Microsporidae). In
R. G. Beutel and R. A. B. Leschen (eds.). Handbuch der Zoologie. Coleoptera, Vol.1, Morphology
and Systematics, pp. 4849. Berlin/New York: Walter de Gruyter. xi + 567 p.
Blackwelder, R. E. 19441957. Checklist of the coleopterous insects of Mexico, Central America, the West
Indies, and South America. Parts 1 through 6. United States National Museum Bulletin. 185:ixii,
11492.
Britton, E. B. 1966. On the larva of Sphaerius and systematic position of the Sphaeriidae (Coleoptera).
Australian Journal of Zoology. 14:11931198.
Lobl, I. 1995. New species of terrestrial Microsporus from the Himalaya (Coleoptera: Microsporidae).
Entomologische Blatter. 91:129138.
Reichardt, H. 1973. A critical study of the suborder Myxophaga, with a taxonomic revision of the
Brazilian Torridincolidae and Hydroscaphidae (Coleoptera). Arquivos de Zoologia 24:73162.
Dytiscidae, Gyrinidae, Haliplidae, and Noteridae (Suborder
Adephaga)
Author(s): Caroline S. Chaboo and William D. Shepard

terms_of_use.
critical research.
88(2), 2015, pp. 146150
Beetles (Coleoptera) of Peru: A Survey of the Families. Dytiscidae,

Gyrinidae, Haliplidae, and Noteridae (Suborder Adephaga)
CAROLINE S. CHABOO1,3 AND WILLIAM D. SHEPARD2
The beetle suborder Adephaga is the second largest in Coleoptera, comprising .40,000
species. An overview of current phylogenetic hypotheses can be accessed online at
Maddison (1995). Adults all exhibit a visible notopleural suture on the prothorax and
six abdominal sterna with the first three fused and divided by the metacoxa. Larvae the
labrum fused and lack mandibular molae.
Carabidae, the largest adephagan family, is treated separately (Erwin et al. in
press.). Here we treat the four other families documented in Peru.
Gyrinidae Latreille, 1802

Diversity in Peru: 3 genera, 19 species.
Recognition: Gyrinidae is a distinctive family of beetles, instantly recognized by
the horizontally divided eyes, very short antennae, and short, paddle-like middle and
hind legs.
Habitat: Adults inhabit lentic habitats and still-water areas of lotic habitats. They
characteristically swim in gyrating motions on the air-water interface, and feed on
insects and spiders swimming or trapped in the surface film.
Notes: Blackwelder (19441957) recorded three genera and 10 species from Peru.
This has been supplemented by Ochs (1953, 1954, 1955), Guignot (1958), Spangler
(1966), and Brinck (1977). All Peruvian species belong to the subfamily Gyrininae.
Checklist:
Andogyrus buculus Brinck, 1977 Gyretes sericeus Laboulbene, 1853
Andogyrus colombicus (Regimbart, 1890) Gyretes sexualis Regimbart, 1883
Andogyrus ohausi Ochs, 1954 Gyretes suturalis Regimbart, 1884
Andogyrus peruvianus Regimbart, 1907 Gyrinus (Neogyrinus) gibbus Aube, 1838
Andogyrus viscus Brinck, 1977 Gyrinus (Oreogyrinus) baeri Regimbart, 1907
Andogyrus zimmermanni Ochs, 1954 Gyrinus (Oreogyrinus) chiliensis Aube, 1838
Gyretes inflatus Regimbart, 1891 Gyrinus (Oreogyrinus) nigellus Ochs, 1954
Gyretes nitidulus Laboulbene, 1853 Gyrinus (Oreogyrinus) peruvianus Regimbart, 1907
Gyretes peruvianus Ochs, 1953 Gyrinus (Oreogyrinus) punctipennis Regimbart,
Gyretes scaphidiformis Regimbart, 1882 1907
1
Division of Entomology, Biodiversity Institute, 1501 Crestline Drive, Suite 140, University of Kansas,
Lawrence, Kansas 66045
2
Essig Museum of Entomology, 1101 Valley Life Sciences Bldg., #4780, University of California,
Berkeley, California 94720
3
Corresponding author. e-mail: cschaboo@ku.edu
Accepted 13 May 2014; Revised 4 August 2015
Haliplidae Aube, 1836

Recognition: Members of Haliplidae may be recognized as such by their broadly
oval body shape, threadlike antennae, legs with long swimming hairs, and
(especially) large metacoxal plates under which most of the abdominal ventrites
(and base of hind femora) are concealed when in repose.
Habitat: Most Haliplidae are found in weedy, slow-moving, permanent water
bodies. Larvae feed primarily on algae, while adults are omnivorous.
Notes: The following checklist of Peruvian Haliplidae was derived from Vondel
and Spangler (2008). Blackwelder (19441957) recorded one species of Haliplidae
from Peru.
Checklist:
Haliplus heppneri Van Vondel and Spangler, 2008 Haliplus peruanus Zimmermann, 1924
Haliplus ornatipennis Zimmermann, 1921
Noteridae C. G. Thomson, 1860

Recognition: Noterids are very similar to the more diverse and abundant family
Dytiscidae, with which they were formerly classified. Members of Noteridae may be
distinguished by having the scutellum concealed (also occurring in some Dytiscidae),
and the presence of the noterid platforma contiguous, abrupt, flat area on the
ventral surface extending from the prosternal process to the hind coxal plates.
Habitat: Noterids are aquatic and carnivorous as both adults and larvae, and
generally occur in still water with aquatic vascular plants. They predominantly feed
on insect eggs and larvae.
Notes: Noteridae was not recorded from Peru in Blackwelder (19441957). The list
below is derived from Nilsson (2011); all belong to Noterinae: Noterini according to
Miller (2009). More species are anticipated to be found in Peru when more collecting is
done for them, especially in Amazonian Peru. Adults are often abundant at light traps.
Checklist:
Hydrocanthus (Hydrocanthus) debilis Sharp, 1882 Suphis minutus Regimbart, 1903
Hydrocanthus (Hydrocanthus) socius R.F. Sahl- Suphisellus grammicus (Sharp, 1882)
berg, 1844
Dytiscidae Leach, 1815

Diversity in Peru: 8 subfamilies, 14 tribes, 23 genera, 73 species.
Recognition: Recognized by the streamlined, watermelon-seed habitus, thread-
like antennae, and presence of long swimming hairs on the modified middle and hind
legs. Noteridae are very similar; see account of that family for tips on separation.
Habitat: Members of Dytiscidae occur in diverse aquatic habitats, from temporary
to permanent, oxygenated and fast-flowing to stagnant and slow-moving, sub-
terranean or surface. Most adults are expert swimmers and are carnivorous (as the
larvae) or scavengers.
Notes: The following checklist of Peruvian Dytiscidae was derived from
Blackwelder (19441957), Gschwendtner in Titschack (1954), Mouchamps (1957),
Spangler (1966), Young (1981, 1990a, b), Tremouilles (1984), Miller (2001, 2002,
2005a, b, c). Blackwelder (19441957) recorded nine species of Dytiscidae from Peru.
Tremouilles (1995) illustrated keys to adults and larvae (in Spanish) for Argentina
may be helpful in identifying Peruvian species.
Checklist:
Agabinae: Agabini
Agametrus monticola (Guignot, 1958) Agametrus rotundatus Brinck, 1948
Agametrus peruvianus (Laporte, 1835) Leuronectes andinus (Guignot, 1958)
Colymbetinae: Colymbetini
Rhantus (Rhantus) advena Sharp, 1882 Rhantus (Rhantus) discicollis (Aube, 1838)
Rhantus (Rhantus) blancasi Guignot, 1955 Rhantus (Rhantus) peruvianus Guignot, 1955
Rhantus (Rhantus) calidus (Fabricius, 1792) Rhantus (Rhantus) signatus (Fabricius, 1775)
Copelatinae: Copelatini
Agaporomorphus grandisinuatus Miller, 2001 Copelatus externus Kirsch, 1873
Agaporomorphus knischi Zimmermann, 1921 Copelatus flavicans Guignot, 1952
Agaporomorphus silvaticus Miller, 2005 Copelatus fryi Balfour-Browne, 1939
Agaporomorphus tambopatensis Miller, 2005 Copelatus mathani Guignot, 1952
Aglymbus eminens (Kirsch, 1873) Copelatus normalis Erichson, 1847
Copelatus amazonicus Regimbart, 1899 Copelatus speciosus Regimbart, 1892
Copelatus blancasi Guignot, 1958 Copelatus substriatus Kirsch, 1873
Dytiscinae: Aciliini
Thermonectus margineguttatus (Aube, 1838)
Thermonectus succinctus (Aube, 1838)
Dytiscinae: Cybistrini
Megadytes (Megadytes) laevigatus (Olivier, 1795)
Megadytes (Megadytes) steinheili (Wehncke, 1876)
Dytiscinae: Eretini
Eretes sticticus (Linnaeus, 1767)
Hydrotinae: Hydrodytini
Hydrodytes opalinus (Zimmermann, 1921)
Hydroporinae: Bidessini
Amarodytes undulates Gschwendtner, 1954 Liodessus acollensis Guignot, 1955
Bidessonotus melanocephalus Regimbart, 1895 Liodessus affinis (Say, 1823)
Bidessonotus obtusatus Regimbart, 1895 Liodessus andinus Guignot, 1957
Bidessonotus tibialis Regimbart, 1895 Liodessus cancellosus Guignot, 1957
Bidessonotus truncates Balfour-Browne, 1947 Liodessus delfini (Regimbart, 1899)
Brachyvatus apicatus (Clark, 1862) Liodessus flavofasciatus
Hypodessus crucifer Guignot, 1939 (Steinall, 1869)
Hypodessus frustrator Spangler, 1966 Neobidessus alvarengai Young, 1981
Hypodessus titschacki (Gschwendtner, 1954) Neobidessus bolivari Young, 1981
Hydroporinae: Hyphydrini
Desmopachria annae Miller, 2005 Desmopachria striga Young, 1990
Desmopachria balionota Miller, 2005 Desmopachria striola Sharp, 1887
Desmopachria pilosa Miller, 2005 Desmopachria tambopatensis Miller, 2005
Hydroporinae: Methlini
Celina latipes (Brulle, 1837)
Hydroporinae: Vatellini
Derovatellus lentus (Wehncke, 1876) Vatellus drymetes Miller, 2005
Derovatellus peruanus Spangler, 1967 Vatellus grandis Buquet, 1840
Derovatellus spangleri Miller, 2005 Vatellus lateralis (Sharp, 1882)
Vatellus annae Miller, 2005
Hydroporinae: tribe unknown

Hydrodessus amazonensis Spangler, 1966
Hydrodessus nanayensis Spangler, 1966
Laccophilinae: Lacophilini
Laccophilus fasciatus terminalis Sharp, 1882 Laccophilus normifer Guignot, 1958
Laccophilus intermedius Regimbart, 1889 Laccophilus oberthueri Regimbart, 1889
Laccophilus mathani Guignot, 1955 Laccophilus plagiatus Regimbart, 1889
Laccophilus nastus Spangler, 1966 Laccophilus ploterus Spangler, 1966
Lancetinae: Lancetini
Lancetes nigriceps (Erichson, 1834)
Lancetes praemorsus (Erichson, 1834)
Acknowledgement
Literature Cited
Indies, and South America. Parts 1 through 6. U. S. Natl. Mus. Bull. 185:ixii, 11492.
Erwin, T. L., C. Micheli, and C. S. Chaboo. In press. Beetles (Coleoptera) of Peru: a survey of the families.
Carabidae Latreille, 1802. J. Kans. Entomol. Soc.
Gschwendtner, L. 1954. Dytiscidae (Col.). Beitrage zur Fauna Perus 4:109115.
Maddison, D. R. 1995. Adephaga. Version 01 January 1995 (under construction). http://tolweb.org/
Adephaga/8875/1995.01.01 in The Tree of Life Web Project, http://tolweb.org/ (Last accessed 28/
06/2014).
Miller, K. B. 2001. Revision of the genus Agaporomorphus Zimmermann (Coleoptera: Dytiscidae). Ann.
Entomol. Soc. Am. 94:520529.
Miller, K. B. 2002. Revision of the subfamily Hydrodytinae Miller (Coleoptera: Dytiscidae) with
description of a new genus. Insect. Syst. Evol. 33:18.
Miller, K. B. 2005a. Revision of the New World and south-east Asian Vatellini (Coleoptera: Dytiscidae:
Hydroporinae) and phylogenetic analysis of the tribe. Zool. J. Linn. Soc. 144:415510.
Miller, K. B. 2005b. Four new species of Desmopachria Babington from Peru (Coleoptera: Dytiscidae).
Zootaxa 1059:3947.
Miller, K. B. 2005c. Two new species of Agaporomorphus Zimmermann (Coleoptera: Dytiscidae) from
Peru. Zootaxa 1059:4959.
Mouchamps, R. 1957. Contribution a la connaissance des Cybisterini (Col. Dytiscidae) du Musee
dHistoire Naturelle de Vienne. Annalen des Naturhistorischen Museums in Wien 61:278287.
Spangler, P. J. 1966. The Catherwood Foundation Peruvian-Amazon Expedition. XIII. Aquatic
Coleoptera (Dytiscidae; Noteridae; Gyrinidae; Hydrophilidae; Dascillidae; Helodidae; Psepheni-
dae; Elmidae). Monogr. Acad. Nat. Sci. 14:377443.
Tremouilles, E. R. 1995. Insecta, Coleoptera, Dytiscidae. Fasciculo 1. Dytiscidae: Methlinae-

Hydroporinae. Fauna de Agua Dulce de la Republica Argentina 37:182.
Young, F. N. 1981. Predaceous water beetles of the genus Neobidessus Young from South America
(Coleoptera: Dytiscidae). Coleopt. Bull. 35:317340.
Young, F. N. 1990a. A review of the classification of the water beetles of the New World genus
Bidessonotus Regimbart (Coleoptera: Dytiscidae: Hydroporinae: Bidessini). Quaest. Entomol.
26:355381.
Young, F. N. 1990b. Predaceous water beetles of the genus Desmopachria Babington: the subgenus
Pachriostrix Guignot (Coleoptera: Dytiscidae). Coleopt. Bull. 44:224228.
Carabidae
Author(s): Terry L. Erwin, Charyn Micheli, and Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 151162
Beetles (Coleoptera) of Peru: A Survey of the Families. Carabidae

TERRY L. ERWIN,1 CHARYN MICHELI,1 AND CAROLINE S. CHABOO2,3
Diversity: 23 tribes, 131 genera, 721 species.

The family Carabidae, commonly known as Ground Beetles, is one of the ten
largest beetle families and is the largest family in the coleopteran suborder
Adephaga. This family was erected by Latreille in 1802 at which time it contained 29
genera. Today, the Carabidae contains an estimated 39,000 described species, in ca.
100 tribes and 1860 genera (Lorenz, 2005). Modern classifications of the Carabidae
include groups that were previously ranked as families, such as Cicindelidae,
Omophronidae, Paussidae, and Rhysodidae.
Recognition: All adult carabids possess three features which distinguish them from
other terrestrial beetles: the hind coxae are fixed in place, the hind coxae divide the
first visible sternite of the abdomen, and they have an internal pair of glands in
the abdomen used for generating defense chemicals. These glands cannot be seen
externally; however, their use produces distinctive and powerful sprays and odors in
many carabid lineages. Two carabid lineages, the Brachinini and Paussinae, are
known as Bombardier and False bombardier beetles, respectively since they have
the ability to explosively discharge defensive chemicals at temperatures of 55100uC
(Aneshansley et al., 1969). This type of defensive system is unique among the beetles
(Bousquet and Larochelle, 1993).
Evolutionary History
Carabid beetles, or ground beetles, constitute an old lineagethey are thought to
have evolved in the early Jurassic, 200 million years ago (Ponomarenko, 1977). The
majority of extant carabid species belong to the subfamily Harpalinae, which is thought
to have radiated in the Cretaceous, some 100 million years ago (Ponomarenko, 1977).
Erwin (1981, 1985) proposed the taxon pulse hypothesis to explain historical
patterns of evolution of lineages within Carabidae. This hypothesis predicts that the
most primitive taxa of carabid lineages were waterside generalists in equatorial
regions. As species dispersed in altitude (e.g., into forest canopies or up mountains
slopes) and out of this region (to the north and south) over evolutionary time, they
became more specialized. Then, these were subsequently replaced by newly-evolved
groups from tropical wetlands. Despite many efforts to infer the phylogeny of
carabid tribes, many deep tribal relationships remain unresolved (see Maddison et al.
(1999) and Maddison (2006, online) for reviews).
1
Department of Entomology, Smithsonian Institution, PO Box 37012, MRC 187, Washington, DC
200137012.
2
Division of Entomology, Natural History Museum and Department of Ecology and Evolutionary
Biology, 1501 Crestline Drive, Suite 140, University of Kansas, Lawrence, KS 66049, USA, E-mail:
cschaboo@ku.edu.
3
Corresponding author.
Accepted 15 May 2014; Revised 15 May 2015
Habits and Habitats

Worldwide, adult carabids range in size from 0.7 to 90.2 mm. Adults of most
species are fast running, night-active, generalist predators of other arthropods or
insect eggs. However, some are specialists. For example: species in the tribes
Peleciini and Promecognathini are millipede hunters; Cychrini and Licinini are snail
hunters; and adults of Notiophilus, Loricera and Leistus eat Collembola; Paussini
and Pseudomorphini live with ants and the former feed on ant workers and ant
brood, and in the genus Pseudomorpha their larvae are fed by ant workers. Other
carabids, such as members of the Harpalini and Zabrini, are seed-eaters, thus
herbivorous.
The larvae of most carabid species are campodeiform with well-developed legs,
antennae and mandibles. They are active and fast-running, and they generally eat
live prey, or seeds. However, some groups, such as the Lebiini, Brachinitae, and
Peleciini, have ectoparasitoid larvae that eat beetle pupae, insect egg clutches, or
young millipedes. Some carabids that are ectoparasitoids in the larval stages, mimic
their adult host in the adult stage (Lindroth, 1971).
Some seed-eating carabids are subsocial; the adults tend clutches of eggs and
young larvae (Brandmayr, 1974). No genera such as this are known from Peru, yet,
but the possibility exists. Tiger beetles have received some attention (Pearson, 1984,
1994; Pearson and Anderson, 1985; Knisley and Hoback, 1994), but unfortunately,
very few way of life studies have been made for Peruvian Carabidae beyond
seasonality that is noted on specimen labels. Some carabids have been studied as
potential biocontrols of agricultural pest herbivores (e.g., Yabar et al., 2006). The
Andean fauna at higher altitudes contain many flightless species (e.g., see Maveety
et al., 2011) while this attribute is very rare in the lowland Amazon portion of the
country.
Because adults of all species of Carabidae have very effective defense glands
located in the posterior part of their abdomen and orifices or turrets to dispense
various types of chemicals (Will et al., 2000), carabids are role models for other
insects without such defense. Aposematism is found in various lineages such as the
bombardier beetles (Brachinini) and both Batesian and Mullerian mimicry
are common. Lindroth (1971) described another type mimicry by disappearance
that involves the genus Lebia and their host chrysomelids. Lebia larvae are
ectoparasitoids on chrysomelid pupae; often the carabid adult closely resembles the
chrysomelid host adult in form and color patterns, suggesting they are associated as
host and parasitoid. Chaboo (2011) shows a remarkable aggressive mimicry between
host Diamphidia leaf beetles and their Lebistina enemy in Africa.
Distribution
Today, carabid species occur between 78u 569 north latitude and 55u south
latitude, and are present on every continent except Antarctica, although they used to
be there, too. They are found in a wide range of microhabitats including the subsoil,
tree tops, sea beaches, cracks in intertidal rocks, at the margins of high mountain
glaciers, and in caves (troglobites), in addition to most soil surface habitats,
especially soil-water interfaces.
Many carabids have well-developed hind wings making them good flyers and
dispersers, which explains their presence on most remote oceanic islands. Some
islands have supported large radiations of certain lineages, for example there are 129
species of the genus Blackburnia Sharp in Hawaii; and 67 and 120 species of the
genus Mecyclothorax Sharp in Tahiti and Hawaii, respectively. On the other hand,
many species of carabids are flightless, their wings reduced by varying degrees. These
species also occur at higher altitudes in the mountains of all continents including
Peru. Flightless carabids rarely occur at lower altitudes, but some do, such as species
of Pelicium Kirby and Asklepia Liebke.
Notes
The carabid fauna known from Peru currently is 721 species, 384 of them being
precinctive. Erwin (1991) reported more than 1000 species of carabids at the lowland
rainforest BIOLAT site, Pakitza, on the Manu River. This area included many
different habitats, but was only a few hundred hectares in size. Most of those species
are undescribed. Maveety et al., (2011) reported 77 morphospecies along an
altitudinal transect in a Peruvian cloud forest in the Kosnipata Valley and estimated
60% of those to be new species.
There are very few keys to species for Peru, those being revisions of isolated genera
that have broader distributions. However, there are keys to carabid genera for South
America (Reichardt, 1977), French Guiana (Erwin et al., 2012), Colombia
(Martnez, 2005), and Costa Rica (Erwin et al., 2002, Link: http://www.inbio.ac.cr/
papers/carabidae/esp/images/CR_carabid_key.doc). The genera covered in these are
mostly also found in Peru.
The species list below indicates that the fauna is far from completely known and
new species are being discovered nearly every year.
Checklist:
Bembidiini
Anillotarsus tetramerus Mateu, 1980
Bembidion allegroi Toledano, 2008 Erwiniana manusculptilis (Erwin, 1994)
Bembidion ancash Toledano, 2008 Erwiniana notesheje (Erwin, 1994)
Bembidion bordoni Toledano, 2008 Erwiniana parkeri (Erwin, 1994)
Bembidion callacalla Toledano, 2008 Erwiniana pfunorum (Erwin, 1994)
Bembidion commissum Erichson, 1847 Erwiniana protosculptilis (Erwin, 1994)
Bembidion guzzettii Toleando, 2008 Erwiniana punctisculptilis (Erwin, 1994)
Bembidion lares Toleando, 2008 Erwiniana quadrata (Erwin, 1994)
Bembidion mallmaense Toledano, 2008 Erwiniana samiria (Erwin, 1994)
Bembidion oxapampa Toledano, 2008 Erwiniana sculpticollis (Bates, 1871)
Bembidion pascoense Toledano, 2008 Geballusa microtreta (Erwin, 1973)
Bembidion peruvianum Brethes, 1920 Gouleta cayennensis (Dejean, 1831)
Bembidion resignatum Erichson, 1847 Gouleta gentryi Erwin, 1994
Bembidion riobamba Toledano, 2008 Gouleta notiophiloides (Erwin, 1973)
Inpa psydroides Erwin, 1978
Bembidion subapterum Darlington, 1934
Liotachys antennatus Bates, 1871
Bembidion zanettii Toledano, 2008
Meotachys platyderus (Bates, 1871)
Cryptocharidius mandubularis Etonti & Mateu, 1992
Pericompsus alcimus Erwin, 1974
Erwiniana am (Erwin, 1994) Pericompsus clitellaris (Erichson, 1847)
Erwiniana angustia (Erwin, 1994) Pericompsus hirsutus Schaum, 1863
Erwiniana crassa (Erwin, 1994) Pericompsus immaculatus Bates, 1871
Erwiniana esheje (Erwin, 1994) Pericompsus rorschachinus Erwin, 1974
Erwiniana eugeneae (Erwin, 1994) Polyderis nympha Erwin, 1984
Erwiniana exigupunctata (Erwin, 1994) Polyderis rapoporti Jeannel, 1962
Erwiniana gruti (Bates, 1871a) Polyderis ucayali Erwin, 1984
Erwiniana iris (Erwin, 1973)
Brachinus atramentarius von Mannerheim, 1837 Brachinus olidus Reiche, 1842

Brachinus bilineatus Laporte de Castelnau, 1836 Brachinus xanthophryus Chaudoir, 1876
Brachinus hylaenus Reichardt, 1967 Pheropsophus aequinoctialis (Linnaeus, 1763)
Calophaeniini
Calophaena acuminata (Olivier, 1790) Calophaena ephippigera Liebke, 1930
Calophaena azurea Liebke, 1938 Calophaena hieroglyphica Liebke, 1930
Calophaena bicincta (Dejean & Boisduval, 1829) Calophaena moseri Liebke, 1930
Calophaena disticnta Chaudoir, 1861 Calophaena peruana Mateu, 1972
Calophaena dupuisi Liebke, 1930 Calophaena pleurostigma Chaudoir, 1861
Carabini
Calosoma abbreviatum Chaudoir, 1869 Oxycheila haenschi W. Horn, 1900
Calosoma fulgens Chaudoir, 1869 Oxycheila oberthueri W. Horn, 1896
Calosoma glabratum Dejean, 1831 Oxycheila pseudoglabra Wiesner, 1999
Calosoma peruviense Mandl, 1971 Oxycheila pseudonigroaenea W. Horn, 1938
Calosoma rufipenne Dejean, 1831 Oxycheila strandi W. Horn, 1913
Catapiesis brasiliensis (Gray, 1832) Oxycheila weyrauchi Mandl, 1967
Catapiesis nitida Solier, 1835 Oxygonia buckleyi Bates, 1872
Homalomorpha castanea Brulle, 1835 Oxygonia delia (Thomson, 1859)
Chlaenius breviusculus Chaudoir, 1876 Oxygonia fleutiauxi W. Horn, 1896
Brasiella argentata (Fabricius, 1801) Oxygonia floridula Bates, 1872
Brasiella balzani (W. Horn, 1907) Oxygonia gloriola Bates, 1872
Brasiella umbrogemmata (W. Horn, 1906) Oxygonia moronensis Bates, 1872
Cheiloxya binotata (Laporte de Castelnau, 1833) Oxygonia oberthueri W. Horn, 1896
Cicindela mathani W. Horn, 1897 Oxygonia prodiga Erichson, 1847
Cicindela rufoaenea W. Horn, 1915 Oxygonia schoenherri (von Mannnerheim, 1837)
Cicindela trifasciata Fabricius, 1781 Oxygonia vuillefroyi Chaudoir, 1869
Cicindela trifasciata Fabricius, 1781 Pentacomia chrysamma Bates, 1872
Cicindela trifasciata Fabricius, 1781 Pentacomia cribrata (Brulle, 1836)
Cylindera malaris (W. Horn, 1896) Pentacomia egregia (Chaudoir, 1835)
Cylindera mixtula (W. Horn, 1915) Pentacomia lacordairei (Gory, 1833)
Cylindera suturalis (Fabricius, 1798 Pentacomia pentacomioides (W. Horn, 1900)
Eucallia boussingaulti Guerin-Meneville, 1843 Pentacomia vallicola Huber, 1999
Iresia egregia (Chaudoir, 1860) Pentacomia ventralis (Dejean, 1825)
Iresia psyche Sumlin, 1994 Phaeoxantha aequinoctialis (Dejean, 1825)
Langea euprosopoides W. Horn, 1901 Phaeoxantha aequinoctialis (Dejean, 1825)
Langea mellicollis Sumlin, 1993 Phaeoxantha klugii (Chaudoir, 1850)
Metriocheila nigricollis Reiche, 1842 Pseudoxycheila andina Cassola, 1997
Odontocheila annulicornis (Brulle, 1836) Pseudoxycheila angustata Chaudoir, 1865
Odontocheila baeri Fleutiaux, 1903 Pseudoxycheila aymara Cassola, 1997
Odontocheila castelnaui Lucas, 1857 Pseudoxycheila bipustulata (Latreille, 1812)
Odontocheila cayennensis (Fabricius, 1787) Pseudoxycheila ceratoma Chaudoir, 1865
Odontocheila confusa (Dejean, 1825) Pseudoxycheila chaudoiri Dokhtouroff, 1882
Odontocheila consobrina Lucas, 1857 Pseudoxycheila confusa Cassola, 1997
Odontocheila consobrina Lucas, 1857 Pseudoxycheila immaculata W. Horn, 1905
Odontocheila cyanella Chaudoir, 1860 Pseudoxycheila inca Cassola, 1997
Odontocheila eximia Lucas, 1857 Pseudoxycheila lateguttata Chaudoir, 1844
Odontocheila ignita Chaudoir, 1860 Pseudoxycheila lateguttata peruviana Cassola, 1997
Odontocheila margineguttata (Dejean, 1825) Pseudoxycheila nitidicollis Cassola, 1997
Odontocheila marginilabris Erichson, 1847 Pseudoxycheila quechua Cassola, 1997
Odontocheila rufiscapis Bates, 1874 Ronhuberia eurytarsipennis (W. Horn, 1905)
Odontocheila tricuspipenis W. Horn, 1932 Tetracha brasiliensis (Kirby, 1819)
Odontocheila trilbyana Thomson, 1857 Tetracha buchardi Naviaux, 2007
Oxycheila barkleyi Wiesner, 1999 Tetracha chilensis (Laporte de Castelnau, 1831)
Oxycheila fleutiauxi W. Horn, 1898 Tetracha flammula (W. Horn, 1905)
Oxycheila germaini Fleutiaux, 1892 Tetracha fulgida (Klug, 1834)
Tetracha horni Ruge, 1892 Tetracha spinosa (Brulle, 1836)

Tetracha inca (Naviaux & Ugarte-Pena, 2006) Tetracha spixii (Brulle, 1836)
Tetracha latreillei (Laporte de Castelnau, 1834) Tetracha spixii (Brulle, 1836)
Tetracha lucifera Erichson, 1847 Tetracha spixii (Brulle, 1836)
Tetracha pilosipennis (Mandl, 1958) Tetracha strandi W. Horn, 1913
Tetracha sobrina (Dejean, 1831) Tetracha suturalis W. Horn, 1900
Collyridini
Ctenostoma agnatum Chaudoir, 1860 Teukrus cruciatus (Bates, 1871)
Ctenostoma agnatum Chaudoir, 1860 Anaulacus ashei Ball & Shpeley, 2002
Ctenostoma compactum Naviaux, 1998 Anaulacus batesi (Chaudoir, 1876)
Ctenostoma crucifrons W. Horn, 1911 Anaulacus erwini Ball & Shpeley, 2002
Ctenostoma deuvei Naviaux, 1998 Anaulacus exiguus Ball & Shpeley, 2002
Ctenostoma formicarium (Fabricius, 1801) Anaulacus sericatus Chaudoir, 1846
Ctenostoma inca Naviaux, 1998 Tetragonoderus deuvei Shpeley & Ball, 2008
Ctenostoma magnum Naviaux, 1998 Tetragonoderus eximius Kirsch, 1873
Ctenostoma metallicum (Laporte de Castelnau, 1834) Tetragonoderus foveicollis Liebke, 1941
Ctenostoma nigrum Chaudoir, 1860 Tetragonoderus rivularis Erichson, 1847
Ctenostoma obliquatum Chaudoir, 1860 Tetragonoderus sivianus Liebke, 1941
Ctenostoma plicaticolle W. Horn, 1911 Tetragonoderus sticticus Erichson, 1847
Ctenostoma regium Naviaux, 1998 Neodrypta costigera (Chaudoir, 1861)
Ctenostoma rugicolle W. Horn, 1904 Ancystroglossus dimidiaticornis Chaudoir,
Ctenostoma sumlini Naviaux, 1998 1863
Amblycoleus peruanus Liebke, 1928 Galerita affinis (Dejean, 1831)
Amblycoleus pluriseriatus (Chaudoir, 1877) Galerita brachinoides Perty, 1830
Askalaphium depressum (Bates, 1871) Galerita inca Reichardt, 1967
Ctenodactyla batesi Chaudoir, 1861 Galerita jelskii Chaudoir, 1877
Ctenodactyla batesi Chaudoir, 1861 Galerita simplicicarinata Reichardt, 1967
Leptotrachelus mexicanus (Chaudoir, 1852) Galerita unicolor Latreille & Dejean, 1824
Harpalini
Amblygnathus bicolor Ball & Maddison, 1987 Pelmatelus sp. C
Amblygnathus georgei Shpeley, 2008 Pelmatelus sp. D
Amblygnathus gilvipes Ball & Maddison, 1987 Pelmatelus sp. E
Amblygnathus janthinus Dejean, 1829 Pelmatelus sp. F
Amblygnathus suturalis Putzeys, 1845 Pelmatelus sp. G
Anisostichus laevis (Curtis, 1839) Pelmatelus sp. H
Gonicellus sp. A Pelmatelus sp. I
Harpalus turmalinus Erichson, 1847 Pelmatelus sp. J
Notiobia aguilarorum Noonan, 1981 Pelmatelus sp. K
Notiobia aulica (Dejean, 1829) Pelmatelus sp. L
Notiobia chalcites (Germar, 1824 Polpochila marginalis Negre, 1963
Notiobia disparilis Bates, 1878 Polpochila sulcata Negre, 1963
Notiobia flavicincta (Erichson, 1847) Polpochila vicina Negre, 1963
Notiobia glabrata Arndt, 1998 Selenophorus punctiger Kirsch, 1873
Notiobia incerta Bates, 1882 Stenolophus badius Erichson, 1847
Notiobia moffetti Noonan, 1981 Stenolophus debilis Erichson, 1847
Notiobia peruviana (Dejean, 1829 Stenolophus lentulus Erichson, 1847
Notiobia schnusei (van Emden, 1953) Stenolophus longicollis Erichson, 1847
Notiobia viridula (Dejean, 1829) Trichopselaphus magnificus Ball, 1978
Pelmatelus sp. A Trichopselaphus subiridescens Chaudoir,
Pelmatelus sp. B 1843
Helluonini
Dailodontus cayennensis (Dejean, 1826) Helluomorpha macroptera Chaudoir, 1850
Helluobrochus brevicollis (Dejean, 1831) Helluomorphoides diana Reichardt, 1974
Helluobrochus cribratus (Reiche, 1843) Helluomorphoides unicolor (Brulle, 1837)
Helluobrochus subrostratus (Bates, 1871) Pleuracanthus inca Reichardt, 1974
Lachnophorini
Anchonoderus montanus Liebke, 1941 Lachnophorus gibbosus Liebke, 1936
Anchonoderus roedingeri Liebke, 1941 Lachnophorus integer Liebke, 1936
Aporesthus anomalus Bates, 1871 Pseudophoriticus sp. A
Aporesthus titschacki (Liebke, 1941) Pseudophoriticus sp. B
Lebiini
Agra acuspina Straneo, 1979 Agra nodicornis Straneo, 1965
Agra acutidens Straneo, 1966 Agra obscuritibiis Straneo, 1965
Agra aeris Erwin, 2000 Agra pala Liebke, 1937
Agra andina Liebke, 1938 Agra pearsoni Erwin, 1984
Agra azureipennis Erwin, 1982 Agra peruana Liebke, 1941
Agra basilewskyi Straneo, 1955 Agra plantipedis Liebke, 1941
Agra biexcavata Straneo, 1958 Agra platyscelis (Chaudoir, 1861)
Agra biolat Erwin, 2000 Agra pseudoboliviana Straneo, 1965
Agra brunneitarsis Straneo, 1958 Agra pseudomoesta Straneo, 1965
Agra ce Erwin, 2010 Agra pseudovarians Straneo, 1965
Agra clavata Liebke, 1941 Agra punctulata Liebke, 1941
Agra cochlearis Liebke, 1938 Agra quadrilamata Straneo, 1965
Agra communis Straneo, 1965 Agra quararibea Erwin, 1993
Agra conhormigas Erwin, 2000 Agra regina Liebke, 1941
Agra conicollis Straneo, 1965 Agra rubrofemorata Straneo, 1958
Agra crassipes Liebke, 1941 Agra rubroviolacea Straneo, 1965
Agra crebrefoveata Straneo, 1965 Agra rufonigra Straneo, 1965
Agra dentipennis Liebke, 1941 Agra saltatrix Erwin, 1982
Agra dominula Bates, 1865 Agra samiria Erwin, 2000
Agra dora Erwin, 1984 Agra satipo Erwin, 1984
Agra exsculpta Liebke, 1941 Agra saundersi Bates 1865
Agra fallax Liebke, 1941 Agra seriefoveata Chaudoir, 1877
Agra fimbriata Straneo, 1965 Agra servatorum Erwin, 2000
Agra foraminosa Liebke, 1938 Agra seticollis Straneo, 1982
Agra grace Erwin, 2010 Agra setigera Liebke, 1938
Agra huambana Straneo, 1982 Agra simillima Straneo, 1965
Agra humboldti Liebke, 1941 Agra sphenarion Erwin, 1982
Agra inca Erwin, 1986 Agra spinicauda Straneo, 1982
Agra iota Erwin, 1984 Agra splendida Dejean, 1829
Agra iquitosana Erwin, 1982 Agra strangulata Chaudoir, 1863
Agra ketschuana Liebke, 1941 Agra striatopunctata Chaudoir, 1866
Agra klugii Brulle 1836 Agra tarapoto Erwin, 1984
Agra lacrymosa Straneo, 1965 Agra tarapotoana Erwin, 1982
Agra laetipes Straneo, 1965 Agra tingo Erwin, 2000
Agra lamproptera Chaudoir, 1847 Agra tingomaria Erwin, 1984
Agra latifemoris Straneo, 1982 Agra titschacki Liebke, 1941
Agra limulus Erwin, 1982 Agra trochanterica Straneo, 1965
Agra lindae Erwin, 2000 Agra tuberculata Straneo, 1965
Agra longelytrata Straneo, 1965 Agra tuitis Erwin, 1987
Agra magnifica Erwin, 2000 Agra vation Erwin, 1983
Agra manu Erwin, 2000 Agra venatrix Liebke, 1941
Agra mathani Straneo, 1965 Agra venustula Straneo, 1958
Agra misella Straneo, 1965 Agra vidua Straneo, 1965
Agra mnemosine Straneo, 1965 Agra viridicollis Straneo, 1965
Agra modesta Straneo, 1979 Agra vulgaris Straneo, 1965
Agra monticola Liebke, 1938 Calleida horni Liebke, 1934
Agra mulieris Liebke, 1941 Calleida jeanneli Liebke, 1935
Agra multipunctata Straneo, 1965 Calleida migratoria Casale, 2002
Agra nigritarsis Straneo, 1982 Calleida mniszechi Chaudoir, 1852
Agra nigrotibiata Straneo, 1965 Calleida prolixa Erichson, 1847
Calleida tibialis Brulle, 1836 Gallerucidia basinotata Chaudoir, 1872

Calleida titschacki Liebke, 1941 Hybopteroides biolat Erwin & Ball, 2012
Calleida viridiaurea Chaudoir, 1877 Hybopteroides karolynae Erwin & Ball, 2012
Catascopus brasiliensis Dejean, 1831 Lebia andina Liebke, 1939
Chelonodema albovariegata (Chaudoir, 1871) Lebia angustula Oberthur, 1883
Chelonodema boliviensis (Chaudoir, 1871) Lebia callanga Liebke, 1935
Chelonodema erotyloides Reichardt, 1972 Lebia declivis Liebke, 1934
Chelonodema sellata (Dejean, 1825) Lebia diehli Liebke, 1941
Chelonodema toroana (Liebke, 1941) Lebia distigma Liebke, 1934
Coptodera acutipennis (Buquet, 1834) Lebia exigua Kirsch, 1873
Coptodera chalcites Bates, 1869 Lebia furcatula Liebke, 1934
Coptodera cupreotincta Bates, 1869 Lebia graphica Liebke, 1938
Coptodera erwini Shpeley & Ball, 1993 Lebia horni Liebke, 1934
Coptodera festiva Dejean, 1825 Lebia jucunda Kirsch, 1873
Coptodera megalops Bates, 1869 Lebia lauta Liebke, 1938
Coptodera nigrostriata (Reiche, 1843) Lebia mesoxantha (Kirsch, 1873)
Coptodera nitidula (Buquet, 1834) Lebia peruana Lutshnik, 1922
Coptodera pakitza Shpeley & Ball, 1993 Lebia reticulata Liebke, 1938
Coptodera picea Dejean, 1826 Lebia sculpticollis Liebke, 1941
Coptodera tripartita Chaudoir, 1869 Lebia silvatica Liebke, 1941
Coptodera tripucntata Shpeley & Ball, 1993 Lebia trullata Liebke, 1938
Coptodera undulata Perty, 1830 Lebia vilcanota Liebke, 1934
Coptodera versicolor Bates, 1869 Lelis quadrisignata (Buquet, 1834)
Coptodera waytkowskii Liebke, 1941 Lelis rutila (Bates, 1869)
Cylindronotum chalceum (Kirsch, 1873) Mimodromius altus Liebke, 1941
Epikastea biolat Erwin, 2004 Mimodromius bicolor (Brulle, 1836)
Epikastea grace Erwin, 2004 Mimodromius leleupi Mateu, 1970
Epikastea mancocapac Erwin, 2004 Mimodromius lojanus Liebke, 1935
Epikastea poguei Erwin, 2004 Mimodromius negrei Mateu, 1960
Eucheila boliviana (Mateu, 1989) Mimodromius peruvianus Mateu, 1955
Eucheila breviformis (Chaudoir, 1872) Mimodromius weyrauchi Mateu, 1970
Eucheila costulata (Chaudoir, 1872) Negrea peruviana Mateu, 1982
Eucheila erwini Ball & Shpeley, 2001 Nemotarsus titschacki (Liebke, 1941)
Eucheila inpa (Ball & Shpeley, 1983) Pseudotoglossa inaequalis (Chaudoir, 1872)
Eucheila marginata Shpeley & Ball, 2001 Pseudotoglossa terminalis (Chaudoir, 1872)
Eucheila mateui Ball & Shpeley, 2001 Stenognathus crenulatus Chaudoir, 1869
Eucheila purpurea (Ball & Shpeley, 1983) Stenognathus dentifer Chaudoir, 1869
Eucheila reichardti Ball & Shpeley, 1983 Stenognathus jauja Shepley & Ball, 2001
Eucheila flavilabris Ball & Shpeley, 1983 Stenognathus luctuosus (Maindron, 1906)
Eurycoleus fofus Reichardt, 1976 Stenognathus melanarius (Dejean 1831)
Eurycoleus poecilopterus (Buquet, 1834) Stenognathus plaumanni Shepley & Ball, 2001
Eurycoleus tredecimpunctatus Chaudoir, 1869 Thoasia rugifrons Liebke, 1939
Licini
Badister amazonus Erwin & Ball, 2011
Metiini
Metius apicalis Straneo, 1987 Metius obtusus Straneo, 1951
Metius auratoides Straneo, 1987 Metius peruvianus Straneo, 1951
Metius cordatulus Straneo, 1951 Metius quadricollis Straneo, 1987
Metius incertus (Putzeys, 1873) Metius striatus (Putzeys, 1873)
Metius loeffleri Straneo, 1987 Metius striolatus Straneo, 1951
Metius mateui Straneo, 1987 Metius submetallicus Straneo, 1987
Metius negrei Straneo, 1977 Metius titschacki Straneo, 1951
Morionini
Morion cordatus (Chaudoir, 1837) Morion simplex Dejean, 1826
Morion cyclomus Chaudoir, 1854 Moriosomus motschulskyi Erwin & Moore, 2007
Odacanthini
Colliuris crispa Klug, 1834 Colliuris maculipennis Liebke, 1930
Colliuris emdeni Liebke, 1930 Colliuris peruana Erichson, 1847
Colliuris excellens Liebke, 1930 Colliuris robusta Liebke, 1930
Colliuris flavipes (Chaudoir, 1850) Colliuris rugicollis (Dejean, 1825)
Colliuris laeviceps Liebke, 1930 Stenocheila lacordairei Laporte de Castelnau, 1832
Oodini
Dercylus buckleyi (Chaudoir, 1882) Dercylus heynei (Kuntzen, 1912)
Dercylus catenatus Kuntzen, 1912 Dercylus mathani Moret, 1995
Ozaenini
Filicerozaena reducticollis Deuve, 2009 Pachyteles sp.
Ozaena manu Ball & Shpeley, 1990 Platycerozaena brevicornis (Bates, 1874)
Pachyteles peruvianus Bates, 1874
Peleciini
Eripus franzi Straneo & Ball, 1989 Stricteripus peruvianus (Straneo, 1955)
Pentagonicini
Pentagonica roedingeri Liebke, 1941
Perigonini
Diploharpus sp. A
Platynini
Dysocolus sp. A Incagonum inca (Moret, 1994)
Dysocolus sp. B Incagonum mateui (Moret, 1994)
Dysocolus sp. C Platynus aemulus (Moret, 1996)
Dysocolus sp. D Platynus ambagiosus (Moret, 1990)
Dysocolus sp. E Platynus callanga (Perrault, 1992)
Dysocolus sp. F Platynus contumaza (Perrault, 1993)
Dysocolus sp. G Platynus cyphopterus (Perrault, 1993)
Dysocolus sp. H Platynus etonii (Moret, 1996)
Dysocolus sp. I Platynus giachinoi (Moret, 1996)
Dysocolus sp. J Platynus interstitialis (Perrault, 1993)
Dysocolus sp. K Platynus jelskii (Perrault, 1990)
Dysocolus sp. L Platynus matucana (Perrault, 1993)
Dysocolus sp. M Platynus nebrianus (Fairmaire, 1878)
Dysocolus sp. N Platynus punoensis (Perrault, 1990)
Glyptolenus sp. A Platynus schunkei (Perrault, 1993)
Pseudomorphini
Manumorpha biolat Erwin & Geraci, 2008 Samiriamorpha grace Erwin & Geraci, 2008
Psydrini
Tropopterus peruvianus Straneo, 1954
Pterostichini
Abaris basistriata Chaudoir, 1873 Abaris erwini Will, 2002
Abaris opaca Will, 2002 Blennidus negrei (Straneo, 1993)

Abaris striolata Bates, 1871 Blennidus nigritulus (Straneo, 1993)
Blennidus abditus Giachino & Allegro, 2011 Blennidus olivaceus (Tschitscherine, 1897)
Blennidus abramalagae Allegro & Giachino, 2011 Blennidus orbicollis (Straneo, 1993)
Blennidus angularis (Straneo, 1993) Blennidus pachycerus (Tschitscherine, 1897)
Blennidus anxius (Tschitscherine, 1898) Blennidus pascoensis (Straneo, 1954)
Blennidus aulacostigma (Tschitscherine, 1897) Blennidus peruvianus (Dejean, 1828)
Blennidus azurescens Straneo, 1986 Blennidus phaenogonus (Tschitscherine, 1898)
Blennidus bellesi (Straneo, 1993) Blennidus pinguis Allegro & Giachino, 2011
Blennidus bordoni (Straneo, 1993) Blennidus procerus Allegro & Giachino, 2011
Blennidus crassus (Straneo, 1993) Blennidus pseudolaevis Allegro & Giachino, 2011
Blennidus curtatus (Straneo, 1993) Blennidus pseudoperuvianus Straneo, 1986
Blennidus cuzcanus Straneo, 1986) Blennidus quadrilmatus Straneo, 1986
Blennidus diminutus (Chaudoir, 1878) Blennidus rectangulus (Straneo, 1993)
Blennidus egens (Tschitscherine, 1898) Blennidus refleximargo (Straneo, 1993)
Blennidus etontii Allegro & Giachino, 2011 Blennidus robustulus (Straneo, 1954)
Blennidus euphaenops (Tschitscherine, 1898) Blennidus straneoi Allegro & Giachino, 2011
Blennidus ferrugineicornis Motschulsky, 1866) Blennidus sublaevis (Straneo, 1993)
Blennidus filicornis (Straneo, 1993) Blennidus tardus (Tschitscherine, 1898)
Blennidus fitzcarraldi Giachino & Allegro, 2011 Blennidus tenenbaumi (Lutshnik, 1927)
Blennidus foveatus Straneo, 1951 Blennidus ticlianus (Straneo, 1993)
Blennidus franzanus (Straneo, 1971) Blennidus unistria (Straneo, 1993)
Blennidus huascarani Allegro, 2010 Blennidus vancoveriensis (Chaudoir, 1878)
Blennidus idioderus (Tschitscherine, 1898) Blennidus vereshaginae (Straneo, 1993)
Blennidus inca (Tschitscherine, 1898) Loxandrus rotundicollis Straneo, 1991
Blennidus jelskii (Tschitscherine, 1897) Loxandrus semperfidelis Will, 2008
Blennidus laevis (Straneo, 1993) Loxandrus sericeus Straneo, 1955
Blennidus languens (Tschitscherine, 1898) Metoncidus epiphytus Will, 2004
Blennidus laurentianus Straneo, 1986) Metoncidus gracilis Will, 2004
Blennidus longiloba (Straneo, 1993) Parhypates herberti Straneo, 1987
Blennidus mateui (Straneo, 1993) Pseudobarys sp.
Blennidus mathani Moret, 1995 Tichonilla peruana (Lutshnik, 1931)
Blennidus mauritii Straneo, 1986 Trirammatus chaudoiri (Guerin-Meneville,
Blennidus mesotibialis (Straneo, 1993) 1838)
Blennidus minor (Straneo, 1993) Trirammatus substriatulus (Straneo, 1967)
Scaritini
Ardistomis seriepunctatus (Brulle, 1837) Scarites aberrans Banninger, 1941
Camptidius ophthalmicus Putzeys, 1866) Scarites illustris Chaudoir, 1880
Camptodontus amazonum Putzeys, 1866 Scarites similis Chaudoir, 1880
Distichus angustiformis (Chaudoir, 1880) Scarites vilcanotanus (Banninger, 1932)
Distichus ebeninus (Lynch Arribalzaga, Schizogenius dyschirioides Putzeys, 1863
1878) Schizogenius impressicollis Putzeys, 1846
Distichus peruvianus (Dejean, 1831) Schizogenius janae Kult, 1950
Distichus semicarinatus (Chaudoir, 1880) Schizogenius putzeysi Kirsch, 1873
Distichus smithi (Linell, 1898) Schizogenius sellatus Putzeys, 1866
Distichus trivialis Chaudoir, 1880) Schizogenius xanthopus (Brulle, 1837)
Dyschirius macrophthalmus (Fedorenko, 1999) Semiardistomis exspectatus Valdes, 2012
Dyschirius peruanus Fedorenko 1991 Semiardistomis glabratus (Putzeys, 1866)
Dyschirius selvas (Fedorenko, 1999) Semiardistomis jedlickai (Kult, 1950)
Dyschirius weyrauchi Kult, 1950 Semiardistomis major Valdes, 2012
Forcipator grandis (Perty, 1830) Semiardistomis pallipes (Dejean, 1831)
Kultianella sulcicollis (Putzeys, 1866) Stratiotes batesi Putzeys, 1866
Oxydrepanus micans Putzeys, 1866 Whiteheadiana minor (Putzeys, 1867)
Trechini
Andinodontis maveetyae Erwin & Maddison, 2010 Andinorites atahualpai Etonti & Mateu, 2000
Andinorites convexus Mateu & Belles, 1981 Trechisibus lamasi Etonti & Mateu, 1992
Andinorites crypticola Mateu & Belles, 1981 Trechisibus laresensis Etonti & Mateu, 1996
Andinorites peruvianus Mateu & Belles, 1981 Trechisibus latellai Mateu & Etonti, 2006
Andinorites striatus Mateu & Belles, 1981 Trechisibus leechi Ueno, 1972
Andinorites troglophilus Mateu & Belles, 1981 Trechisibus liberatrix Mateu, 1979
Andinorites vilchezi Mateu & Belles, 1981 Trechisibus loeffleri Jeannel, 1958
Aputrechisibus dubius Trezzi, 2007 Trechisibus martinezi Mateu, 1978
Incatrechus lagunensis Mateu & Etonti, 2006 Trechisibus maucauensis Etonti & Mateu,
Incatrechus pilosus Mateu & Belles, 1982 20025
Incatrechus rattii Etonti & Mateu, 2000 Trechisibus michelbacheri Ueno, 1972
Incatrechus tenuis Etonti & Mateu, 2002 Trechisibus minutus Etonti & Mateu, 1996
Luyatrechus cuelapensis Etonti & Mateu, 2000 Trechisibus missionis Allegro, Giachino & Sciaky,
Oxytrechus cyathiderus Jeannel, 1954 2008
Oxytrechus gitzeni Etonti, 2002 Trechisibus nicki Schweiger, 1959
Oxytrechus paredesi Etonti & Mateu, 1992 Trechisibus obesus Jeannel, 1954
Perucharidius andinus Etonti & Mateu, 2002 Trechisibus olympicus Allegro, Giachino & Sciaky,
Pseudotrechisibus sphaericus Mateu & Belles, 1982 2008
Putzeysius coecus Sciaky & Grottolo, 1996 Trechisibus orophilus Etonti & Mateu, 2002
Trechisibus acutangulus Mateu & Belles, 1981 Trechisibus ovalis Jeannel, 1958
Trechisibus alticola Mateu, 1979 Trechisibus pascoensis Mateu, 1978
Trechisibus amesi Etonti & Mateu, 1992 Trechisibus patarcochae Allegro, Giachino &
Trechisibus amplipennis Etonti & Mateu, 1996 Sciaky, 2008
Trechisibus arduus Mateu, 1979 Trechisibus peruvianus Jeannel, 1927
Trechisibus bellesi Etonti & Mateu, 2002 Trechisibus pierrei Mateu, 1979
Trechisibus bohorquezae Etonti & Mateu, 19920 Trechisibus pluvialis Etonti & Mateu, 2000
Trechisibus bordoni Mateu, 1978 Trechisibus punaensis Etonti & Mateu, 2000
Trechisibus brevicornis Ueno, 1976 Trechisibus quierocochensis Mateu, 1978
Trechisibus callanganus Jeannel, 1937 Trechisibus quietus Etonti & Mateu, 2000
Trechisibus chacasinus Allegro, Giachino & Sciaky, Trechisibus rarianus Etonti & Mateu, 1996
2008 Trechisibus rossi Ueno, 1972
Trechisibus chaudoiri Jeannel, 1954 Trechisibus schmidti Ueno, 1971
Trechisibus chucurensis Trezzi, 2007 Trechisibus spelaeus Mateu & Belles, 1981
Trechisibus convexiusculus Jeannel, 1962 Trechisibus subglobosus Mateu & Belles, 1981
Trechisibus crassipes Ueno, 1972 Trechisibus tenuitarsis Mateu, 1979
Trechisibus cuzcoensis Etonti & Mateu, 1996 Trechisibus theresiae Etonti & Mateu, 19969
Trechisibus decensii Allegro, Giachino & Sciaky, Trechisibus ticliensis Trezzi, 2007
2008 Trechisibus tripunctatus Jeannel, 1958
Trechisibus dispar Jeannel, 1937 Trechisibus trisetosus Jeannel, 1958
Trechisibus eleonorae Allegro, Giachino & Sciaky, Trechisibus ukupachensis Trezzi, 2007
2008 Trechisibus valenciai Etonti & Mateu, 1992
Trechisibus franzi Mateu & Negre, 1972 Trechisibus variicornis (Putzeys, 1870)
Trechisibus gigas Trezzi, 2007 Trechisibus veneroi Etonti & Mateu, 1992
Trechisibus gonzalesi Etonti & Mateu, 1992 Trechisibus ventricosus Jeannel, 1958
Trechisibus homaloderoides Ueno, 1976 Trechisibus vivesi Mateu, 2000
Trechisibus inca Mateu, 1979 Trechisibus wachucochae Allegro, Giachino &
Trechisibus incertus Etonti & Mateu, 19962 Sciaky, 2008
Trechisibus infuscatus Mateu, 1978 Trechisibus yanamensis Allegro, Giachino &
Trechisibus lacunensis Mateu, 1978 Sciaky, 2008
Acknowledgements
We acknowledge NSF-EPSCoR # 66928 (to Caroline S. Chaboo) for supporting
the Beetles of Peru project and the University of Kansas Ecology Evolutionary
Biology-General Research Fund (to Chaboo) for funding this publication.
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Bousquet, Y., and A. Larochelle. 1993. Catalogue of the Geadephaga (Coleoptera: Trachypachidae,
Rhysodidae, Carabidae including Cicindelini) of America north of Mexico. Memoirs of the
Entomological Society of Canada 167:1397.
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Chaboo, C. S. 2011. Defensive behaviors in leaf beetles: from the unusual to the weird. In J. M. Vivanco
and T. Weir (eds.). Chemical Biology of the Tropics, Signaling and Communication in Plants
8:5969. doi:10.1007/978-3-642-19080-3_4, Springer-Verlag, Berlin, Heidelberg.
Erwin, T. L. 1981. Taxon pulses, vicariance, and dispersal: an evolutionary synthesis illustrated by carabid
beetles. In G. Nelson and D. Rosen (eds.). Vicariance Biogeography: A Critique, pp. 159183.
Columbia University Press, New York.
Erwin, T. L. 1985. The taxon pulse: a general pattern of lineage radiation and extinction among carabid
beetles. In G. E. Ball (ed.). Taxonomy, phylogeny, and zoogeography of beetles and ants: A volume
dedicated to the memory of Philip Jackson Darlington Jr.19041983, pp. 437472. Dr. W. Junk b.v.
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Erwin, T. L. 1991. Natural history of the carabid beetles at the BIOLAT Rio Manu Biological Station,
Pakitza, Peru. Revista Peruana de Entomologia 33:185.
Erwin, T. L., C. Micheli, and G. Hevel. 2012. Neotropical beetles of Guyane: Genera of the family
Carabidae (Coleoptera) with notes on species richness currently known from the literature and
recent collections. Le Coleopteriste 5:188.
Knisley, C. B., and W. H. Hoback. 1994. Nocturnal roosting of Odontocheila confusa Dejean in the
Peruvian Amazon. The Coleopterists Bulletin 48:353354.
Lindroth, C. H. 1971. Disappearance as a protective factor: a supposed case of Batesian mimicry among
beetles (Carabidae and Chrysomelidae). Entomologica Scandinavica 2:4148.
Lorenz, W. 2005. Systematic list of extant ground beetles of the world (Insecta, Coleoptera Geadephaga:
Trachypachidae and Carabidae incl. Paussinae, Cicinidelinae, Rhysodinae). 2nd Edn, iv + 530 pp.
Privately published, W. Lorenz, Tutzing, Germany.
Maddison, D. R. 2006. Carabidae. Ground beetles and tiger beetles. Version 11 April 2006 (under
construction). http://tolweb.org/Carabidae/8895/2006.04.11 in The Tree of Life Web Project, http://
tolweb.org/ [accessed 4 April 2014]
Maddison, D. R., M. D. Baker, and K. A. Ober. 1999. Phylogeny of carabid beetles as inferred from 18S
ribosomal DNA (Coleoptera: Carabidae). Syst. Entomol. 24:103138.
Martnez, C. 2005. Introduction a los escarabajos Carabidae (Coleoptera) de Colombia, Colombia,
Diversa por Naturaleza. 546 pp.
Maveety, S. A., R. A. Browne, and T. L. Erwin. 2011. Carabidae diversity along an altitudinal gradient in
a Peruvian cloud forest (Coleoptera). In T. L. Erwin (ed.). Proceedings of a symposium honoring
the careers of Ross and Joyce Bell and their contributions to scientific work, pp. 651666.
Burlington, Vermont, 1215 June 2010. ZooKeys 147: ixii, 689 pp.
Pearson, D. L. 1984. The tiger beetles of the Tambopata Reserved Zone, Madre de Dios, Peru. Revista
Peruana de Entomologia 27:1524.
Pearson, D. L. 1994. Escarabajos tigre del Peru (Coleoptera: Cicindelidae). Lista preliminary de especies
y clave para generos. Revista Peruana de Entomologia 36:5558.
Pearson, D. L., and J. J. Anderson. 1985. Perching heights and nocturnal communal roosts of some tiger
beetles (Coleoptera: Cincidellidae) in southeastern Peru. Biotropica 17:126129.
Ponomarenko, A. G. 1977. Suborder Adephaga. In Arnoldi, L. V., V. V. Zherikhin, L. M. Nikritin, and
A. G. Ponomarenko (eds.). Mesozoic Coleoptera [in Russian], pp. 17104. Trudy Paleontolo-
gicheskogo Instituta Akademiya Nauka SSSR No. 161. 204 pp. [Note. An English translation of
the book was published in 1992 by the Smithsonian Institution Libraries.]
Reichardt, H. 1977. A synopsis of the genera of Neotropical Carabidae. Quaestiones entomologicae
13:346493.
Will, K. W., A. B. Attygalle, and K. Herath. 2000. New defensive chemical data for ground beetles
(Coleoptera: Carabidae): interpretations in a phylogenetic framework. Biological Journal of the
Linnean Society 71:459481.
Yabar, E., E. Castro, L. Melo, and E. Gianoli. 2006. Predacion de Bembidion sp., Notiobia peruviana
(Dejean) y Metius sp. (Coleoptera: Carabidae) sobre huevos de Premnotrypes latithorax (Pierce)
(Coleoptera: Curculionidae) en condiciones de laboratorio. Revista peruana Entomologia
45:9194.
Dryopidae, Limnichidae, Luctrochidae, Psphenidae, and
Ptilodactylidae
Author(s): William D. Shepard and Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 163166
Beetles (Coleoptera) of Peru: A Survey of the Families. Dryopidae,

Limnichidae, Luctrochidae, Psphenidae, and Ptilodactylidae
WILLIAM D. SHEPARD1 AND CAROLINE S. CHABOO2,3
Dryopidae
Diversity in Peru: 3 genera, 3+ species.
Recognition: All adult dryopids have short, pectinate antennae. Adult Dryops and
Onopelmus are covered with erect setae that float them to the surface if submerged.
Adult Elmoparnus have a shiny black exterior. Larvae are terrestrial and found in the
sand banks of streams; very few are known. Dryops occurs world-wide with many
species, including a host of undesecribed species. There is little external differentiation
and the species are generally distinguished with genitalic dissections.
Habitat: Some dryopids are aquatic and found in the stream substrates while
others occur on emergent vegetation and sticks protruding from streams. Many are
collected at lights at night; for these the habitat is unknown. A few dryopids are
entirely terrestrial with the adults and larvae found in forest leaf litter.
Notes: Since much of Peru has never been collected systematically, many more
genera and species will be found to occur there. The following is a checklist of
Peruvian Dryopidae, based on the publications of Spangler (1980) and Spangler and
Perkins (1977), and on personal collections. * 5 endemic to Peru.
Checklist:
Dryops undetermined species
Elmoparnus dasycheilus Spangler and Perkins, 1977
Onopelmus inca Spangler, 1980*
Limnichidae
Diversity in Peru: 6 genera, 13 species
Recognition: Adult limnichids are very small (24 mm), with a compact ovoid
body, with the head capable of being retracted into the thorax, and usually covered
with multicolored scale-like setae or short erect hair-like setae. Their antennae curl
around the eyes and are held close to the head. When disturbed, they withdraw their
legs into grooves in the exoskeleton. Very few larvae have ever been described but
they resemble elmid larvae and are found in the water miniscus around emergent
wood, or in the interstitial water of sandy banks of steam and lakes.
Habitat: Adults occur on the banks of streams and lakes and on vegetation
overhanging water bodies. They are often in abundance at lights at night. The few
larvae that have been found occur in sandy banks of water bodies or in well-rotted
wood protruding from the water.
1
Berkeley, California, USA, 94720.
2
Lawrence, Kansas, USA, 66045.
3
Corresponding author. Email: cschaboo@ku.edu
Accepted 20 February 2014; Revised 20 August 2015
Notes: The checklist below comes from Spangler et al. (2001) and Ribera and
Hernando (2000). Blackwelder (19441957) did not record the family from Peru. All
known Peruvian taxa belong to the Subfamily Limnichinae. Many more species are
anticipated to be found in Peru when it is better collected. * 5 endemic to Peru.
Checklist:
Byrrhinus magnus Wooldridge, 1987 Limnichoderus conspectus Wooldridge, 1981
Cephalobyrrhinus robustus Wooldridge, 1986 Limnichoderus similis Wooldridge, 1981*
Corrinea parva Wooldridge, 1980 Phalacrichus atomarius Sharp, 1902
Corrinea splendida Wooldridge, 1980 Phalacrichus duras Wooldridge, 1982
Eulimnichus ater (LeConte, 1854) Phalacrichus latus Wooldridge, 1982
Eulimnichus langleyae Wooldridge, 1979 Phalacrichus max Ribera & Hernando, 2000*
Eulimnichus plebius Sharp, 1902
Lutrochidae
Diversity in Peru: 1 genus, 5 species (including 4 undescribed).
Recognition: Lutrochids are recognized by their adaptations to fast-flowing
streams, which include a dense pelage of golden or black setae, compact antennae,
and long, last tarsomeres with large claws. Larvae are well sclerotized, compact and
have a terminal, ventral operculum hiding the tracheal gill tufts.
Habitat: Generally lutrochids occur in shallow rapids of clear streams, often in
association with submerged wood (adults and larvae). Some lutrochids are found in
leaf packs caught on objects projecting from the stream or on vegetation
overhanging streams and partially in the water. The species with golden pelage
often are associated with travertine deposits in streams.
Notes: The following is a list of Peruvian Lutrochidae. Blackwelder (19441957)
reported only the Hinton species, but four additional, undescribed species are known
to occur in Peru (C.A. Maier, personal communication).
Checklist:
Lutrochus gigas Hinton, 1939 Lutrochus undescribed sp. 3
Lutrochus undescribed sp. 1 Lutrochus undescribed sp. 4
Lutrochus undescribed sp. 2
Psephenidae
Recognition: Larval psephenids are flat, oval and brownish in color, hence the
common name of water penny beetles. Adults have long palps, soft elytra and dark
colors. Larvae cannot be distinguished beyond subfamily. Adult identifications to
the species level require genitalic dissections of males. However, some genera can be
recognized externally in adults.
Habitat: Larvae occur on rocks and in the gravel of stream substrates. Adults are
found on the exposed surfaces of rocks in riffles and on vegetation overhanging
streams. Adults often occur where the sun is sparkling on the irregular water surface
of riffles. Larvae feed on algae and detritus found on submerged rocks while adults
are short-lived and do not feed.
Notes: One species has been recorded from Peru (Spangler, 1966). This family was
not reported from Peru by Blackwelder (19441957). Recent collections have found
additional species, including at one site where there were 3 morpho-species of larvae
in the substrate. Because much of Peru has not been systematically collected many
more species should be found in the future. * 5 endemic to Peru.
Checklist:
Psephenus robacki Spangler, 1966* (actually belongs to the genus Psephenops)
Ptilodactylidae
Recognition: Adult ptilodactylids are recognized by their pronotum, which is
anteriorly narrowed and rounded and has a crenulate base, and by the serrate to
ramose antennae and cordate scutellum (Ivie, 2002). Larval ptilodactylids are long,
narrow wire-worms, with 10 visible abdominal segments, the tenth often is not
visible dorsally and it bears anal hooks and sometimes osmoregulatory papillae.
Aquatic larvae may also have tufts of anal osmobranichiae, gill-like structures which
may have an osmoregulatory function.
Habitat: Most species are entirely terrestrial; a few have aquatic larvae. Adults are
usually found on vegetation. Often they are abundant at lights at night. Larvae are
either in leaf litter and dead wood, or they are in streams and seeps where they feed
on dead leaves and rotting wood. The life cycle and biology of the aquatic
Anchytarsus palpalis is described by Spangler (1983)
Notes: The following checklist of Peruvian Ptilodactylidae was assembled using
a combination of Blackwelder (19441957) and Striblings (1986a) revision of
Anchytarus and his (1986b) unpublished dissertation. Blackwelder (19441957)
reported 26 species (and subspecies) from Peru.
Checklist:
Anchytarsus palpalis (Champion, 1897) Ptilodactyla luteitarsus Pic, 1947
Anchytarsus species B (Stribling, 1986a) Ptilodactyla minuta Kirsch, 1873
Aploglossa aureonotata Pic, 1913 Ptilodactyla nigra Kirsch, 1873
Aploglossa baeri Pic, 1913 Ptilodactyla nodieri Pic, 1947
Aploglossa collaris reducta Pic, 1934 Ptilodactyla obesa Erichson, 1847
Aploglossa collaris testaceicollis Pic, 1913 Ptilodactyla obscura Kirsch, 1873
Aploglossa testaceipes Pic, 1934 Ptilodactyla omospila Kirsch, 1873
Bradytoma lyciformis (Erichson, 1847) Ptilodactyla pallescens Kirsch, 1873
Chelonariomorphus undescribed sp. (Stribling, 1986b) Ptilodactyla peruviana Pic, 1923
New genus, undetermined sp. (Stribling, 1986b) Ptilodactyla peruviensis Pic, 1947
Ptilodactyla angusta Kirsch, 1874 Ptilodactyla praecellens Kirsch, 1873
Ptilodactyla atricolor Pic, 1947 Ptilodactyla probanda Kirsch, 1873
Ptilodactyla baeri Pic, 1916 Ptilodactyla punosa Pic, 1947
Ptilodactyla cisteloides Kirsch, 1874 Ptilodactyla scutellaris Kirsch, 1873
Ptilodactyla compressa Pic, 1947 Ptilodactyla secedens Kirsch, 1873
Ptilodactyla decumana Erichson, 1847 Ptilodactyla simulans Kirsch, 1873
Ptilodactyla heterophya Kirsch, 1873 Ptilodactyla tricoloricornis Pic, 1923
Ptilodactyla impressipennis Pic, 1916 Ptilodactyla truncaticollis Pic, 1947
Ptilodactyla lamellifera Kirsch, 1873 Ptilodactyla vilis Kirsch, 1873
Acknowledgements
Evolutionary Biology-General Research Fund (PI: CS Chaboo). We thank Crystal
Maier and Matthew Gimmel for help improving the manuscript.
Literature Cited
Indies, and South America, part 3. Bulletin of the United States National Museum 185:343550.
Ivie, M. A. 2002. Ptilodactylidae Laporte 1836. In Arnett, R. H., Jr., M. C. Thomas, P. E. Skelley, and J.
H. Frank (eds.). American Beetles. Vol. 2. Polyphaga: Scarabaeoidea through Curculionidea, pp.
135138. CRC Press, New York. 861 pp.
Ribera, I., and C. Hernando. 2000. Phalacrichus max n. sp. (Coleoptera, Limnichidae) from Peru
(America). Miscellania Zoologica 23:103106.
Spangler, P. J. 1966. The Catherwood Foundation Peruvian-Amazon Expedition. XIII Aquatic
Coleoptera (Dytiscidae; Noteridae; Gyrinidae; Hydrophilidae; Dascillidae; Helodidae; Psepheni-
dae; Elmidae). Monographs of the Academy of Natural Sciences of Philadelphia No. 14:377443.
Spangler, P. J. 1980. Onopelmus, a new genus of dryopid beetle from Peru (Coleoptera: Dryopidae).
Proceedings of the Entomological Society of Washington 82:6368.
Spangler, P. J. 1983. Immature stages and biology of Tetraglossa palpalis Champion (Coleoptera:
Ptilodactylidae). Entomological News 94:161175.
Spangler, P. J., and P. D. Perkins. 1977. Three new species of the Neotropical water beetle genus
Elmoparnus (Coleoptera: Dryopidae). Proceedings of the Biological Society of Washington
89:743760.
Spangler, P. J., C. L. Staines, P. M. Spangler, and S. L. Staines. 2001. A checklist of the Limnichidae and
the Lutrochidae (Coleoptera) of the world. Insecta Mundi 15:151165.
Stribling, J. B. 1986a. Revision of Anchytarsus (Coleoptera: Dryopoidea) and a key to the New World
genera of Ptilodactylidae. Annals of the Entomological Society of America 79:219234.
Stribling, J. B. 1986b. World Generic Revision of Ptilodactylidae (Coleoptera: Dryopoidea). Unpublished
dissertation, The Ohio State University, xv + 200 pp.
Beetles (Coleoptera) of Peru: A Survey of the Families. Elmidae

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88(2), 2015, pp. 167168
Beetles (Coleoptera) of Peru: A Survey of the Families. Elmidae

Diversity in Peru: 2 subfamilies, 17 genera, 51 species.

Recognition: These are commonly called riffle beetles because the larvae and adults
frequent running water habitats, aided by unusual respiratory structures. Adults
have a plastron and larvae have brancheal gills that open externally from under
a terminal operculum. Both adults and larvae have well developed tarsal claws.
Habitat: Elmids are associated with cool, well oxygenated, quickly flowing waters.
Both adults and larvae scrape algae off rocks for food; only a few genera consume
decaying wood in streams. Laraine adults are found in white-water areas where their
morphology catches submerged air bubbles for respiration. Elminae adults and the
larvae of both subfamilies depend on dissolved oxygen which diffuses into either the
plastron or the gills. Some adults and larvae are found in leaf-packs caught on emergent
objects in steams. Most elmids are found within the gravel and rocks that make up the
stream substrate. A heterogenous substrate supports a greater diversity of species.
Notes: Blackwelder (19441957) reported 12 species from Peru. More recent
citations include: Spangler (1966), Spangler and Santiago (1987), Spangler and
Perkins (1989), Dos Passos et al. (2010), and Segura et al. (2013). The checklist below
is derived from W. Shepard (personal communication). * 5 endemic to Peru.
Checklist:
Larainae
Disersus ambocheilus Spangler and Santiago, 1987 Hexanchorus tibialis Hinton, 1935
Disersus quincemil Spangler and Santiago, 1987* Phanocerus charopus Spangler, 1966*
Elminae
Austrelmis condimentarius (Philippi, 1864)* Cylloepus araneolus (Muller, 1806)*
Austrelmis confluenta (Hinton, 1940)* Cylloepus atys Hinton, 1945*
Austrelmis confusa (Hinton, 1940)* Cylloepus caicus Hinton, 1945*
Austrelmis consors (Hinton, 1940) Cylloepus consobrinus Grouvelle, 1896
Austrelmis dorotae Wiezlak, 1987* Cylloepus drymus Hinton, 1945*
Austrelmis gilsoni (Hinton, 1940) Cylloepus olenus Hinton, 1945
Austrelmis glabra (Hinton, 1940)* Cylloepus palpalis Hinton, 1937*
Austrelmis lata (Hinton, 1940)* Cylloepus silius Hinton, 1945*
Austrelmis peruana (Hinton, 1937) Cylloepus tuberculatus Hinton, 1940
Austrelmis steineri (Spangler, 1980)* Cylloepus ulpianus Hinton, 1945*
Austrelmis woytkowskii (Hinton, 1937) Heterelmis glabra (Horn, 1870)
Austrolimnius formosus (Sharp, 1882) Heterelmis obesa Sharp, 1882
Austrolimnius tarsalis Hinton, 1941 Heterelmis simplex Sharp, 1882
1
2
Lawrence, Kansas, 66045, USA, E-mail: cschaboo@ku.edu
3
Accepted 23 May 2015; Revised 25 August 2015
Heterelmis vilcanota Spangler, 1980* Microcylloepus steffani Bug, 1973*

Hexacylloepus nothrus Spangler, 1966 Neoelmis apicalis (Sharp, 1882)
Hintonelmis sandersoni Spangler, 1966* Notelmis bifoveolata Deleve, 1968
Macrelmis aeolis (Hinton, 1946) Onychelmis leleupi Deleve, 1968
Macrelmis amanus (Hinton, 1946) Stegoelmis andersoni Spangler, 1990
Macrelmis amazonica (Hinton, 1945) Stenhelmoides pubipes Spangler and Perkins,
Macrelmis graniger (Sharp, 1882) 1989
Macrelmis peruviana (Hinton, 1945)* Stenhelmoides rufulus (Hinton, 1934)
Macrelmis striata (Sharp, 1882) Stenhelmoides strictifrons (Grouvelle, 1908)
Microcylloepus angustus Hinton, 1940 Xenelmis leechi Perkins and Steiner, 1981*
Microcylloepus latus Bug, 1973* Xenelmis marcapata Perkins and Steiner, 1981*
Acknowledgements
We acknowledge NSF-EPSCoR # 66928 (PI: CS Chaboo) and the University of
Kansas Department of Ecology and Evolutionary Biology-General Research Fund
(PI: CS Chaboo) for supporting the Beetles of Peru project. We thank Cheryl Barr
and Matthew Gimmel for help assembling names. We also thank two anonymous
reviewers for helping to improve the manuscript.
Literature Cited
Indies, and South America. Parts 16. United States National Museum Bulletin 185:ixii, 11492.
Dos Passos, M. I. S., A. S. Fernandes, N. Hamada, and J. L. Nessimian. 2010. Insecta, Coleoptera,
Elmidae, Amazon region. CheckList 6(4):538545.
Segura, M. A., M. I. Dos Passos, A. A. Fonseca-Gessner, and C. G. Froehlich. 2013. Elmidae Curtis, 1830
(Coleoptera, Polyphaga, Byrrhoidea) of the Neotropical region. Zootaxa 3731(1):157.
Spangler, P. J. 1966. XIII Aquatic Coleoptera (Dytiscidae; Noteridae; Gyrinindae; Hydrophilidae;
Dascillidae; Helodidae; Psephenidae; Elmidae). In Patrick, R. et al. (eds.). The Catherwood
Foundation Peruvian-Amazon Expedition. Monographs of the Academy of Natural Sciences of
Philadelphia, No. 14:377443.
Spangler, P. J., and P. D. Perkins. 1989. A revision of the Neotropical aquatic beetle genus Stenhelmoides
(Coleoptera: Elmidae). Smithsonian Contributions to Zoology No. 79, 63 pp.
Spangler, P. J., and S. Santiago. 1987. A revision of the Neotropical aquatic beetle genera Disersus,
Pseudodisersus, and Potamophilops (Coleoptera: Elmidae. Smithsonian Contributions to Zoology
No. 446, 40 pp.
Epimetopidae, Hydrochidae, Hydrophilidae (Hydrophiloidea)

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88(2), 2015, pp. 169172
Beetles (Coleoptera) of Peru: A Survey of the Families. Epimetopidae,

Hydrochidae, Hydrophilidae (Hydrophiloidea)
The Superfamily Hydrophiloidea has varied greatly in its makeup over time. The
most recent decisions have restricted it to six families: Epimetopidae, Hydrochidae,
Georissidae, Helophoridae, Sperchidae, and Hydrophilidae. Three families have
known Peruvian distributions and are described below.
The family Georissidae is monogeneric and cosmopolitan in distribution.
Georissus humeralis Pic, 1923 is known from Argentina, Brazil, and Bolivia, and
a Georissus sp. is known from Chile. Thus, the family may occur in Peru and may be
found on sandy shores of lakes and streams.
Spercheidae is monogeneric and widely distributed globally. Two species occur in
South America: Spercheus fimbriicollis Bruch, 1915 from Brazil and Argentina and
Spercheus halophilus Archangelsky, 2001 from Argentina. Spercheus may be found in
Peru, especially in Amazonian areas where they would be found in stagnant waters.
The monogeneric Helophoridae has approximately 190 species that are largely
Holarctic in distribution. This family is not likely to be found in Peru.
Epimetopidae
Recognition: This small family contains three genera and ,30 described species
world-wide. Epimetopus is restricted to the New World. Epimetopids are small (14
mm), with a pronotum that is anteriorly broad with the middle overlapping the head
and posteriorly narrower than the elytra. The eyes are divided by a median canthus.
The first abdominal sternite is very short. The pronotum and elytra are often heavily
ornamented or sculptured.
Habitat: The ecology of epimetopids is poorly known. Some species have been
found associated with drying streams. Perkins (2012) notes that epimetopids are
associated with the margins of water bodies, often in the organic matter found there.
Notes: Epimetopids are found in the Peruvian Andes, but not yet in Amazonia
(Perkins, 2012). This may change with more and better collecting. Often adults are
collected at lights at night. The following checklist is taken from Perkins (2012).
Checklist:
Epimetopus angustus Perkins, 2012 Epimetopus peruvianus Perkins, 2012
Epimetopus inaequalis Perkins, 2012 Epimetopus spatulus Perkins, 2012
Epimetopus mendeli Fikacek, Barclay and Perkins, Epimetopus transversoides Perkins,
2011 2012
1
Berkeley, California, 94720.
2
Lawrence, Kansas, 66045, USA, E-mail: cschaboo@ku.edu
3
Accepted 12 May 2014; Revised 13 May 2015
Hydrochidae
Recognition: Long considered a subfamily of Hydrophilidae, this small,
cosmopolitan group of species is easily separated from typical hydrophilids by their
small size, narrow body, round bulging eyes, tiny scutellum, and pronotum which is
wider anteriorly. Today, about 200 species in Hydrochus are known world-wide.
Habitat: Hydrochids live mainly in stagnant or slowly flowing waters where they
are often found holding onto plants.
Notes: The following species is taken from the world catalogue of Hansen (1999).
Checklist:
Hydrochus pupillus dOrchymont, 1939
Hydrophilidae
Diversity in Peru: 5 subfamilies, 7 tribes, 17 genera, 47 species.
Recognition: Aquatic hydrophilids are usually recognized by their dark-colored
shiny oval bodies, short clubbed antennae and palps that are longer than the
antennae. The antennae have 79 antennomeres of which the last three form a club
which is preceded by a cup-like antennomere. Terrestrial hydrophilids have the
antennae longer than the palps and the first tarsomere of the hind tarsus is long.
Habitat: Most hydrophilids are aquatic as adults and larvae and common in
shallow still water. Some aquatic species inhabit vertical surfaces with sheet flow,
such as in seeps and splash zones near waterfalls. Peruvian species of the Subfamily
Sphaeridiinae is terrestrial and found in dung, carrion and decaying vegetation.
Notes: The following checklist of Peruvian Hydrophilidae is based on data from the
world catalogue of Hansen (1999), supplemented by Komarek (2005), Short and
Hebauer (2006), Fernandez et al. (2000), Fikacek (2009), Fikacek et al. (2009), Fikacek
and Vondracek (2014), and Short and Fikacek (2011). The classification follows
Short and Fikacek (2013). Hydrophilids readily come to lights which facilitates
collecting.
Checklist:
Hydrophilinae: Berosini
Berosus (Enoplurus) ghanicoides dOrchymont, Hemiosus hapalus Spangler, 1966
1941 Hemiosus hartmanni Mouchamps,
Berosus (Berosus) domitus Spangler, 1966 1957
Derallus argutus dOrchymont, 1940 Hemiosus maculatus Sharp, 1882
Hemiosus aequatorialis Oliva, 1994 Hemiosus morlestus dOrchymont, 1940
Hemiosus funditus Spangler, 1966 Hemiosus tarsalis Oliva and Short, 2010
Hydrophilinae: Laccobiini
Paracymus limbatus Wooldridge, 1973
Hydrophilinae: Hydrophilini
Hydrophilus (Dibolocelus) smaragdinus Brulle, Tropisternus (Pristoternus) robustus Sharp,
1837 1883
Hydrophilus (Hydrophilus) foveolatus (Regimbart, Tropisternus (Strepitornus) collaris (Fabricius,
1901) 1775)
Hydrophilus (Hydrophilus) guarani (Bachmann, Tropisternus (Tropisternus) bidentatus
1966) dOrchymont, 1922
Tropisternus (Pleurhomus) sahlbergi (Sharp, 1883) Tropisternus (Tropisternus) lateralis limbatus
Tropisternus (Pristoternus) baeri dOrchymont, (Brulle, 1837)
1921 Tropisternus (Tropisternus) setiger (Germar, 1824)
Chaetarthriinae: Chaetarthriini
?Chaetarthria panda dOrchymont, 1939
Chaetarthriinae: Anacaenini
Anacaena attigua dOrchymont, 1942 Crenitulus solstitialis (Kirsch, 1873)
Crenitulus punctata (Kirsch, 1873) Crenitulus suturalis (LeConte, 1866)
Enochrinae
Enochrus (Methydrus) lampros Knisch, 1924 Enochrus (Hugoscottia) orchymonti Mouchamps,
Enochrus (Methydrus) obscurus (Sharp, 1882) 1956
Enochrus (Methydrus) waterhousei Blair, Enochrus (Hugoscottia) peruvianus dOrchymont,
1933 1941
Acidocerinae
Quadriops depressus Hansen, 1999 Quadriops politus Hansen, 1999
Sphaeridiinae: Coelostomatini
Dactylosternum subdepressum subdepressum Dactylosternum subrotundum (Fabricius,
(Castelnau, 1840) 1792)
Dactylosternum subdepressum striatopunctatum Phaenostoma kontax Gustafson and Short, 2010
(Mulsant, 1844) Phaenostoma posticatum (Sharp, 1887)
Sphaeridiinae: Megasternini
Australocyon lescheni Hansen, 2003 Oosternum cicatricosum Fikacek, 2009
Australocyon peruvianus Hansen, 2003 Pelosoma brunneum (Kirsch, 1873)
Cercyon (Cercyon) variegatus Sharp, 1882 Sacosternum lebbinorum Fikacek and Short,
Cercyon (Paracercyon) proximus dOrchymont, 2010
1941 Sacosternum sp. A (Fikacek and Short, 2010)
Oosternum aequinoctiale (Motschulsky, 1855)
Acknowledgements
Evolutionary Biology-General Research Fund (PI: CS Chaboo) for funding this
publication. We thank Mathew Gimmel and Andrew Short for help assembling
species names and two anonymous reviewers for helping to improve the final
manuscript.
Literature Cited
Archangelsky, M. 2001. A new Neotropical species of Spercheus Kugelann, and its larval stages
(Coleoptera: Hydrophiloidea: Sperchidae). Studies on Neotropical Fauna and Environment
36:199204.
Indies, and South America. Parts 1 through 6. United States National Museum Bulletin 185:ixii,
11492.
Fernandez, L. A., A. O. Bachmann, and M. Archangelsky. 2000. Nota sobra Hydrophilidae neotropicales
(Coleoptera) II. Nuevos taxiones de Tropisternus. Revista de la Sociedad entomologica argentina
59:185197.
Fikacek, M. 2009. Taxonomic revision of the New World species of the genus Oosternum Sharp III. A new
species of the O. aequinoctiale species group from Costa Rica (Coleoptera: Hydrophilidae:
Sphaeridiinae). Koleopterologische Rundschau 79:179187.
Fikacek, M., F. Hebauer, and M. Hansen. 2009. Taxonomic revision of New World species of the genus
Oosternum Sharp (Coleoptera: Hydrophilidae: Sphaeridiinae) I. Definition of species groups and
revision of the Oosternum aequinoctiale group. Zootaxa 2054:137.
Fikacek, M., and D. Vondracek. 2014. A review of Pseudorygmodus (Coleoptera: Hydropphilidae), with
notes on the classification of the Anacaenini and on distribution of genera endemic to southern
South America. Acta Entomologica Musei Nationalis Pragae 54:479514.
Hansen, M. 1999. Hydrophiloidea (s.str.) (Coleoptera). World Catalogue of Insects 2:1416.
Komarek, A. 2005. Taxonomic revision of Anacaena Thomson, 1859. II. Neotropical species (Coleoptera:
Hydrophilidae) Koleopterologische Rundschau 75:253301.
Perkins, P. D. 2012. A revision of Epimetopus Lacordaire, the new world hooded shore beetles
(Coleoptera: Epimetopidae). Zootaxa 3531:195.
Short, A. E. Z., and M. Fikacek. 2011. World catalogue of the Hydrophiloidea (Coleoptera): additions
and corrections II (20062010). Acta Entomologica Musei Nationalis Pragae 51:83122.
Short, A. E. Z., and M. Fikacek. 2013. Molecular phylogeny, evolution and classification of the
Hydrophilidae (Coleoptera). Systematic Entomology 38:723752.
Short, A. E. Z., and F. Hebauer. 2006. World catalogue of Hydrophiloidea additions and corrections, 1
(19992005) (Coleoptera). Koleopterologische Rundschau 76:315359.
Histeridae
Author(s): Alexey K. Tishechkin and Nicolas Dgallier

terms_of_use.
critical research.
88(2), 2015, pp. 173179
Beetles (Coleoptera) of Peru: A Survey of the Families. Histeridae

ALEXEY K. TISHECHKIN1,3 AND NICOLAS DEGALLIER2

Recognition: Histeridae represent a monophyletic clade of staphyliniform beetles,
classified as part of Hydrophiloidea or a major component of Histeroidea,
a superfamily closely related to the former one. Histerids are small to medium-
sized beetles (0.825 mm, most are within 25 mm range), with solid compact oval or
sub-sphaerical convex bodies and short, robust, more or less flattened legs. When
disturbed, histerids fall into thanatosis, retracting the head and legs into specialized
cavities in pronotum and ventral sides of the body and remaining motionless for
minutes. Body colors are predominantly uniform in histerids, being almost
invariably uniformly brown or black, occasionally paler, yellowish or light brown,
or with metallic luster, while a very few histerids have pale spots or bands on elytra.
Deviations from the typical histerid body forms occur in some xylophilous and
myrmecophilous histerids. Extreme xylophilous specialists living under bark of dead
trees or in galleries of wood-boring insects may have the bodies, respectively,
strongly flattened (Histerinae: Hololeptini and some Exosternini, most Dendrophi-
linae: Paromalini in Peru) or almost cylindrical (Abraeinae: Teretriinae and
Trypanaeinae in Peru). Some histerid guests of army ants have extremely elongate,
stick-shaped femora and tibiae. These long legs are functional for the beetles in
running alongside the ants during emigrations and/or attaching themselves to worker
ants for riding during emigrations or acquiring colony odors. Myrmeco- and
termitophilous histerids also often have some other specialized structures, not found
in other histerids, e.g., strikingly developed setose body cover, trichomes, pronotal
and ventral projections and/or invaginations.
Diagnostic morphological features distinguishing adult histerids, along with
compact robust body shape, are: geniculate antennae with long scape and distinct
sphaerical or short oval 3-segmented club and slightly shortened elytra, leaving
uncovered the propygidium and pygidium (6th and 7th abdominal tergites).
Additional histerid diagnostic characters include special structures acting as locks
between elytral apices and 4th abdominal tergite, complete separation of procoxae
by prosternal keel, and hypognathous, fully retractable head.
Histerid larvae are relatively poorly known and rarely collected. They occur in the
same microhabitats as adults and have very fast development with only two instars
present. Histerid larvae may be recognized in the field as staphyliniform larvae with
relatively small and narrow heads, and with 2-segmented urogomphi.
Habitat: Histeridae occur primarily in lowland, premontane and lower montane
forests. However, all the habitats in Peru will harbor some Histeridae, and such
1
Systematic Entomology Laboratory USDA, c/o National Museum of Natural History, MRC-168,
P.O. Box 37012, Washington, DC 20013-7012, USA
2
120 rue de Charonne, 75011 Paris, France.
3
Corresponding Author E-mail: atishe8@gmail.com
Accepted 29 May 2014; Revised 25 November 2014
extreme habitats as open deserts, sea beaches and alpine scrubs and grasslands above
3000 m have some specialized histerid species. Adults of most histerids are predators,
both as larvae and adults, and mostly feed on soft-bodied insect larvae (e.g., Diptera,
Siphonaptera), and also on mites and springtails. Several genera in Abraeinae, Den-
drophilinae and Tribalinae are known or strongly suspected to feed on microfungi in
rotten wood, subcortical spaces of decaying trees and forest litter.
Most of the histerids are associated with some kind of decomposing organic
matter, e.g., animal dung, carrion, rotting vegetation and forest litter. Substantial
numbers of histerid species have an inquilinous life style and are regular and diverse
inhabitants of vertebrate nests and burrows and colonies of social insects, primarily
ants, but also termites and social bees. Also, there are numerous and often abundant
specialized inhabitants of subcortical spaces of dead trees and galleries of wood-
boring insects. Finally, a few histerids are known to be sand dune and sea beach
dwellers, where they live in rhizospheres of dune plants and under decomposing dead
animals and beached algae. The family is known to be poorly represented at higher
elevations (above 20002500 m in Peru) and in forest canopies, but there is evidence
of the existence of specialized histerid communities in both habitats.
Histerids are readily collected by direct inspection of their prime habitats and
niches, but passive trapping, baited pitfall traps and especially flight intercept traps,
are productive methods to collect these beetles. Most histerids are not attracted by
light traps, although a few taxa occasionally are collected by this method, especially
in more open habitats.
Notes: The following checklist was derived from Erichson (1847), Blackwelder
(1944), and Mazur (2011), and supplemented by Degallier et al. (2012), Leivas et al.
(2012), Caterino and Tishechkin (2013a, b, 2014), Moura and Almeida (2013).
Seventeen species are reported here as new records for Peru:
Degallierister barberoi Gomy, 2001: Cuzco, Cock of the Rock Lodge, NE

Paucartambo, 13u03.39S, 71u32.79W, 1120 m, 4-9-XI-2007, flight intercept trap, D.
Brzoska. Junn, Pampa Hermosa Lodge, 22 km N San Ramon, 10u59.39S,
75u25.59W, 1220 m, flight intercept trap, 24-27-XI-2007, D. Brzoska (SEMC).
Degallierister hielkemai Gomy, 2013: Loreto, Teniente Lopez, 2u35.669S,

76u06.929W, 23-VII-1993, flight intercept trap, R. Leschen (SEMC); El Porvenir,
45 km W Iquitos, Rio Nanay, 3u549020S, 73u339250W, flight intercept trap, 10-VII-
2011, G. Lamarre (CHND); same locality, flight intercept trap, 5-VII-2011, G.
Lamarre (CHND); same locality at 3u549S, 73u339W, flight intercept trap, VII-2011,
G. Lamarre (CHND); Rio Uyacali, Jenaro, 145 km SW Iquitos, terra firme, 4u549S,
73u40W, 100 m, flight intercept trap, VIII-2011, G. Lamarre (CHND); same locality,
flight intercept trap, 26-VII-2011, G. Lamarre (CHND); same locality, flight
intercept trap, 31-VII-11, G. Lamarre (CHND). Madre de Dios, Pantiacolla Lodge,
5.5 km NW El Mirador Trail, Alto Madre de Dios River, 12u399100S, 71u159280W,
500 m, 23-26-X-2000, flight intercept trap, R. Brooks (SEMC); same locality at
12u389300S, 71u169410W, 800 m, R. Brooks (SEMC); same locality, Alto Madre de
Dios River, 12u39.39S, 71u13.99W, 420 m, 14-19-XI-2007, flight intercept trap, D.
Brzoska (SEMC); CICRA Field Station, Experimental Plot, South Transect,
12.55261uS 70.11008W, 295 m, flight intercept trap, 9-11-VII-2010, C. Chaboo
team (SEMC); same locality, 11-13-VII-2010, C. Chaboo team (SEMC); same
locality, North Transect, 9-11-VII-2010, C. Chaboo team (SEMC); same locality,

11-13-VII-2010 (SEMC); same locality, 13-15-VII-2010, C. Chaboo team (SEMC).
Degallierister surinamensis Gomy, 2013: Loreto, El Porvenir, 45 km W Iquitos, Rio
Nanay, 3u549020S, 73u339250W, flight intercept trap, 10-VII-2011, G. Lamarre
(CHND); Rio Uyacali, Jenaro; 145 km SW Iquitos, flight intercept trap, 23-VIII-
2011 G. Lamarre (CHND).
Euspilotus azurescens (Marseul, 1855): Junn, left bank of Rio Tambo ca. 10 km E
Puerto Ocopa, lower Rio Shima, 6-16-I-2006, A. V. Petrov (LSAM). Loreto, 68 road
km from Iquitos to Nauta, 4-II-2007, A. V. Petrov (LSAM).
Hister diadema Marseul, 1854: Junn, ca. 5 km NNW Satipo, Rio Negro, 800 m,
2-III-2005, A.V.Petrov (LSAM); ca. 4 km NE Puerto Ocopa, Santa Cruz, 11u6.809S,
74u16.989W, 1150 m, 12-I-2006, A.V.Petrov (LSAM). Loreto, Rio Ucayali, Jenaro,
145 km SW Iquitos, 4u549S,73u409W, 100 m, terra firme, flight intercept trap, 26-VII-
2011, G. Lamarre (CHND).
Eutidium dentifrons (Desbordes, 1916): Loreto, Porvenir, Rio Nanay, terra firme,
flight intercept trap, 14-VI-2011, G. Lamarre (CHND).
Eutidium williamsi (Dillon, 1935): Loreto, Porvenir, Rio Nanay, flight intercept trap,
31-VII-2011, G. Lamarre (CHND).
Hololepta devia Marseul, 1853: Loreto, Porvenir, Rio Nanay, flight intercept trap,
28-VIII-2011, G. Lamarre (CHND).
Hololepta punctulata (Marseul, 1853): Loreto, Porvenir, Rio Nanay, flight intercept
trap, 14-VI-2011, G. Lamarre (CHND).
Hololepta quadridentata (Olivier, 1789): Loreto, Porvenir, Rio Nanay, flight
intercept trap, 10-VII-2011, G. Lamarre (CHND).
Hololepta reichii (Marseul, 1853): Loreto, Porvenir, Rio Nanay, flight intercept trap,
14-VI-2011, G. Lamarre (CHND).
Omalodes bifoveolatus Marseul, 1854: Cusco, Villa Carmen Research Station at
12.55236uS 70.10989uW, 295 m, flight intercept trap, 11-13-VI-2011, C. Chaboo
Team (SEMC); same locality at 12.89250uS, 71.41915uW, 555 m, flight intercept
trap, 22-24-V-2011, D. J. Bennet & E. Razuri (SEMC); same locality at 12.89221uS,
71.41946uW, 557 m, flight intercept trap, 20-22-V-2011, D. J. Bennet & E. Razuri
(SEMC).
Omalodes anthracinus Marseul, 1854: Junn, Satipo, X-XII-1935, F. Waytkowski
(CHND).
Operclipygus quadratus Caterino & Tishechkin, 2013: Cusco, Villa Carmen Field
Station, ,1.7 km W cafeteria at 12.89250uS, 70.10989uW, 555 m, flight intercept
trap, 22-24-V-2011, D. J. Bennett & E. Razuri (SEMC).
Oxysternus maximus (L., 1767): Junn, ca. 7.5 km NE Puerto Ocopa, Cananeden,
11u4.929S, 74u16.109W, 1185 m, 2-II-2006, A. V. Petrov (LSAM). Loreto, ca. 20 km
NW Iquitos, Gen Gen on Rio Momon, 5-6.II.2007, A. V. Petrov (LSAM).
Terapus marseuli Westwood, 1874: Madres de Dios, Rio Los Amigos, CICRA Field
Station, 700 m, flight intercept trap, 17-XI-2006, A. Asenjo (MUSM).
Studies on the material in several major collections have proved the presence in Peru
of unidentified and/or undescribed species of the following genera: Acritus LeConte,
Aeletes Horn and Trypolister Bickhardt (Abraeinae), Anapleus Horn, Degallierister
Gomy and Bacanius Le Conte (Dendrophilinae), Aemulister Reichensperger,
Alienodites Tishechkin, Alloiodites Reichensperger, Aristomorphus Lewis, Cachexia
Lewis, Convivister Reichensperger, Cyclechinus Bickhardt, Daitrosister Helava,
Daptesister Helava, Euclasea Lewis, Eurysister Helava, Fistulaster Helava, Helavadites
Tishechkin, Hemicolonides Reichensperger, Homalopygus Boheman, Iugulister Rei-
chensperger, Leptosister Helava, Metasynodites Reichensperger, Mesynodites
Reichrdt, Nevermannister Reichensperger, Nymphister Reichensperger, Opadosister
Helava, Plaumannister Reichensperger, Psalidister Reichensperger, Pulvinister Rei-
chensperger, Reninus Lewis, Scapicoelis Marseul, Sternocoelopsis Reichensperger,
Symphilister Reichensperger, Synoditinus Reichensperger, Synoditulus Reichensper-
ger, Terapus Marseul, Termitoxenus Schmidt, Thaumataerius Mann, Trichoreninus
Lewis, Troglosternus Bickhardt, Tylois Marseul and Voratister Helava (Haeteriinae),
Hypobletus Schmidt and Yarmister Wenzel (Histerinae), Peploglyptus LeConte
(Onthophilinae), Euspilotus Lewis (Saprininae), Epierus Erichson (Tribalinae),
Coptotrophis Lewis and Xylonaeus Lewis (Trypanaeinae).
Records of Operclipygus dubius (Lewis), O. hospes (Lewis) (Mazur, 2011) and
Troglosternus ecitonis Mann (Tishechkin, 2003; Mazur, 2011) are based on incorrect
identifications (Caterino and Tishechkin, 2013b; Tishechkin, unpubl.) and so these
species are not included in the list.
Checklist:
Abraeinae
Acritini
Halacritus instabilis (Marseul, 1869)
Teretriini
Teretrius peruanus Erichson, 1847
Dendrophilinae
Bacaniini
Degallierister barberoi Gomy, 2001 new record Degallierister peruvianus Mazur, 1988
Degallierister hielkemai Gomy, 2013 new record Degallierister surinamensis Gomy, 2013 new record
Paromalini
Carcinops assimilis Wenzel, 1944 Paromalus leleupi Wenzel, 1976
Carcinops exigua Wenzel, 1944 Paromalus oculipygus Marseul, 1870
Carcinops peruviana Kirsch, 1873
Haeteriinae
Cheilister lucidulus Reichensperger, 1924
Paratropinus scalptus Reichensperger, 1935
Terapus marseuli Westwood, 1874 new record
Histerinae
Exosternini
Baconia aenea Caterino & Tishechkin, 2013 Operclipygus impressifrons Caterino & Tishechkin,
Baconia angulifrons Caterino & Tishechkin, 2013 2013
Baconia clemens Caterino & Tishechkin, 2013 Operclipygus impressistrius Caterino & Tishechkin,
Baconia dentipes Caterino & Tishechkin, 2013 2013
Baconia dives (Marseul, 1861) Operclipygus impuncticollis (Hinton, 1935)
Baconia emarginata Caterino & Tishechkin, 2013 Operclipygus inflatus Caterino & Tishechkin 2013
Baconia gibbifer Caterino & Tishechkin, 2013 Operclipygus juninensis Caterino & Tishechkin,
Baconia incognita Caterino & Tishechkin, 2013 2013
Baconia lescheni Caterino & Tishechkin, 2013 Operclipygus kerga (Marseul, 1870)
Baconia mustax Caterino & Tishechkin, 2013 Operclipygus lama Mazur, 1988
Baconia plebeia Caterino & Tishechkin, 2013 Operclipygus latipygus Caterino & Tishechkin,
Baconia punctiventer Caterino & Tishechkin, 2013 2013
Baconia subtilis Caterino & Tishechkin, 2013 Operclipygus longidens Caterino & Tishechkin,
Baconia tenuipes Caterino & Tishechkin, 2013 2013
Baconia tuberculifer Caterino & Tishechkin, 2013 Operclipygus olivensis Caterino & Tishechkin, 2013
Conchita propygidiale Mazur, 1994 Operclipygus panamensis (Wenzel & Dybas, 1941)
Crenulister dentatus Caterino & Tishechkin, 2014 Operclipygus parallelus Caterino & Tishechkin,
Crenulister explanatus Caterino & Tishechkin, 2013
2014 Operclipygus peregrinus Caterino & Tishechkin,
Crenulister impar Caterino & Tishechkin, 2014 2013
Kaszabister ferrugineus (Kirsch, 1873) Operclipygus petrovi Caterino & Tishechkin, 2013
Operclipygus angustisternus (Wenzel, 1944) Operclipygus plicicollis (Schmidt, 1893)
Operclipygus arnaudi Degallier, 1982 Operclipygus profundipygus Caterino & Tishech-
Operclipygus bickhardti Caterino & Tishechkin, kin, 2013
2013 Operclipygus prolixus Caterino & Tishechkin, 2013
Operclipygus brooksi Caterino & Tishechkin, 2013 Operclipygus punctipleurus Caterino & Tishechkin,
Operclipygus callifrons Caterino & Tishechkin, 2013
2013 Operclipygus pygidialis (Lewis, 1908)
Operclipygus communis Caterino & Tishechkin, Operclipygus quadratus Caterino & Tishechkin,
2013 2013 new record
Operclipygus confertus Caterino & Tishechkin, Operclipygus rupicolus Caterino & Tishechkin,
2013 2013
Operclipygus confluens Caterino & Tishechkin, Operclipygus schlingeri Caterino & Tishechkin,
2013 2013
Operclipygus conquisitus (Lewis, 1902) Operclipygus siluriformis Caterino & Tishechkin,
Operclipygus crenatus (Lewis, 1888) 2013
Operclipygus depressus (Hinton, 1935) Operclipygus simplicipygus Caterino & Tishechkin,
Operclipygus disconnectus Caterino & Tishechkin, 2013
2013 Operclipygus tiputinus Caterino & Tishechkin,
Operclipygus distinctus (Hinton, 1935) 2013
Operclipygus dytiscoides Caterino & Tishechkin, Operclipygus validus Caterino & Tishechkin, 2013
2013 Phelister bipulvinatus Marseul, 1854
Operclipygus ecitonis Caterino & Tishechkin, 2013 Phelister bistriatus Hinton, 1935
Operclipygus extraneus Caterino & Tishechkin, Phelister brevis Bickhardt, 1917
2013 Phelister carinifrons Schmidt, 1893
Operclipygus falini Caterino & Tishechkin, 2013 Phelister condor Mazur, 1988
Operclipygus fossipygus (Wenzel, 1944) Phelister egincola Marseul, 1889
Operclipygus foveipygus (Bickhardt, 1918) Phelister erraticus Marseul, 1887
Operclipygus hirsutipes Caterino & Tishechkin, Phelister fractistrius Lewis, 1908
2013 Phelister latus Wenzel & Dybas, 1941
Operclipygus ignifer Caterino & Tishechkin, 2013 Phelister praecox (Erichson, 1847)
Operclipygus impositus Caterino & Tishechkin, Phelister pusio (Erichson, 1847)
2013 Scaptorus pyramus Caterino & Tishechkin, 2014
Histerini
Hister akatanga Caterino, 1999 Hister denysi Marseul, 1870

Hister cavifrons Marseul 1854 Hister diadema Marseul, 1854 new record
Hister curvatus Erichson, 1834 Hister incisifrons Marseul, 1863
Hololeptini
Eutidium dentifrons (Desbordes, 1916) new record Hololepta minuta Erichson, 1834
Eutidium peruanum Lewis, 1907 Hololepta patula (Lewis, 1906)
Eutidium williamsi (Dillon, 1935) new record Hololepta punctulata (Marseul, 1853) new record
Hololepta andicola Schmidt, 1893 Hololepta quadridentata (Olivier, 1789) new record
Hololepta devia Marseul, 1853 new record Hololepta reichii (Marseul, 1853) new record
Hololepta guidonis Marseul, 1860 Hololepta truxillana Marseul, 1860
Hololepta intersecta (Lewis, 1904) Oxysternus maximus (Linnaeus, 1767) new record
Omalodini
Omalodes amazonius Marseul, 1861 Omalodes lucidus peruvianus Marseul, 1861

Omalodes anthracinus Marseul, 1854 Omalodes mazuri Moura & Almeida, 2013
Omalodes bifoveolatus Marseul, 1854 Omalodes mestino Lewis, 1904
new record Omalodes pulvinatus Erichson, 1834
Omalodes foveola Erichson, 1834 Scapomegas auritus Marseul, 1855
Omalodes gagatinus Erichson, 1847 Sphyracus peruanus Bickhardt, 1914
Saprininae
Euspilotus amazonius Desbordes, 1923 Euspilotus lepidus (Erichson, 1847)

Euspilotus azurescens (Marseul, 1855) new record Phoxonotus lectus Lewis, 1902
Euspilotus decoratus (Erichson, 1834) Saprinus caerulescens (Hoffmann, 1803)
Euspilotus excavata Arriagada in Degallier Satrapister nitens Bickhardt, 1912
et al. 2012 Xerosaprinus chiliensis (Marseul, 1855)
Euspilotus flaviclava (Marseul, 1870)
Tribalinae
Epierus planulus Erichson, 1834 Plagiogramma hastata (Marseul, 1855)

Idolia punctisternum Lewis, 1885 Plagiogramma marseulii (Kirsch, 1873)
Idolia scitula Lewis, 1888 Plagiogramma mundus (Erichson, 1834)
Plagiogramma arciger (Marseul, 1855) Plagiogramma peruana (Schmidt, 1889)
Trypanaeinae
Trypanaeus ensifer Marseul, 1856

Trypanaeus volvulus Erichson, 1834
Acknowledgements
We are grateful to Matthew Gimmel for initiating the species list. We wish to
extend our gratitude to Angelico Asenjo, Caroline Chaboo, Zack Falin, Greg
Lamarre, Alexander Petrov and Jennifer Thomas, who provided some recent
Peruvian histerid material for our study. We acknowledge NSF-EPSCoR #66928 (to
Caroline S. Chaboo) for supporting the Beetles of Peru project and the University
of Kansas Ecology Evolutionary Biology-General Research Fund (to Chaboo) for
funding this publication.
Literature Cited
Caterino, M. S., and A. K. Tishechkin. 2013a. A systematic revision of Baconia Lewis (Coleoptera,
Histeridae, Exosternini). ZooKeys 343:1297.
Caterino, M. S., and A. K. Tishechkin. 2013b. A systematic revision of Operclipygus Marseul (Coleoptera:
Histerdiade: Exosternini). ZooKeys 271:1401.
Caterino, M. S., and A. K. Tishechkin. 2014. New genera and species of Neotropical Exosternini
(Coleoptera, Histeridae). ZooKeys 381:1178.
Degallier, N., G. Arriagada, M. S. Caterino, P. Kanaar, D. P. Moura, A. K. Tishechkin, and W. B.
Warner. 2012. Coleoptera Histeridae de Guyane. VII. Complements au catalogue avec des donnees
sur la faune du Surinam et une contribution a la connaissance des Saprininae. In J. Touroult (ed.).
Contribution a lEtude des Coleopteres de Guyane. Tome VI. Supplement au Bulletin de Liaison
dA.C.O.R.E.P., pp.3352. A.C.O.R.E.P., Paris (France). 88 pp.
Erichson, W. F. 1847. Conspectus Insectorum Coleopterorum quae in Republica Peruana observata sunt.
Archiv fur Naturgeschichte 13:67185.
Leivas, F. W. T., C. L. Bicho, N. Degallier, and D. P. Moura. 2012. Revision of the genus Scapomegas
Lacordaire (Coleoptera: Histeridae: Omalodini). Zootaxa 3482:3346.
Mazur, S. 2011. A concise catalogue of the Histeridae (Coleoptera). Warsaw University of Life Sciences
S.G.G.W. Press, Warsaw (Poland). 332 pp.
Moura, D. P., and L. M. Almeida. 2013. Three new species of Omalodes (Omalodes) from South America.
ZooKeys 335:8599.
Tishechkin, A. K. 2003. New distribution records of Neotropical Hetaeriinae (Coleoptera: Histeridae).
Sociobiology 41:673683.
Hydraenidae
Author(s): Philip D. Perkins and Caroline S. Chaboo

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88(2), 2015, pp. 180181
Beetles (Coleoptera) of Peru: A Survey of the Families. Hydraenidae

PHILIP D. PERKINS1 AND CAROLINE S. CHABOO2,3

Recognition: Hydraenids, commonly called minute moss beetles, may be
recognized by their small size (generally ,3 mm), very short nine-segmented
antennae with a five-segmented club, and six or seven abdominal ventrites.
Habitat: These beetles are usually found along margins of streams (especially
sandy streams), but sometimes along the margins of lentic habitats (including rock
pools). Adults often live interstitially among sand grains and gravel, or occupy the
region just below the water surface along the shore or on half-submerged rocks.
Most species can be collected by the shore washing method, in which water is
splashed on the shoreline, and flows back into the stream or pond. Beetles are
dislodged from the sand and gravel, and float to the surface, where they can be
collected with a fine mesh net. Standard aquatic nets often have meshes that are too
large, and the hydraenids pass through.
Notes: The following list of Hydraenidae is derived from Perkins (1980, 2011).
Blackwelder (19441957) did not list any species for Peru. There are certainly more
species of Hydraenidae to be discovered in Peru.
Checklist:
Hydraena multispina Perkins, 2011
Gymnochthebius bartyrae Perkins, 1980
Hydraena novacula Perkins, 2011
Gymnochthebius peruvianus (J. Balfour-
Browne, 1971) Hydraena pavicula Perkins, 1980
Hydraena amazonica Perkins, 2011 Hydraena peru Perkins, 1980
Hydraena cordispina Perkins, 2011 Hydraena scintillamima Perkins, 2011
Hydraena curvosa Perkins, 2011 Hydraena scintillarca Perkins, 2011
Hydraena d-concava Perkins, 2011 Hydraena tridigita Perkins, 2011
Acknowledgements
Evolutionary Biology-General Research Fund (PI: CS Chaboo). We thank Matthew
Gimmel for help improving the manuscript.
Literature Cited
11492.
1
Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA, 02138.
2
3
Corresponding author, E-mail: cschaboo@ku.edu
Accepted 28 August 2014; Revised 03 August 2015
Perkins, P. D. 1980. Aquatic beetles of the family Hydraenidae in the Western Hemisphere: classification,
biogeography and inferred phylogeny (Insecta: Coleoptera). Quaestiones Entomologicae 16:3554.
Perkins, P. D. 2011. New records and description of fifty-four new species of aquatic beetles in the genus
Hydraena Kugelann from South America (Coleoptera: Hydraenidae). Zootaxa 3074:1198.
Beetles (Coleoptera) of Peru: A Survey of the Families. Ptiliidae
Heer, 1843
Author(s): Michael Darby and Caroline S. Chaboo

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88(2), 2015, pp. 182183
Beetles (Coleoptera) of Peru: A Survey of the Families. Ptiliidae

Heer, 1843
MICHAEL DARBY1 AND CAROLINE S. CHABOO2,3

Recognition: Members of Ptiliidae may be recognized by their extremely small size
(usually in the 0.51.5 mm range), the clubbed antennae whose antennomeres each
contain a whorl of long apical setae, and the feather-like hind wings (strap-like
membrane with fringe of extremely long hairs). The elytra often incompletely cover
the abdomen.
Habitat: Ptiliidae occur most often in moist, decaying organic matter (rotten
wood, mammal nests, dung, tree holes, seaweed piles, leaf litter) or fungi. Some
species are adapted for life in ant or termite nests.
Notes: Apparently only one species has been recorded from Peru (from Hall,
2003). These poorly-studied beetles are almost certainly very diverse in Peru. In our
survey of the arthropod fauna of phytotelmata Zingiberales plants (Jalinsky et al.,
2014), we found ptiliid adults accounting for 47% of the total adult beetle fauna in
the floral phytotelmata of Calathea and Heliconia plants. Here we report Acrotrichis
as an additional genus: PERU: Madre de Dios Dept., CICRA Field Station, ,50 m
W on Tr. 1, 12.569211uS, 70.100261uW, 27 m, 9.VI.2011, ex leaf roll, Heliconia
stricta, Jalinksy, Radocy, Wertenberger, PER-11-JJ-007 [1, SEMC].
Checklist:
Acrotrichinae
Acrotrichis undetermined species, New Record
Ptiliinae
Limulopteryx loebli Hall, 2003
Acknowledgements
publication. We thank Matthew Gimmel for help assembling species names.
Literature Cited
Hall, W. E. 2003. Limulopteryx, a new genus of Neotropical featherwing beetles (Coleoptera:
Staphylinoidea:Ptiliidae) and comments on pterycine ptiliids. In G. Cuccodoro and R. A. B.
Leschen (eds.). Systematics of Coleoptera: Papers Celebrating the Retirement of Ivan Lobl.
1
Department of Life Sciences (Entomology), Natural History Museum, London SW7 5BD.
2
3
Corresponding author, e-mail: cschaboo@ku.edu
Memoirs on entomology international, Vol. 17, pp. 85102. Associated Publishers, Gainesville,
FL. 955 pp.
Jalinsky, J., C. S. Chaboo, T. A. Radocy, and R. Wertenberger. 2014. Insect diversity in phytotelmata
habitats of two host plants, Heliconia stricta Huber Heliconiaceae) and Calathea lutea Schult
(Marantaceae) in the south-east Amazon of Peru. J. Kans. Entomol. Soc. 87:299311.
Silphidae
Author(s): Derek S. Sikes and Caroline S. Chaboo

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88(2), 2015, pp. 184185
Beetles (Coleoptera) of Peru: A Survey of the Families. Silphidae

DEREK S. SIKES1 AND CAROLINE S. CHABOO2,3

Recognition: Silphidae may be recognized by their large size (usually 1220 mm
long), conical procoxae, either truncate or costate elytra, and clubbed antennae
(lamellate in Nicrophorinae) (Sikes, 2005). Other characters, images, and citations
are available online at Sikes et al. (2005)
Habitat: Members of Silphidae are found associated with vertebrate carcasses, at
lights, or dung. Those in the genus Nicrophorus bury small carcasses in order to rear
larvae. Some species breed communally and others show biparental care (Eggert and
Muller, 1997).
One of the simplest ways to sample silphids is to bait containers with rotted fish or
chicken, then nail each container to a tree about chest height at dusk. Most silphids
fly at sundown so the majority of the nights catch will be there within an hour.
Notes: The list below is derived from Peck and Anderson (1985). Ten species and
varieties were reported in Blackwelder (19441957), but five have since been
relegated to synonymy. Some species, e.g., Oxelytrum erythrurum, are known from
single Peru records so these might be outliers.
Checklist:
Nicrophorinae
Nicrophorus didymus Brulle, 1840
Nicrophorus scrutator Blanchard, 1840
Silphinae
Oxelytrum anticola (Guerin-Meneville, 1855)
Oxelytrum cayennense (Sturm, 1826)
Oxelytrum discicolle (Brulle, 1840)
Oxelytrum erythrurum (Blanchard, 1840)
Acknowledgements
We acknowledge acknowledge NSF-EPSCoR #66928 (PI: CS Chaboo) for
supporting the Beetles of Peru project and the University of Kansas Department of
Ecology and Evolutionary Biology-General Research Fund (PI: CS Chaboo) for
funding this publication. We thank Matthew Gimmel for his help assembling some
species names.
1
University of Alaska Museum, 907 Yukon Dr., University of Alaska, Fairbanks, Alaska, USA 99775.
2
Lawrence, Kansas, USA 66045.
3
Accepted 27 May 2014; Revised 9 July 2015
Literature Cited
Indies, and South America. Parts 1 through 6. U. S. Natl. Mus. Bull., 185:ixii, 11492.
Eggert, A. -K., and J. K. Muller. 1997. Biparental care and social evolution in burying beetles: Lessons
from the larder. In J. C. Choe and and B. J. Crespi (eds.), Social Behavior in Insects and
Arachnids, pp. 216236. Cambridge University Press, Cambridge, New York & Oakleigh., vxiii,
1541.
Peck, S. B., and R. S. Anderson. 1985. Taxonomy, phylogeny and biogeography of the carrion beetles of
Latin America (Coleoptera: Silphidae). Quaest. Entomol. 21:247317.
Sikes, D. S. 2005. Silphidae. In Kristensen, N. P., and R. G. Beutel (eds.). Handbook of Zoology, Vol. IV
Arthropoda: Insecta Part 38, Coleoptera, Beetles Vol I: Morphology and Systematics
(Archostemmata, Adephaga, Myxophaga, Polyphaga partim) (Beutel, R. G., and R. A. B.
Leschen, eds.), pp. 288296. Walter de Gruyter, Berlin, New York xi + 567 pp.
Sikes, D. S., S. T. Trumbo, and S. B. Peck. 2005. Silphidae Latreille 1807. Large carrion and burying
beetles. Version 07 February 2005. http://tolweb.org/Silphidae [Accessed 27 May 2014].
Scarabaeoidea
Author(s): B. C. Ratcliffe, M. L. Jameson, L. Figueroa, R. D. Cave, M. J.
Paulsen, Enio B. Cano, C. Beza-Beza, L. Jimenez-Ferbans, and P. Reyes-Castillo

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88(2), 2015, pp. 186207
Beetles (Coleoptera) of Peru: A Survey of the Families. Scarabaeoidea

B. C. RATCLIFFE,1,10 M. L. JAMESON,2 L. FIGUEROA,3 R. D. CAVE,4 M. J. PAULSEN,5
ENIO B. CANO,6 C. BEZA-BEZA,7 L. JIMENEZ-FERBANS,8 AND P. REYES-CASTILLO9
ABSTRACT: The 1042 species of Scarabaeoidea known to occur in Peru are listed with their
taxonomic placement in families, subfamilies, and tribes.
Peru is known for high species richness, endemism, and habitat complexity, but
despite this interest in Peruvian biodiversity, little is known about the beetle fauna of the
country (Larsen et al., 2011). The following checklist reflects our most current
knowledge about the composition of the superfamily Scarabaeoidea in Peru. Compared
with the neighboring country of Chile in which 247 species and 66 genera of scarabaeoid
beetles are recorded (Elgueta, 2000), we record 1045 species of Peruvian scarabaeoid
beetles, but we know the number will increase with additional exploration and discovery.
This checklist was assembled by compiling data from print and online catalogs,
the primary literature that describes new species or reviews genera, some research
collections, and personal communications from some entomologists who have
personally conducted research in Peru. Clearly, there are other species of
scarabaeoids that are not yet recorded from Peru because their data have never
been included in the literature. Conversely, there may be a few species recorded from
Peru in the older catalogs that do not actually occur in Peru because they were
incorrectly identified. Moreover, disparate levels of knowledge exist between scarab
groups because some have undergone considerable study while others have not.
In the lists below, an asterisk (*) indicates a species that is known only from Peru.
Based upon our data, there are large numbers of scarabs endemic to Peru. This
suggests that the scarab beetle fauna of Peru is amazingly rich, and that further
exploration and discovery will reveal many more species. Scarab beetles are an
important component of Peruvian biodiversity because they are important
pollinators of plants, decomposers of waste, economically important pests, and
some are valuable culturally for adornment or as a food source. Knowledge about
1
Systematics Research Collections, University of Nebraska, Lincoln, Nebraska 68588-0514, USA; e-mail:
bratcliffe1@unl.edu.
2
Department of Biological Sciences, Wichita State University, Wichita, Kansas 67260-0026, USA;
e-mail: Maryliz.jameson@gmail.com.
3
Departmento de Entomologa, Museo de Historia Natural de la UNMSM, Lima, PERU; e-mail:
luis_thecell@hotmail.com.
4
Indian River Research and Education Center, University of Florida, Fort Pierce, Florida 34945-3138,
USA; e-mail: rdcave@ufl.edu.
5
Systematics Research Collections, University of Nebraska, Lincoln, Nebraska 68588-0514, USA;
e-mail: mjpaulsen@unl.edu.
6
Systematic Entomology Laboratory, Universidad del Valle de Guatemala, GUATEMALA; e-mail:
ecano@uvg.edu.gt.
7
Department of Biological Sciences, University of Memphis, Memphis, Tennessee 38152, USA; e-mail:
cfbeza@memphis.edu.
8
Universidad de La Guajira, Riohacha, COLOMBIA; e-mail: larryjimenezferbans@gmail.com.
9
Instituto de Ecologa, A. C., Xalapa 91070, Veracruz, MEXICO; e-mail: pedro.reyes@inecol.mx.
10
Corresponding Author E-mail: bratcliffe1@unl.edu.
Accepted 21 April 2014; Revised 1 December 2014
Peruvian biodiversity is pivotal for conserving and managing natural resources, food
security, poverty reduction, health, biosecurity, new industrial product development,
and ecotourism (Smith et al., 2011).
Geotrupidae
Diversity in Peru: 1 subfamily, 4 genera, and 11 species.
Recognition: The body shape is oval or round, and the head is not deflexed. The
antennae are 11-segmented with a 3-segmented, opposable club with all antennomeres
tomentose. The eyes are completely or partially divided by a canthus. The clypeus is
often with a tubercle or horn. The labrum is truncate, prominent, and produced beyond
the apex of the clypeus. The mandibles are prominent and produced beyond the apex of
the labrum. The pronotum is convex with a base wider than or subequal to the elytral
base and with or without tubercles, ridges, horns, or sulci. The elytra are convex, with
or without striae. The pygidium is concealed by the elytra (Jameson, 2002a).
Habitat: Life histories of the geotrupids are diverse, and food habits vary from
saprophagous to coprophagous and mycetophagous. Adults of most species are
secretive, living most of their life in burrows. Although adults do not tend larvae,
adults provision food for larvae in brood burrows. Adults dig vertical burrows (15
200 cm in depth) and provision larval cells with dead leaves, cow dung, horse dung, or
humus. Burrows of some species extend to a depth of 3.0 m (Jameson, 2002a).
Notes: The family Geotrupidae includes 68 genera and about 620 species (Scholtz
and Browne, 1996). The subfamily Geotrupinae does not occur in South America.
The following checklist of Peruvian Scarabaeidae is from Howden (1985, 2002) and
Howden and Martnez (1978).
BOLBOCERATINAE
Athyreini
Athyreus bicornus Howden, 2002* Neoathyreus fallolobus Howden, 2006
Athyreus larseni Howden, 2002* Neoathyreus ornatus Howden, 1985*
Athyreus martinezi Howden, 1995 Neoathyreus rufobrunneus Howden, 1985
Athyreus pyriformis Howden and Martnez, 1978* Neoathyreus rufoventris Howden, 1985*
Athyreus tribuliformis Felsche, 1909*
Bolboceratini
Bolboceras baeri Boucomont, 1902
Zefevazia peruana (Boucomont, 1902)*
Lucanidae
Diversity in Peru: 1 subfamily, 14 genera, 40 species.
Recognition: The head is prognathous and not deflexed. The antennae are
geniculate or straight, 10-segmented, and with a 37 segmented club (all
antennomeres unopposable and tomentose). The first antennomere is often subequal
in length to the remaining antennomeres.
Habitat: Lucanids are usually associated with decaying wood and logs in
coniferous and deciduous forests. Adults of some species are attracted to lights at
night and some feed at sap flows from fluxing trees. The eggs are usually laid in
crevices in bark or logs, and the larvae feed on decaying wood.
Notes: The stag beetle family Lucanidae includes four subfamilies, 108 genera, and
around 1,500 species worldwide. In the New World, there are 41 genera and 223
species with representatives of each of the four subfamilies (Paulsen and Ratcliffe,
2005). Most New World taxa are Neotropical. The following checklist of Peruvian
Lucanidae is from Paulsen (2013).
LUCANINAE
Brasilucanini
Brasilucanus acomus Ratcliffe, 1984
Chiasognathini
Sphaenognathus alticollis Mollenkamp, 1912 Sphaenognathus peruvianus (Waterhouse, 1869)
Sphaenognathus gaujoni (Oberthur, 1885) Sphaenognathus prionoides Buquet, 1838
Sphaenognathus giganteus Boileau, 1911 Sphaenognathus xerophilus Bartolozzi and Onore,
Sphaenognathus monguilloni Lacroix, 1972* 2006*
Sclerostomini
Aegognathus aguirei Arnaud and Bomans, 2007* Incadorcus damasoi Arnaud and Bomans, 2006*
Aegognathus confusus Arnaud and Bomans, 2006* Incadorcus michelleae Arnaud and Bomans, 2006*
Aegognathus leuthneri leuthneri Van der Poll, 1886 Incadorcus shaunai Arnaud et al., 2007*
Aegognathus leuthneri damasoi Arnaud and Incadorcus zugeri Arnaud and Bomans, 2006*
Bomans, 2006* Metadorcinus beneshi (Martnez, 1953)
Aegognathus similis Arnaud and Bomans, 2006* Metadorcinus lineatus (Deyrolle, 1864)*
Aegognathus soulai Arnaud and Bomans, 2004* Metadorcinus yamauchii Arnaud et al., 2008*
Aegognathus waterhousei Leuthner, 1883* Metadorcus ebeninus Deyrolle, 1864
Andinolucanus inesae Arnaud and Bomans, 2006* Onorelucanus boileaui (Weinreich, 1960)*
Arnaudius bomansi Grossi and Bartolozzi, 2011* Onorelucanus marujae (Arnaud et al., 2008)*
Arnaudius digennaroi (Arnaud et al., 2008)* Onorelucanus noguchii (Arnaud and Bomans,
Arnaudius koikei (Arnaud et al., 2007)* 2007)*
Auxicerus platyceps Waterhouse, 1883* Pseudoscortizus incredibilis Arnaud et al., 2008
Cantharolethrus azambrei Boileau, 1897 Sclerostomus bartolozzii Arnaud and Bomans,
Cantharolethrus elongatus Lacroix, 1982* 2007*
Cantharolethrus steinheili Parry, 1875* Sclerostomus damasoi Arnaud and Bomans, 2006*
Incadorcus ashaninka Grossi, 2011* Sclerostomus wendyae Arnaud and Bomans, 2006*
Incadorcus cuzcoensis Arnaud et al., 2007* Scortizus prodigiosus Arnaud et al., 2007*
Passalidae
Diversity in Peru: 1 subfamily, 2 tribes, 6 genera, 50 species.
Recognition: The body shape is elongate-cylindrical and depressed, and the length is
1380 mm. Their color is black and shiny. The head is prognathous, narrower than the
thorax, and often with a dorsomedian horn. The antennae have ten antennomeres
including a 3 to 5-segmented club that is not opposable and not geniculate but is
capable of being rolled together. The mandibles are large and project beyond the apex
of the labrum, and the apex of the labrum is large, rounded, and toothed. The
pronotum is broader than the head, quadrate, and the surface is smooth with a median,
longitudinal groove. The elytra are elongate, parallel sided, with rounded apices and
well-developed striae. The pygidium is concealed by the elytra (Schuster, 2002).
Habitat: Passalid adults live in well-decayed logs and stumps with their larvae in
subsocial family groups. All stages are found in galleries of wood that are excavated
by the adults. Eggs are usually placed together in a nest of frass. Adults and larvae
communicate by stridulating and can produce different calls. Adults care for larvae
and prepare food by chewing it and presumably mixing it with saliva (Schuster,
2002). One species (Ptichopus angulatus [L.]) inhabits the waste chamber in nests of
leaf cutter ants, Atta species.
Notes: Most New World taxa occur in the Neotropics. Hincks and Dibb (1935)
cited passalid species from Peru, and Van Doesburg (1942) provided a checklist of
Peruvian Passalidae.
PASSALINAE
Passalini
Passalus abortivus Percheron, 1835 Passalus plicatus Percheron, 1835
Passalus aduncus Erichson, 1847 Passalus prominens Gravely, 1918*
Passalus anguliferus Percheron, 1835 Passalus pubicostatus (Kuwert, 1898)
Passalus arrowi Hincks, 1934 Passalus pugionatus Burmeister, 1847
Passalus barrus Boucher and Reyes-Castillo, 1991 Passalus pugionifer (Kuwert, 1891)*
Passalus caelatus Erichson, 1847 Passalus punctiger Le Peletier and Servillle, 1825
Passalus coniferus Eschscholtz, 1829 Passalus rhodocanthopoides (Kuwert, 1891)
Passalus convexus Dalman, 1817 Passalus rotundatus Hincks, 1940*
Passalus elfriedae Luderwaldt, 1931 Passalus rusticus Percheron, 1835
Passalus episcopus (Kuwert, 1898)* Passalus schneideri (Kuwert, 1898)*
Passalus huebneri (Kuwert, 1898)* Passalus spinifer Percheron, 1835
Passalus inca (Zang, 1905)* Passalus zangi Hincks, 1934
Passalus interruptus (L., 1758) Paxillus camerani (Rosmini, 1902)
Passalus interstitialis Eschscholtz, 1829 Paxillus.forsteri Luderwaldt, 1927
Passalus latifrons Percheron, 1841 Paxillus leachi MacLeay, 1819
Passalus occipitalis Eschscholtz, 1829 Spasalus crenatus (MacLeay, 1819)
Passalus peruvianus (Kuwert, 1898) Spasalus kaupi Boucher, 2004*
Proculini
Popilius amazonicus Gravely, 1918* Veturius inca Boucher, 2006
Popilius marginatus (Percheron, 1835) Veturius libericornis Kuwert 1891
Verres furcilabris (Eschscholtz, 1829) Veturius sinuosus (Drapiez, 1820)
Veturius amazonicus Boucher, 2006 Veturius spinipes (Zang, 1905)
Veturius arawak Boucher, 2006 Veturius standfussi Kuwert, 1891
Veturius cephalotes (Le Peletier and Serville, 1825) Veturius tarsipes Boucher, 2006*
Veturius ecuadoris Kuwert, 1898 Veturius unicornis Gravely, 1918
Veturius guntheri Kuwert, 1898 [5 peruvianus of Veturius yahua Boucher, 2006
Arrow]
Trogidae
Diversity in Peru: 1 subfamily, 2 genera, 10 species.
Recognition: Adult trogids are recognized by their warty, brown to gray to black,
dirt-encrusted appearance and flat abdominal sternites.
Habitat: Adults and larvae are among the last scavengers that visit the dry remains
of dead animals, where they feed on feathers, fur, and skin. They also feed on
organic matter found in the nests of mammals and birds.
Notes: The world fauna consists of 4 genera and 300 species (Scholtz, 1982). The
following checklist of Peruvian trogids is from Scholtz (1982).
TROGINAE
Omorgus howelli (Howden and Vaurie, 1957) Polynoncus bullatus (Curtis, 1845)
Omorgus suberosus (Fabricius, 1775) Polynoncus ecuadorensis (Vaurie, 1962)
Omorgus persuberosus (Vaurie, 1962) Polynoncus gordoni (Steiner, 1981)*
Polynoncus aeger (Guerin-Meneville, 1844)* Polynoncus gemmingeri (Harold, 1872)
Polynoncus aricensis (Gutierrez, 1950) Polynoncus peruanus (Erichson, 1847)
Polynoncus brevicollis (Eschscholtz, 1822) Polynoncus sallei (Harold, 1872)
Ochodaeidae
Diversity in Peru: 1 subfamily, 1 genus, 1 species.
Recognition: Ochodaeids are elongate and convex. Their color is testaceous,
brown, reddish brown, brown, black, or occasionally bicolorous. The head is not
deflexed. The antennae have 9 or 10 antennomeres, with a 3-segmented, opposable
club, and all antennomeres are tomentose. The clypeus is simple or tuberculate on
the anterior margin. The labrum is produced beyond the apex of the clypeus, often
bilobed and emarginate, and prominent. The prominent mandibles are produced
beyond the apex of the labrum. The pronotum is convex, subquadrate, mostly
punctate and setose and without tubercles, ridges, horns, or sulci. The mesotibia at
the apex has at least one apical spur pectinate. The elytra are convex, with or without
striae, often punctate, or granulate and setose, but some are smooth. The pygidium is
exposed or concealed by the elytra (Carlson, 2002).
Habitat: Ochodaeidae are most often collected at lights, sometimes in large
numbers. Adults of a few species are active during the day. Many species prefer
sandy areas. Adults may spend the day in subterranean burrows, where they may
feed on fungi. Little is known about the biology of Ochodaeidae. There are few
observations of adult or larval habits except that adults of most species are nocturnal
and are attracted to lights (Carlson, 2002).
Notes: The family Ochodaeidae includes 10 genera and about 80 species worldwide
(Carlson and Paulsen, 2012).
OCHODAEINAE
Parochodaeus bituberculatus (Erichson, 1847)*
Hybosoridae
Recognition: Hybosorids are light brown to black and glossy. The head is not
deflexed. The antennae are 10-segmented with a 3-segmented, opposable club with
the last 2 antennomeres tomentose. The first antennomere of the club is hollowed to
receive antennomeres 2 and 3. The clypeus generally lacks a tubercle or horn. The
prominent labrum is truncate and produced beyond the apex of the clypeus. The
large mandibles project beyond the apex of the labrum and have the external edge
rounded with the apices pointed.
Habitat: Little life history information is known for hybosorids. Adults feed on
both invertebrate and vertebrate carrion in the early stages of decomposition. Some
species are found in dung, and others are attracted to lights at night (Jameson,
2002b). Ceratocanthines are commonly found in clusters of dead leaves and can be
collected using a beating sheet.
Notes: The family Hybosoridae worldwide contains 4 subfamilies, 35 genera, and
about 220 species (Ocampo, 2006; Ocampo and Ballerio, 2006). Species are widely
distributed in the tropics. The classification below is from Ocampo and Ballerio (2006).
ANAIDINAE
Anaides onofrii Ocampo, 2006 Chaetodus allsoppi Martnez, 1988*
Anaides rugosus Robinson, 1948* Chaetodus asuai Martnez, 1956
Chaetodus mimi Ocampo, 2006* Hybochaetodus erugocarinatus Ocampo, 2006*

Chaetodus smithi Ocampo, 2006* Hybochaetodus flaco Ocampo, 2002*
Chaetodus tricarinatus Ocampo, 2006 Hybochaetodus obscurus Arrow, 1909*
Hybochaetodus disruptus Ocampo, 2006*
CERATOCANTHINAE
Anopsiostes punctatus Paulian, 1982 Ceratocanthus suturalis (Lansberge, 1887)
Astaenomoechus americanus (Boucomont, 1936) Germarostes antiquus (Erichson, 1843)
Astaenomoechus criberrimus Paulian, 1982* Germarostes aphodioides (Illiger, 1800)
Ceratocanthoides undatus (Petrovitz, 1973) Germarostes carltoni Howden and Gill, 2005*
Ceratocanthus clypealis (Lansberge, 1887)* Germarostes geayi Paulian, 1982*
Ceratocanthus inca Paulian, 1982* Germarostes macleayi (Perty, 1830)
Ceratocanthus mathani Paulian, 1982* Germarostes semituberculatus (Germar, 1843)
Ceratocanthus perpunctatus Paulian, 1982* Germarostes sulcipennis Harold, 1875*
Ceratocanthus politus (Erichson, 1843) Trachycrusus lescheni Howden and Gill, 1995*
Ceratocanthus punctulatus Lansberge, 1887* Trachycrusus striatulus Howden and Gill, 1995*
Ceratocanthus seriatus (Erichson, 1843)
HYBOSORINAE
Coilodes punctipennis Arrow, 1909
Scarabaeidae
Diversity in Peru: 10 subfamilies, 194 genera, and 898 species.
Recognition: Scarabs are 2.0160.0 mm in length with variable shape and color,
with or without metallic reflections, and with or without vestiture. The antennae
have 10 antennomeres (some 712) with a 3 to 5-segmented, opposable club; the
club has the apical antennomeres nearly glabrous (Melolonthinae, Dynastinae,
Rutelinae, Cetoniinae) or with all antennomeres tomentose (Aphodiinae,
Scarabaeinae). The clypeus is with or without a tubercle or horn. The labrum is
distinct in most taxa, and produced beyond the apex of the clypeus or not. The
mandibles are variable, produced beyond the apex of the labrum or not. The
pronotum is variable, with or without horns or tubercles. The elytra are convex or
flattened and with or without striae. The pygidium is concealed by the elytra
(Aphodiinae, Scarabaeinae) or exposed (Scarabaeinae, Melolonthinae, Dynasti-
nae, Rutelinae, Cetoniinae). The scutellum is exposed or not, and its shape is
triangular or parabolic. The legs have transverse or conical coxae; the protibiae
are tridentate, bidentate, or serrate on the outer margin; the meso- and metatibia
at the apex have 1 or 2 spurs; the tarsi are 5-5-5, and the anterior tarsi are absent
in some Scarabaeinae; the tarsal claws are variable, equal in size or not, and
simple or toothed. The abdomen has 6 free sternites; the 7 functional abdominal
spiracles are situated in the pleural membrane (Aphodiinae, Scarabaeinae) or in
the sternites and tergites (Melolonthinae, Dynastinae, Rutelinae, and Cetoniinae).
Habitat: Scarab beetles occupy many habitats ranging from deserts to tropical
rainforests and have diverse life histories. Depending on the group, adults feed on
foliage, dung, flowers, or sap while the larvae are phytophagous, coprophagous,
or detritivorous. Some are diurnal, while many are nocturnal in their activity
patterns.
Notes: Members of the subfamily Aphodiinae, known as the aphodiine dung
beetles, are primarily dung and detritus feeders. All of the species are small, rarely
exceeding 12 mm in length. Worldwide, the subfamily contains 12 tribes,
approximately 280 genera, and 3,200 species with 9 tribes, 128 genera and 816 species
in the New World (Skelley, 2008). In Peru, there are 24 genera and 63 species
(Skelley, 2008).
The subfamily Scarabaeinae is commonly referred to as dung beetles. Most
species feed on mammal dung, but others specialize upon the dung of other
vertebrates and invertebrates as well as on carrion, mushrooms, rotting fruit, and
other decomposing plant material. The world fauna includes over 5,000 described
species in 234 genera, with about 1,800 species just in the genus Onthophagus (Gill,
2005). The tribal classification below follows that of Hanski and Cambefort (1991).
Peru is incredibly rich in dung beetles, and over 150 species are recorded from
a single lowland forest site in Madre de Dios (Los Amigos, Peru) (Trond Larsen,
personal communication to BCR, February 2014). Larsen and Genier (2008a, b)
provided color guides for identifying dung beetles at two different biological
stations in Peru. Peru has 40 genera and 278 species, but many more will be
recognized in the future, especially in the genera Ateuchus, Canthidium,
Dichotomius, and Uroxys. In comparison, neighboring Brazil includes 49 genera
and 618 species (Vaz-de-Mello, 2000).
The Melolonthinae, often called May beetles, June beetles, or leaf chafers, are
cosmopolitan in distribution and one of the largest and most diverse subfamilies of
Scarabaeidae. There are approximately 750 genera and over 11,000 species
worldwide, with about 90 genera in the New World (Evans and Smith, 2005). Peru
has 20 genera and 82 species, but this number will increase as more species are
recognized. This is not now easily accomplished because identification of some
genera and many species remains in a state of confusion. The tribes are still
unresolved due to the lack of definition and inconsistent use of characters (Evans
and Smith, 2005). Reliable identifications are difficult since the majority of
descriptions prior to 1940 lacked illustrations or comparisons with other similar
species.
The subfamily Rutelinae (leaf chafers) includes 7 tribes, 2 of which do not occur in
Peru (Adoretini from the Old World and Alvarengiini from southern Brazil), and
approximately 4,300 species. Based on this checklist, the leaf chafers of Peru
encompass a rich fauna with 52 genera, 270 species and subspecies, and 150 endemic
species. The neighboring country of Ecuador includes 53 genera, 298 species, and 92
endemic species (Paucar-Cabrera, 2005). The last compilation of ruteline species of
Peru was a product of the Hamburg Southern Peru Expedition in 1936, and this
yielded 29 genera and 85 species (Ohaus, 1952).
Because of the economic importance and interest in the Rutelinae, classification
and nomenclature of the group is undergoing extensive revisions. This checklist
reflects changes in the tribes Anomalini (Ramrez-Ponce and Moron, 2009; Moron
and Ramrez-Ponce, 2012), Anoplognathini (Smith, 2003), Anatistini (previously
Spodochlamyini; Jameson and Ratcliffe, 2011), Geniatini (Hawkins and Jameson,
2005; Soula, 2010), and Rutelini (Soula, 1998, 2002a, b, 2003, 2005, 2006a, b, 2008,
2009, 2010, 2011a, b; Jameson, 1998; Moore and Jameson, 2013). It should be noted
that Soulas species concept was typological as well as topological; specimens that
displayed variation from the type specimen and/or specimens from unique localities
were often described as new species, subspecies, or varieties, thus reflecting a stamp
collector view of biodiversity. As a result, Soula greatly overestimated ruteline
diversity by describing many new genera and species that we believe will become
invalid. In this checklist, Soulas taxa comprise nearly half of the ruteline diversity
(119 species and 31 subspecies) in Peru. Because Soulas names were established in
accordance with the rules of zoological nomenclature, the names must be considered
as available. We do not address validity of Soulas species and genera here, but we
caution the user in accepting Soulas classification and species concepts. Because of
Soulas overestimation of ruteline diversity, lack of survey throughout South
America, and poor taxonomic foundation, the Rutelinae are currently poor
bioindicators for habitat conservation. Rutelines are important ecologically as
pollinators and decomposers; agriculturally as consumers of crop leaves, roots, and
fruits; and culturally as food and decorative adornment. Species in the subfamily are
often highly metallic and large (up to 50 mm in length). Because of their beauty and
abundance, the Shuar people (also Achuar, apach, inkis, jvaros, or jbaros) of the
eastern Amazonian areas in Peru and Ecuador use the bodies of some rutelines
(especially Chrysophora chrysochlora) to create headpieces and necklaces (Ratcliffe,
2006). These decorative pieces are now popular among tourists and local people
alike.
The subfamily Dynastinae is one of the most conspicuous subfamilies of the beetle
family Scarabaeidae. Members of the subfamily occur in all the major biogeographic
areas of the world (except the polar regions), although most species are found in the
tropics, specifically the New World tropics. Dynastines comprise 196 genera and
about 1,500 species grouped among 8 tribes. Six tribes, 87 genera, and at least 800
species occur in the New World, and most of these species occur in the Neotropics.
Our checklist for Peru currently has 39 genera and 182 species. By comparison,
Mexico, another large and megadiverse country, has 30 genera and 196 species
(Ratcliffe et al., 2013), while Costa Rica and Panama together have 36 genera and
158 species (Ratcliffe, 2003).
Adult dynastines are small (4 mm) to very large (160 mm) beetles. The males in
some species (principally Dynastini, Agaocephalini, and Oryctini) possess prominent
and often spectacular horns on the head and/or prothorax which, together with their
large size, have given rise to such popular names for them as rhinoceros,
elephant, hercules, and unicorn beetles. In fact, the entire subfamily is usually
referred to as the rhinoceros beetles even though the majority of species do not
possess horns. The adults of nearly all species are nocturnal or crepuscular,
and many are readily attracted to lights at night. Adult dynastines are known to
feed on ripe or rotting fruits, slime fluxes, and plant roots. The adults of some
Cyclocephalini are important pollinators of palms and aroids when they feed on the
floral parts of these plants.
The subfamily Cetoniinae, or flower chafers, are most abundant in Africa and
Asia. In the New World they comprise 6 tribes in 41 genera with approximately 275
species (Krajcik, 2012). In Peru there are 4 tribes containing 8 genera and 18 species.
The genus Gymnetis is still in need of revision, and the species names below have
presumed synonyms that may turn out to be valid species.
The following checklist of Peruvian Scarabaeidae is extracted from many sources,
principal among them are Ocampo and Colby (2009) (Allidiostomatinae); Skelley
(2008), Smith and Skelley (2007), and Stebnicka (2005, 2007, 2009) (Aphodiinae);
Edmonds (1994, 2000), Edmonds and Zidek (2010, 2012), and Figueroa et al. (2014)
(Scarabaeinae in part); Evans and Smith (2009) (Melolonthinae); Machatschke
(1972), Villatoro (2002), Jameson (1998), and Soula (19982011) (Rutelinae); Colby
(2009) (Orphninae), Endrodi (19661985) (Dynastinae); Erichson (1847) (Peru);

Blackwelder (1944) (Scarabaeoidea), and Krajcik (2012). The older checklists in
particular should be used with caution because classification and some names may
have changed.
APHODIINAE
Aphodiini
Aidophus infuscatopennis (Schmidt, 1909) Neotrichaphodioides woytkowskii Dellacasa, Dellacase,
Aphodius pseudolividus Balthasar, 1941 and Skelley, 2010*
Blackburneus laxepunctatus (Schmidt, 1910) Orodaliscoides rugosiceps (Harold, 1859)
Gonaphodiellus castanescens (Petrovitz, 1973) Paranimbus peruanus (Erichson, 1834)
Gonaphodiellus chapini (Hinton, 1934) Trichaphodiellus brasiliensis (Laporte, 1840)
Gonaphodiellus nigrinus (Schmidt, 1916)*
Eupariini
Ataeniopsis regulis (Balthasar, 1947) Ataenius petrovitzi Balthasar, 1960
Ataenius abancay Stebnicka, 2005* Ataenius picinus Harold, 1868
Ataenius aequatorialis Petrovitz, 1961 Ataenius platensis (Blanchard, 1846)
Ataenius atramentarius (Erichson, 1847) Ataenius rubrotessellatus (Blanchard, 1843)
Ataenius attenuator Harold, 1874 Ataenius santarosae Stebnicka, 2007
Ataenius buenavistae Stebnicka, 2001 Ataenius sculptilis Harold, 1868
Ataenius carinator Harold, 1874 Ataenius siminasus Petrovitz, 1973
Ataenius catenulatus (Erichson, 1847)* Ataenius strigicaudus Bates, 1887
Ataenius columbicus Harold, 1889 Ataenius tambopatae Stebnicka, 2001*
Auperia andamanensis (Koshantschikov, 1916)
Ataenius complicatus Harold, 1869
Auperia capitosus (Harold, 1867)
Ataenius costulifer Balthasar, 1941*
Auperia huebneri (Petrovitz, 1970)
Ataenius crenaticollis Petrovitz, 1973
Auperia iquitosae (Stebnicka, 2002)
Ataenius gracilis (Melsheimer, 1844)
Auperia loretoensis (Stebnicka, 2002)*
Ataenius huanus Stebnicka, 2007 Euparixia boliviana Gordon and McCleve, 2003
Ataenius icanus Balthasar, 1951* Euparixiodes johnsoni Stebnicka, 1998
Ataenius impiger Schmdt, 1916 Lomanoxoides mapitunari Stebnicka and Skelley,
Ataenius lamarensis Stebnicka, 2007 2005*
Ataenius montanus Schmidt, 1911 Odontolytes iquitosae (Stebnicka, 2007)
Ataenius morator Harold, 1869 Passaliolla corticalis (Bates, 1887)
Ataenius napoensis Stebnicka, 2007 Saprosites dentipes Harold, 1867
Ataenius nugator Harold, 1880 Saprosites parallelus Harold, 1867
Ataenius palmaritoensis Stebnicka, 2007 Saprosites sulcatus Harold, 1869
Odontolochini
Amerilochus cinereus Skelley, 2007* Saprositellus denticulatus Balthasar, 1967
Saprolochus tambopatae Stebnicka and Galante, Saprositellus peruanus Stebnicka, 2003*
2007 Stebnickiella zosterixys Skelley, 2007*
Saprolochus tridentatus Skelley, 2007
Psammodiini
Mysarus peruanus Petrovitz, 1962*
Rhyparini
Aschnarhyparus peregrinus (Hinton, 1934)
Stereomerini
Termitaxis holmgreni Krikken, 1970*
SCARABAEINAE
Onthophagini
Digitonthophagus gazella (Fabricius, 1787) Onthophagus osculatii Guerin-Meneville, 1855
Onthophagus bidentatus Drapiez, 1819* Onthophagus ptox Erichson, 1847*
Onthophagus clypeatus Blanchard, 1846 Onthophagus ranunculus Arrow, 1913
Onthophagus coscineus Bates, 1887 Onthophagus rhinophyllus Harold, 1868
Onthophagus haematopus Harold, 1875 Onthophagus rubrescens Blanchard, 1846
Onthophagus marginicollis Harold, 1880 Onthophagus schunckei Paulian, 1936*
Onthophagus onorei Zunino and Halffter, 1998 Onthophagus sp. aff. tristis Harold, 1873
Onthophagus onthochromus Arrow, 1913 Onthophagus xanthomerus Bates, 1887
Onthophagus ophion Erichson, 1847 Onthophagus sp. aff. xanthomerus Bates, 1887
Oniticellini
Eurysternus caribaeus (Herbst, 1789) Eurysternus lanuginosus Genier, 2009
Eurysternus cayennensis Laporte, 1840 Eurysternus marmoreus Laportee, 1840
Eurysternus contractus Genier, 2009 Eurysternus nigrovirens Genier, 2009
Eurysternus foedus Guerin-Meneville, 1830 Eurysternus plebejus Harold, 1880
Eurysternus gracilis Genier, 2009 Eurysternus squamosus Genier, 2009
Eurysternus hamaticollis Balthasar, 1939 Eurysternus streblus Genier, 2009
Eurysternus howdeni Genier, 2009 Eurysternus strigilatus Genier, 2009
Eurysternus hypocrita Balthasar, 1939 Eurysternus vastiorum Martnez, 1988
Eurysternus inca Genier, 2009 Eurysternus wittmerorum Martnez, 1988
Eurysternus inflexus Germar, 1824
Canthonini
Anisocanthon villosus (Harold, 1868) Canthon septemmaculatus septemmaculatus
Canthon aberrans (Harold, 1868) (Latreille, 1807)
Canthon aequinoctialis Harold, 1868 Canthon septemmaculatus histrio LePeletier and
Canthon angustatus Harold, 1867 Serville, 1828
Canthon balteatus Boheman, 1858 Canthon sericatus Schmidt, 1922
Canthon bimaculatus Schmidt, 1922 Canthon simulans Martnez, 1950
Canthon brunneus Schmidt, 1922 Canthon smaragdulus (Fabricius, 1781)
Canthon chiriguano Martinez and Halffter, 1972 Canthon subhyalinus Harold, 1867
Canthon coloratus Schmidt, 1922 Canthon triangulare Drury, 1870
Canthon conformis Harold, 1868 Canthon unicolor Blanchard, 1843
Canthon fulgidus Redtenbacher, 1867 Canthon velutinus Harold, 1868
Canthon fuscipes Erichson, 1847 Canthon virens chalybaeus Blanchard, 1843
Canthon gemellatus Erichson, 1847 Canthonella sp. aff. amazonica Ratcliffe and Smith,
Canthon helleri Schmidt, 1922 1999
Canthon janthinus Blanchard, 1843 Canthonella barrerai Halffter and Martnez, 1968
Canthon juvencus Harold, 1868 Canthonella sp. aff. catharinensis Pereira and
Canthon laesum Erichson, 1847 Martnez, 1956
Canthon lituratus Germar, (1813) Canthonella cf. gomezi Halffter and Martnez, 1968
Canthon luteicollis Erichson, 1847 Canthonidia rubromaculata (Blanchard, 1846)
Canthon sp. aff. matthewsi Martnez and Halffter, Cryptocanthon campbellorum Howden, 1973
1972 Deltochilum aequinoctiale Buquet, 1844
Canthon monilifer Blanchard, 1843 Deltochilum amazonicum Bates, 1887
Canthon mutabilis Lucas, 1857 Deltochilum cf. aureopilosum Paulian, 1939
Canthon muticus Harold, 1867 Deltochilum burmeisteri Harold, 1867
Canthon pallidus Schmidt, 1922 Deltochilum carinatum Westwood, 1837
Canthon sp. aff. pallidus Schmidt, 1922 Deltochilum crenulipes Paulian, 1938
Canthon paraguayanum Balthasar, 1939 Deltochilum erodioides Harold, 1867
Canthon quadriguttatus (Olivier, 1789) Deltochilum fuscocupreum Bates, 1870
Canthon quinquemaculatus Laporte, 1840 Deltochilum granulatum Bates, 1870
Canthon rubrescens Blanchard, 1843 Deltochilum howdeni Martnez, 1955
Canthon semiopacus Harold, 1868 Deltochilum hypponum Buquet, 1844
Deltochilum sp. aff. komareki Balthasar, 1939 Scybalocanthon aereus (Schmidt, 1922)
Deltochilum mexicanum Burmeister, 1848 Scybalocanthon imitans (Harold, 1868)
Deltochilum orbiculare Lansberge, 1874 Scybalocanthon moniliatus (Bates, 1887)
Deltochilum peruanum Paulian, 1939 Scybalocanthon pinopterus (Kirsch, 1873)
Deltochilum pretiosum Harold, 1875 Scybalocanthon sexspilotus (Guerin-Meneville,
Deltochilum pseudoicarus Balthasar, 1939 1855)
Deltochilum robustus Molano and Gonzalez, 2009 Scybalocanthon trimaculatus (Schmidt, 1922)
Deltochilum tessellatum Bates, 1870 Scybalocanthon zischkai Martnez, 1949
Deltochilum valgum Burmeister, 1873 Scybalophagus rugosus (Blanchard, 1846)
Deltochilum sp. aff. valgum Burmeister, 1873 Streblopus opatroides Lansberge, 1874
Malagoniella astyanax (Olivier, 1789) Streblopus punctatus Balthasar, 1938*
Malagoniella cupreicollis Waterhouse, 1890* Sylvicanthon bridarollii (Martnez, 1949)
Pseudocanthon felix (Arrow, 1913) Sylvicanthon furvus (Schmidt, 1921)
Pseudocanthon xanthurum (Blanchard, 1843)
Dichotomiini
Anomiopus andrei Canhedo, 2006 Canthidium sp. aff. centrale Boucomont, 1928
Anomiopus batesi (Waterhouse, 1891) Canthidium coerulescens Balthasar, 1939
Anomiopus brevipes (Waterhouse, 1891) Canthidium cupreum (Blanchard, 1846)
Anomiopus cambeforti Canhedo, 2006* Canthidium sp. aff. deyrollei Harold, 1867
Anomiopus foveicollis Canhedo, 2006 Canthidium discolor Harold, 1867
Anomiopus gilli Canhedo, 2006* Canthidium cf. dohrni Harold, 1867
Anomiopus idei Canhedo, 2006* Canthidium escalerai Balthasar, 1939
Anomiopus intermedius (Waterhouse, 1891) Canthidium sp. aff. funebre Balthasar, 1939
Anomiopus pictus (Harold, 1862) Canthidium gerstaeckeri Harold, 1867
Anomiopus pishtaco Edmonds and Figueroa, 2013* Canthidium sp. aff. gigas Balthasar, 1939
Anomiopus pumilius Canhedo, 2006 Canthidium histrio Balthasar, 1939
Anomiopus smaragdinus (Westwood, 1842) Canthidium kirschi Harold, 1875
Anomiopus validus Canhedo, 2006* Canthidium lentum Erichson, 1847
Ateuchus aeneomicans (Harold, 1868) Canthidium miscellum Harold, 1883
Ateuchus cereus (Harold, 1868) Canthidium cf. onitoides (Perty, 1830)
Ateuchus columbianus (Harold, 1868) Canthidium sp. aff. quadridens Harold, 1867
Ateuchus connexus (Harold, 1868) Canthidium cf. ruficolle (Germar, 1824)
Ateuchus sp. aff. laevicollis (Harold, 1868) Canthidium thalassinum (Erichson, 1847)
Ateuchus peruanus (Balthasar, 1939)* Dichotomius adrastus (Harold, 1875) *
Ateuchus pygidialis (Harold, 1868) Dichotomius apicalis (Luderwaldt, 1931)
Ateuchus sp. aff. pygidialis Harold, 1868 Dichotomius batesi (Harold, 1869)
Ateuchus scatimoides (Balthasar, 1939) Dichotomius belus (Harold, 1869) *
Ateuchus sp. aff. setulosus Balthasar, 1939 Dichotomius bicornis (Waterhouse, 1891)*
Ateuchus simplex (Le Peletier and Serville, 1828) Dichotomius bicuspis Germar, 1824
Ateuchus striatulus (Preudhomme de Borre, 1886) Dichotomius calcaratus (Arrow, 1913)
Ateuchus substriatus (Harold, 1868) Dichotomius camargoi (Martnez, 1955)
Ateuchus viridimicans (Boucomont, 1935) Dichotomius conicollis (Blanchard, 1843)
Besourenga horacioi (Martnez, 1967) Dichotomius cotopaxi (Guerin, 1855)
Bdelyrus cochabambae Cook, 2000 Dichotomius cuprinus (Felsche, 1901)*
Bdelyrus howdeni Cook, 1998 Dichotomius diabolicus (Harold, 1875)
Bdelyrus iquitosensis Cook, 2000* Dichotomius fissus (Harold, 1867)
Bdelyrus lobatus Cook, 1998* Dichotomius sp. aff. fonsecae (Luderwaldt, 1924)
Bdelyrus parvus Cook, 1998 Dichotomius globules (Felsche, 1901)
Bdelyrus pecki Cook, 1998 Dichotomius inachus (Erichson, 1847)
Bdelyrus peruviensis Cook, 1998* Dichotomius lucasi (Harold, 1869)
Bradypodidium adisi (Ratcliffe, 1980) Dichotomius mamillatus (Felsche, 1901)
Canthidium angusticeps (Bates, 1887) Dichotomius melzeri (Luderwaldt, 1922)
Canthidium atramentarium Balthasar, 1939 Dichotomius ocellapunctatus (Felsche, 1901)
Canthidium basipunctatum Balthasar, 1939 Dichotomius ohausi (Luderwaldt, 1924)
Canthidium batesi Harold, 1867 Dichotomius planicollis (Gillet, 1911)
Canthidium bicolor Boucomont, 1928 Dichotomius prietoi Martnez and Martnez, 1982
Dichotomius problematicus (Luderwaldt, 1924) Ontherus obliquus Genier, 1996

Dichotomius protectus (Harold, 1867) Ontherus pubens Genier, 1996
Dichotomius pullus (Felsche, 1910)* Ontherus raptor Genier, 1996
Dichotomius quinquelobatus (Felsche, 1901) Ontherus rectus Genier, 1996*
Dichotomius robustus (Luderwaldt, 1935) Ontherus sulcator (Fabricius, 1775)
Dichotomius satanas (Harold, 1867) Ontherus tenuistriatus Genier, 1996*
Dichotomius semiaeneus (Germar, 1824) Ontherus ulcopygus Genier, 1996
Dichotomius simplicicornis (Luderwaldt, 1924) Scatimus cucullatus Erichson, 1847*
Dichotomius virescens (Luderwaldt, 1924) Scatimus monstrosus Balthasar, 1939
Dichotomius worontzowi (Pereira, 1942) Scatimus sp. aff. onorei Genier and Kohlmann,
Genieridium cryptops (Arrow, 1913 2003
Homocopris torulosus (Eschscholtz, 1822) Scatimus quadricuspis Genier and Kohlmann,
Onoreidium cristatum (Arrow, 1931) 2003*
Ontherus alexis (Blanchard, 1846) Scatimus strandi Balthasar, 1939
Ontherus aphodioides Burmeister, 1847 Sinapisoma minuta Boucomont, 1928
Ontherus ashei Genier, 1996* Trichillum externepunctatum (Preudhomme de
Ontherus azteca Harold, 1869 Borre, 1880)
Ontherus brevipennis Harold, 1867 Uroxys bahianus Boucomont, 1927
Ontherus bridgesi Waterhouse, 1891 Uroxys elongatus Harold, 1868
Ontherus sp. aff. bridgesi Waterhouse, 1891 Uroxys sp. aff. kratochvili Balthasar, 1947
Ontherus carinifrons Luderwaldt, 1930 Uroxys sp. aff. minutus Harold, 1868
Ontherus edentulus Genier, 1996 Uroxys peruanus Balthasar, 1940*
Ontherus howdeni Genier, 1996 Uroxys sp. aff. simplex Waterhouse, 1891
Ontherus laminifer Balthasar, 1938 Uroxys variabilis Robinson, 1951
Phanaeini
Coprophanaeus callegarii Arnaud, 2002* Megatharsis buckleyi Waterhouse, 1891
Coprophanaeus degallieri Arnaud, 1997 Oruscatus davus (Erichson, 1847)
Coprophanaeus ignecinctus (Felsche, 1909) Oxysternon conspicillatum (Weber, 1801)
Coprophanaeus lancifer (L., 1767) Oxysternon lautum (MacLeay, 1819)
Coprophanaeus larseni Arnaud, 2002 Oxysternon silenus Laporte, 1840
Coprophanaeus ohausi (Felsche, 1911) Oxysternon spiniferum Laporte, 1840
Coprophanaeus parvulus (Olsoufieff, 1924) Phanaeus achilles Boheman, 1858
Coprophanaeus suredai Arnaud, 1996 Phanaeus bispinus Bates, 1868
Coprophanaeus telamon (Erichson, 1847) Phanaeus cambeforti Arnaud, 1982
Dendropaemon angustipennis Harold, 1869 Phanaeus chalcomelas (Perty, 1830)
Dendropaemon telephus Waterhouse, 1891 Phanaeus haroldi Kirsch, 1871
Diabroctis mimas (L., 1758) Phanaeus lecourti Arnaud, 2000
Gromphas aeruginosa (Perty, 1830) Phanaeus lunaris Taschenberg, 1870
Gromphas amazonica (Bates, 1870) Phanaeus meleagris Blanchard, 1843
Phanaeus sororibispinus Edmonds and Zidek, 2012 Sulcophanaeus actaeon (Erichson, 1847)*
Sulcophanaeus faunus (Fabricius, 1775) Tetramereia convexa (Harold, 1869)
ORPHNINAE
Aegidinus petrovi Colby, 2009* Paraegidium costalimai Vulcano, Pereira, and
Aegidinus teamscaraborum Colby, 2009 Martnez, 1966
ALLIDIOSTOMATINAE
Allidiostoma simplicifrons (Fairmaire, 1885)
Parallidiostoma tricornum Ocampo and Colby, 2009*
MELOLONTHINAE
Sericini
Astaena andicola Frey, 1973* Astaena exquisita Frey, 1973*
Astaena biciliata Saylor, 1946* Astaena glabroclypealis Frey, 1974
Astaena moseri Frey, 1973* Astaena pygidia Saylor, 1946*

Astaena negligens Frey, 1973* Astaena setosa Frey, 1973*
Astaena penai Frey, 1973* Astaena tridentata Erichson, 1847
Astaena peruana Moser, 1918* Raysymmela huanuca Saylor, 1947*
Astaena peruensis Frey, 1973* Symmela varians Erichson, 1847*
Astaena pottsi Saylor, 1946*
Diplotaxini
Liogenys leechi Frey, 1967*
Melolonthini
Phyllophaga austera (Erichson, 1847)* Phyllophaga pachypyga (Burmeister, 1855)
Phyllophaga jumberea Saylor, 1942* Phyllophaga peruana (Moser, 1918)
Phyllophaga marcapatana Moser, 1918* Phyllophaga umbrosa (Erichson, 1847)*
Macrodactylini
Ancistrosoma hilare Arrow, 1913 Isonychus cervinalis Frey, 1965*
Ancistrosoma intermedium Arrow, 1913* Isonychus cervinoides Frey, 1965*
Ancistrosoma klugii Curtis, 1836* Isonychus cervinus Erichson, 1847
Ancistrosoma reductum Frey, 1964* Isonychus egregius Frey, 1965*
Ancistrosoma vittigerum Erichson, 1847 Isonychus fraudulentus Frey, 1969*
Barybus compacta (Erichson, 1847)* Isonychus nitens Moser, 1921*
Barybus peruana Moser, 1918* Isonychus nubeculus Frey, 1969*
Barybus squamiger Frey, 1967* Isonychus ovinus Erichson, 1847*
Calodactylus abendrothii Kirsch, 1873* Isonychus pavonii Erichson, 1847*
Calodactylus heterosquamosus Frey, 1973* Isonychus peruanus Moser, 1921*
Ceraspis innotata (Blanchard, 1850)* Isonychus pulchellus Moser, 1918*
Ceraspis penai Frey, 1964* Isonychus rosettae Frey, 1969*
Ceraspis rubiginosa (Latreille, 1811)* Isonychus saylori Frey, 1969*
Ceraspis rufoscutellata Moser, 1919* Isonychus similis Frey, 1973*
Ceraspis squamulifera (Moser, 1919) Isonychus simulator Frey, 1969*
Chariodactylus sublaevicollis Moser, 1919* Macrodactylus bilineolatus Moser, 1919*
Clavipalpus peruanus Moser, 1918* Macrodactylus bistriatus Moser, 1919*
Clavipalpus spadiceus (Burmeister, 1855)* Macrodactylus brenskei Moser, 1918*
Ctilocephala asperula (Perty, 1830) Macrodactylus cinereus Blanchard, 1850*
Dicrania peruana Frey, 1972* Macrodactylus discipennis Moser, 1918*
Isonychus arbusticola Erichson, 1847* Macrodactylus marginicollis Moser, 1919*
Isonychus cervicapra Frey, 1965* Macrodactylus peruanus Moser, 1919*
Macrodactylus sapphirinus Moser, 1919* Plectris molesta (Kirsch, 1873)*
Macrodactylus sulcicollis Moser, 1919* Plectris penaella Frey, 1967*
Macrodactylus vittipennis Moser, 1919* Plectris penai Frey, 1967*
Plectris aberrans Frey, 1964* Plectris sculptipennis Frey, 1974*
Plectris candezei Frey, 1967 Plectris tenebrosa Frey, 1967*
Plectris kochi Frey, 1967 * Plectris tolimana (Moser, 1921)
Plectris lanata Frey, 1964* Plectris tuberculata (Moser, 1919)
Pachydemini
Diaphylla hispida Erichson, 1847*
Leuretra pectoralis Erichson, 1847
RUTELINAE
Anatistini
Spinochlamys macropus (Benderitter, 1921)* Spodochlamys latipes Arrow, 1946
Spodochlamys feyeri Ohaus, 1908 Spodochlamys peruvianus Soula, 2010*
Spodochlamys iheringi Ohaus, 1905
Rutelini
Acraspedon bernierei Soula, 2002* Lagochile delassisei Soula, 2005*
Acraspedon peruvianus Soula, 2002* Lagochile ebrardi Soula, 2010*
Aequatoria cludtsi Soula, 2002* Lagochile fuscoviridis Bouchard and Soula, 2005
Aequatoria davidi Soula, 2005* Lagochile peruana peruana Ohaus, 1898
Aequatoria lequericai Soula, 2006* Lagochile peruana huallagensis Soula, 2005*
Anticheiroides adamsii nevinsoni (Fowler, 1906)* Lagochile peruana occidentalis Soula, 2005*
Anticheiroides davidi Soula, 2006 Lagochile pottgensi Demez and Soula, 2010
Catoclastus chevrolatii Solier, 1851* Lagochile rodriguezi Soula, 2009*
Catoclastus jaumesi Soula, 2010* Lagochile santacruzis santacruzis (Machatschke,
Catoclastus rabinovichi Martnez, 1971* 1972)*
Chlorota callegariorum Soula, 2005* Lagochile santacruzis chanchomayoensis Soula,
Chlorota chaparroi Curoe and Soula, 2005* 2005*
Chlorota chavezlopezi Soula, 2006* Lagochile solimoensis solimoensis Ohaus, 1903
Chlorota columbica columbica (Ohaus, 1912) Lagochile solimoensis oberthuri Soula, 2005
Chlorota columbica peruana Soula, 2002* Lagochile solimoensis wadai Soula, 2005*
Chlorota nasuta Ohaus, 1905 Lagochile tibialis (Ohaus, 1935)*
Chlorota sergiocastroi Soula, 2008* Lagochile trigona trigona (Herbst, 1790)
Chlorota surinama surinama (Ohaus, 1898) Lagochile trigona mancocapaci Soula, 2005*
Chlorota surinama iquitosensis Soula, 2005* Lagochile trigona pozuzoensis Soula, 2005*
Chrysina argenteola (Bates, 1888) Lagochile vergaracobianae Soula and Curoe, 2005*
Chrysophora chrysochlora (Latreille, 1811) Lagochile vasseli Soula, 2010*
Cnemida retusa (Fabricius, 1801) Lagochile villatoroae villatoroae Soula, 2005*
Dorysthetus andicola Ohaus, 1905 Lagochile villatoroae tingomariaensis Soula, 2005*
Dorysthetus fulgida (Waterhouse, 1881) Lasiocala detingomaria Soula, 2006*
Dorysthetus mezai Soula, 2005* Lasiocala dioni Soula, 2006*
Dorysthetus peruanus (Ohaus, 1905) Lasiocala jenseni Soula, 2006*
Epichalcoplethis benjamini Bouchard and Soula, Lasiocala josei Soula, 2006*
2006 Lasiocala lamasi Soula, 2006*
Epichalcoplethis gilleti Soula, 2010 Lasiocala schmiti Soula, 2006*
Epichalcoplethis santistebani Bouchard and Soula, Lasiocala vasseli Soula, 2010*
2006* Macraspis andicola Burmeister, 1844
Epichalcoplethis schiffleri Bouchard and Soula, 2006* Macraspis assimilis Ohaus, 1908*
Eremophygus philippi Ohaus, 1910 Macraspis bicincta bicincta Burmeister, 1844
Exanticheira vidua (Ohaus, 1922) Macraspis chalcea Burmeister, 1844
Exothyridium filippii Soula, 2002* Macraspis chloraspis chloraspis Laporte, 1840
Exothyridium mercieri Soula, 2002* Macraspis chloraspis subandina Soula, 1998
Heterochlorota colini Soula, 2008* Macraspis chrysis (L., 1764)
Heterochlorota mathildae mathildae (Ohaus, 1908) Macraspis concoloripes ratcliffi (Soula, 2005)
Heterochlorota mathildae peruana Soula, 2002* Macraspis festiva Burmeister, 1844
Homonyx maurettei Soula, 2010 * Macraspis maculata maculata Burmeister, 1844
Homonyx peruanus Ohaus, 1913* Macraspis maculicollis Ohaus, 1905*
Homonyx zovii Demez and Soula, 2011 Macraspis martinezi martinezi Soula, 2003
Hypaspidius costatus (Burmeister, 1844)* Macraspis melanaria (Blanchard, 1850)
Lagochile aequatorialis aequatorialis Ohaus, 1898 Macraspis peruviana Ohaus, 1898*
Lagochile aequatorialis raimondii Soula, 2005* Macraspis pseudochrysis Landin, 1956
Lagochile amazona (Thunberg, 1822) Macraspis stirpita Ohaus, 1914*
Lagochile andicola andicola Ohaus, 1903 Macraspis testaceipennis Ohaus, 1898
Lagochile andicola condori Soula, 2005* Macraspis willersi Soula, 2010*
Lagochile anophrys (Ohaus, 1914) Macraspis xanthosticta Burmeister, 1844
Lagochile brunnea brunnea (Perty, 1830) Mecopelidnota arrowi Bates, 1904
Lagochile brunnea satipoensis Soula, 2005* Mecopelidnota dewynteri Soula, 2008*
Lagochile brunnea tenaensis Soula, 2005* Mecopelidnota marxi Soula, 2008
Lagochile brusteli Soula, 2005* Mecopelidnota mezai Soula, 2008*
Lagochile cachetica cachetica Ohaus, 1903 Mesomerodon spinipenne Ohaus, 1905
Lagochile cachetica fusciventris Ohaus, 1912* Microrutela campa (Ohaus, 1922)
Lagochile cachetica orientalis Soula, 2005 Microrutela ucalayiensis Jameson, 1997
Lagochile ciliata ciliata Ohaus, 1908 Minidorysthetus hoehnei (Ohaus, 1914)*
Minidorysthetus tingomariaensis Soula, 2006* Pseudohypaspidius silvestrei Soula, 2002*

Minidorysthetus ucayaliensis Soula, 2006* Pseudomacraspis affinis affinis (Laporte, 1840)
Minidorysthetus vandemergheli Soula, 1998* Pseudomacraspis affinis amazonica Soula, 2002
Oryctomorphus maculicollis Guerin-Meneville, 1838 Pseudomacraspis beryllina beryllina (Erichson, 1847)*
Parachlorota estebani Demez and Soula, 2010* Pseudothyridium (Pseudothyridium) bouchardi
Parachlorota josei Soula, 2005* Soula, 2002*
Parachlorota morettoi Soula, 2002* Pseudothyridium buckwaldi (Ohaus, 1912)
Paradorysthetus signatipennis signatipennis (Ohaus, Pseudothyridium ericki Soula, 2006*
1908)* Pseudothyridium hirtum (Kirsch, 1870)
Paraptenomela amazona tingomariaensis Soula, 2002* Pseudothyridium juanjosei Soula, 2006*
Paraptenomela opalescens (Ohaus, 1935)* Pseudothyridium minettii Soula, 2002*
Paratelaugis robusta robusta (Kirsch, 1871) Pseudothyridium oblongum oblongum (Ohaus, 1905)*
Pelidnota angiae Demez and Soula, 2009* Pseudothyridium quentini Soula, 2002*
Pelidnota bondili Soula, 2006* Ptenomela grangesi Soula, 2006*
Pelidnota brusteli Soula, 2010* Ptenomela tavakiliani Soula, 2002*
Pelidnota chlorana Erichson, 1847 Ptenomela toulgoeti Soula, 2006*
Pelidnota dobleri Frey, 1967 Rutela heraldica Perty, 1832
Pelidnota fusciventris fusciventris Ohaus, 1905* Rutela histrio Sahlberg, 1823
Pelidnota halleri Demez and Soula, 2010* Rutela histrioparilis Jameson, 1997
Pelidnota hernanlequericai Soula, 2006* Rutela laeta (Weber, 1801)
Pelidnota hoefigi Ohaus, 1912* Rutela lineola (Linnaeus, 1767)
Pelidnota incerta Soula, 2006* Rutela tricolorea (Ohaus, 1905)
Pelidnota injantepalominoi Demez and Soula, 2010* Sorocha bousqueti Soula, 2006*
Pelidnota lacazei Soula, 2010* Sorocha carloti Demez and Soula, 2010*
Pelidnota mezai Soula, 2009* Sorocha castroi Soula, 2008*
Pelidnota neitamorenoi neitamorenoi (Soula, 2006) Sorocha champenoisi Soula, 2006*
Pelidnota neitamorenoi rodriguezdemendozaensis Sorocha chapellei Demez and Soula, 2010*
Soula, 2010* Sorocha damasoi Soula, 2006*
Pelidnota ohausi ohausi Frey, 1976 Sorocha jeanmaurettei Demez and Soula, 2010*
Pelidnota ohausi piurensis (Soula, 2006)* Sorocha lamasi lamasi Soula, 2006*
Pelidnota peslieri Soula, 2009* Sorocha lamasi satipoensis Soula, 2006 *
Pelidnota polita (Latreille, 1811) Sorocha maylini Soula, 2006*
Pelidnota porioni Soula, 2010* Sorocha similis (Ohaus, 1908)*
Pelidnota satipoensis Demez and Soula, 2010* Sorocha yelamosi Soula, 2010*
Pelidnota schneideri Soula, 2010* Telaugis aenescens aenescens Burmeister, 1844
Pelidnota subandina subandina Ohaus, 1905 Telaugis aenescens subandina Soula, 1998*
Pelidnota testaceovirens felipemezai Soula, 2006* Theuremaripa buchei (Soula, 2002)*
Pelidnota toulgoeti Soula, 2006* Theuremaripa imitatrix (Ohaus, 1903)
Pelidnota uncinata Ohaus, 1930 Theuremaripa meyeri (Soula, 2005)*
Pelidnota unicolor unicolor (Drury, 1778) Thyriochlorota lassalei Soula, 2002*
Pelidnota unicolor subandina Soula, 2009* Thyriochlorota villosa (Ohaus, 1908)*
Pelidnota werneri Soula, 2006* Tipicha champanheti Soula, 2002*
Pelidnota zovii Soula, 2010* Tipicha joliveti Soula, 2002*
Pseudochlorota peruana Ohaus, 1905* Vayana melzeri melzeri Ohaus, 1928
Pseudohypaspidius antoinei Soula, 1998* Vayana melzeri subandina Soula, 1998*
Anomalini
Callistethus aequatorialis huanapensis (Ohaus, Callistethus suratus (Burmeister, 1844)*
1908)* Callistethus tricostatus (Ohaus, 1897)
Callistethus antis (Ohaus, 1903)* Paranomala cincta viridicollis Burmeister, 1844
Callistethus cicatricosa (Perty, 1832) Paranomala hylobia (Ohaus, 1897)*
Callistethus eckhardti (Ohaus, 1897)* Paranomala inconstans Burmeister, 1844
Callistethus kulzeri (Frey, 1968)* Paranomala undulata peruviana Guerin-Meneville,
Callistethus marginatus (Fabricius, 1792) 1838*
Callistethus penai (Frey, 1968)* Strigoderma marginata (Olivier, 1789)
Callistethus pyritosus (Erichson, 1847)* Strigoderma peruviensis Blanchard, 1850
Callistethus rufomicans (Ohaus, 1897) Strigoderma sulcipennis sumtuosa Burmeister, 1844
Anoplognathini
Platycoelia abdominalis Ohaus, 1904* Platycoelia insolita Smith, 2003*
Platycoelia aenigma Smith, 2003* Platycoelia kirschi (Ohaus, 1904)*
Platycoelia alternans Erichson, 1847 Platycoelia laelaps (Gutierrez, 1951)*
Platycoelia baessleri (Ohaus, 1904)* Platycoelia lutescens Blanchard, 1851
Platycoelia burmeisteri Arrow, 1899 Platycoelia marginata Burmeister, 1844
Platycoelia convexa Smith, 2003 Platycoelia peruviana Smith, 2003
Platycoelia flavostriata (Latreille, 1813) Platycoelia pomacea Erichson, 1847
Platycoelia gaujoni Ohaus, 1904 Platycoelia prasina Erichson, 1847
Platycoelia helleri (Ohaus, 1904) Platycoelia rufosignata Ohaus, 1904
Platycoelia inca Smith, 2003* Platycoelia selanderi Martnez and Martnez, 1994
Platycoelia inflata Ohaus, 1904
Geniatini
Bolax albopilosa Ohaus, 1917* Leucothyreus lazarus Ohaus, 1918
Bolax andicola Burmeister, 1844* Leucothyreus saparus Ohaus, 193*
Bolax boliviensis Ohaus, 1898 Lobogeniates bicolor Ohaus, 1917
Bolax cupreoviridis Ohaus, 1931* Trizogeniates aphilus Villatoro, 2002*
Bolax disgamia Ohaus, 1917* Trizogeniates apicalis Ohaus, 1917*
Bolax glabripennis Ohaus, 1917* Trizogeniates barrerai Martnez, 1965*
Bolax gonzalofideli Soula, 2010* Trizogeniates caiporae Villatoro, 2002
Bolax incogitata Dohrn, 1883 Trizogeniates catsus Villotoro, 2002
Bolax malkini Soula, 2010* Trizogeniates crispospinatus Villatoro, 2002*
Bolax nigriceps Ohaus, 1917* Trizogeniates laticollis Ohaus, 1931
Bolax robackeri Soula, 2010* Trizogeniates ohausi Villatoro, 2002
Bolax rutila Erichson, 1847* Trizogeniates planipennis Ohaus, 1917
Trizogeniates temporalis Ohaus, 1917
Geniates balzapamae Ohaus, 1917
Trizogeniates tibialis Ohaus, 1917
Leucothyreus baeri Ohaus, 1917*
Trizogeniates trivittatus Ohaus, 1917
Leucothyreus demetrius Ohaus, 1918*
DYNASTINAE
Cyclocephalini
Acrobolbia macrophylla Ohaus, 2012 Cyclocephala brevis Hohne, 1923
Ancognatha castanea Erichson, 1847 Cyclocephala colasi Endrodi, 1964
Ancognatha erythrodera (Blanchard, 1841) Cyclocephala confusa Endrodi, 1966
Ancognatha humeralis Burmeister, 1847 Cyclocephala contracta Kirsch, 1873
Ancognatha lutea Erichson, 1847 Cyclocephala couturieri Dechambre, 1999
Ancognatha scarabaeoides Erichson, 1847 Cyclocephala dilatata (Prell, 1934)
Ancognatha vulgaris Arrow, 1911 Cyclocephala diluta Erichson, 1847
Aspidolea brunnea Hohne, 1922 Cyclocephala discolor (Herbst, 1792)
Aspidolea collaris Endrodi, 1965* Cyclocephala dispar (Herbst, 1792)
Aspidolea fuliginea (Burmeister, 1847) Cyclocephala flavoscutellaris Hohne, 1923
Cyclocephala flora Arrow, 1911
Aspidolea laticeps Harold, 1869
Cyclocephala freyi Endrodi, 1964
Aspidolea lindae Ratcliffe, 1978
Cyclocephala fulgurata Burmeister, 1847
Aspidolea mimethes (Hohne, 1922)*
Cyclocephala fulvipennis Burmeister, 1847
Aspidolea notaticollis Hohne, 1922
Cyclocephala genieri Joly, 2010*
Aspidolea singularis Bates, 1888
Cyclocephala goetzi Endrodi, 1966*
Aspidolea suturalis Hohne, 1922 Cyclocephala guycolasi Dechambre, 1992
Augoderia freyi Endrodi, 1976* Cyclocephala hirsuta Hohne, 1923
Chalepides paradytis Ponchel and Dechambre, 2003 Cyclocephala inca Endrodi, 1966
Cyclocephala affinis Endrodi, 1966 Cyclocephala isabellina Hohne, 1923
Cyclocephala almitana Dechambre, 1992 Cyclocephala kaszabi Endrodi, 1964
Cyclocephala altamontana Dechambre, 1999 Cyclocephala ligyrina Bates, 1888
Cyclocephala amazona (L., 1767) Cyclocephala lineigera Hohne, 1923
Cyclocephala bicolor Laporte,1840 Cyclocephala liomorpha Arrow, 1911
Cyclocephala lunulata Burmeister, 1847 Cyclocephala spilopyga Erichson, 1847

Cyclocephala macrophylla Erichson, 1847 Cyclocephala stictica Burmeister, 1847
Cyclocephala mannheimsi Endrodi, 1964 Cyclocephala testacea Burmeister, 1847
Cyclocephala marginalis Kirsch,1870 Cyclocephala tronchonii Martnez, 1975*
Cyclocephala mecynotarsis Hohne, 1923 Cyclocephala tylifera Hohne, 1923
Cyclocephala melanocephala (Fabricius, 1775) Cyclocephala verticalis Burmeister, 1847
Cyclocephala molesta Endrodi, 1969 Cyclocephala viridis Dechambre, 1982
Cyclocephala moreti Dechambre, 1992 Cyclocephala zurstrasseni Endrodi, 1964*
Cyclocephala morphoidina Prell, 1937 Dyscinetus dubius (Olivier, 1798)
Cyclocephala munda Kirsch, 1870 Dyscinetus dytiscoides (Arrow, 1911)
Cyclocephala obscura Endrodi, 1966* Dyscinetus olivaceus Hohne, 1923
Cyclocephala ocellata Bolvar y Pieltain, Jimenez- Dyscinetus paradytis Ponchel and Dechambre, 2003
Asua, and Martnez, 1963 Erioscelis peruana Saylor, 1946*
Cyclocephala octopunctata Burmeister, 1847 Erioscelis proba (Sharp, 1877)
Cyclocephala panthera Dechambre, 1979 Harposcelis paradoxus Burmeister, 1847
Cyclocephala paraflora Martnez, 1978 Mimeoma signatoides Hohne, 1923
Cyclocephala paraguayensis Arrow, 1903 Stenocrates bicarinatus Robinson, 1948
Cyclocephala peruana Endrodi, 1966* Stenocrates carbo Prell, 1937
Cyclocephala pilosa Dupuis, 2006* Stenocrates celatus Prell, 1937
Cyclocephala prolongata Arrow, 1902 Stenocrates clipeatus Endrodi, 1966
Cyclocephala pugnax Arrow, 1914 Stenocrates cognatus Endrodi, 1966
Cyclocephala quadripunctata Hohne, 1923 Stenocrates cultor Burmeister, 1847
Cyclocephala rufovaria Arrow, 1911 Stenocrates haackae Ratcliffe, 1979
Cyclocephala rustica municipalis Hohne, 1923 Stenocrates holomelanus (Germar, 1824)
Cyclocephala saltini Ratcliffe, 2008 Stenocrates minutus Endrodi, 1966
Cyclocephala scarabaeina (Gyllenhal, 1817) Stenocrates nasutus Dechambre, 1979*
Cyclocephala sexpunctata Laporte, 1840 Stenocrates popei Endrodi, 1971
Cyclocephala simulatrix Hohne, 1923
Pentodontini
Bothynus entellus (LePeletier and Serville, 1828) Oxyligyrus peruanus Endrodi, 1966
Bothynus lancifer Dechambre, 1981* Parapucaya amazonica Prell, 1934
Diloboderus abderus (Sturm, 1826) Parapucaya nodicollis (Kirsch, 1873)
Heteroglobus obesus Dupuis and Dechambre, 2008* Pentodina peruviana Endrodi, 1968
Hylobothynus obesus Ohaus, 1910 Piscoperus paracanicola Ratcliffe and Giraldo, 2014*
Oxyligyrus contractus Dupuis, 2010* Tomarus bituberculatus (Palisot de Beauvois, 1805)
Tomarus burmeisteri (Steinheil, 1872) Tomarus maternus (Prell, 1937)
Tomarus ebenus (DeGeer, 1774) Tomarus peruvianus (Endrodi, 1970)
Tomarus gyas Erichson, 1847 Tomarus similis (Endrodi, 1968)
Tomarus maimon Erichson, 1847 Tomarus villosus (Burmeister, 1847)
Oryctini
Coelosis biloba (L., 1767) Heterogomphus ulysses Burmeister, 1847
Enema pan (Fabricius, 1775) Megaceras brevis Dechambre, 1999
Heteroglobus obesus Dupuis and Dechambre, 2008 Megaceras briansaltini Ratcliffe, 2007*
Heterogomphus arrowi Prell, 1912* Megaceras endroedii Dechambre, 1998
Heterogomphus dilaticollis Burmeister, 1847 Megaceras inflatum Prell, 1934
Heterogomphus hirticollis Prell, 1912* Megaceras laevipenne Prell, 1914
Heterogomphus hirtus Prell, 1912 Megaceras morpheus Burmeister, 1847
Heterogomphus incornutus Prell, 1912* Megaceras philoctetes (Olivier, 1789)
Heterogomphus mirabilis Prell, 1912* Megaceras porioni Dechambre, 1981
Heterogomphus ochoai Martnez, 1966 Megaceras quadraticollis Dechambre, 1975
Heterogomphus orsilochus Erichson, 1847 Podischnus oberthueri Sternberg, 1907
Heterogomphus peruanus Endrodi, 1976* Podischnus sexdentatus (Taschenberg, 1870)
Heterogomphus pilosus Dechambre, 1998 Strategus aloeus (L., 1758)
Heterogomphus porioni Dechambre, 1998 Strategus jugurtha Burmeister, 1847
Heterogomphus rugicollis Prell, 1912 Strategus surinamensis hirtus Sternberg, 1910
Heterogomphus rubripennis Prell, 1912
Phileurini
Amblyodus castroi Grossi and Grossi, 2011 Homophileurus waldenfelsi Endrodi, 1978
Amblyoproctus chalumeaui Endrodi, 1977 Microphileurus caviceps Kolbe, 1910
Amblyoproctus piliger (Perty, 1830) Microphileurus subulo Prell, 1912*
Amblyoproctus rugosus (Erichson, 1847) Oryctophileurus armicollis Prell, 1911
Archophileurus aper Endrodi, 1977 Palaeophileurus carbo Ratcliffe, 2002
Archophileurus burmeisteri (Arrow, 1908) Palaeophileurus erebus Ratcliffe, 2002
Archophileurus oedipus (Prell 1912) Palaeophileurus marcusoni Ratcliffe, 1998
Archophileurus peruanus Endrodi, 1977 Palaeophileurus ocampoi Neita and Ratcliffe,
Archophileurus sus Dechambre, 2006* 2012
Hemiphileurus brasiliensis Endrodi, 1978 Palaeophileurus proximus Dechambre, 1997
Hemiphileurus depressus (Fabricius, 1801) Palaeophileurus sclateri (Bates, 1887)
Hemiphileurus elongatus Dupuis and Dechambre, Phileucourtus bicornutus Dechambre, 2008*
2000* Phileurus angustatus Kolbe, 1910
Hemiphileurus howdeni Endrodi, 1978* Phileurus didymus (L., 1758)
Hemiphileurus isabellae Dupuis, 2004* Phileurus excavatus Prell, 1911
Hemiphileurus kahni Dupuis and Dechambre, Phileurus kaszabi Endrodi, 1978
2000* Phileurus valgus (Olivier, 1789)
Homophileurus quadrituberculatus (Palisot de
Beauvois, 1805)
Agaocephalini
Aegopsis chaminadei Dechambre, 2000 Brachysiderus quadrimaculatus Waterhouse, 1881
Aegopsis peruvianus Arrow, 1941* Mitracephala humboldti Thomson, 1859
Dynastini
Dynastes hercules (L., 1758) Golofa eacus Burmeister, 1847
Dynastes neptunus (Quensel, 1806) Golofa spatha Dechambre, 1989
Golofa aegeon (Drury, 1773) Golofa testudinarius (Prell, 1934)*
Golofa clavigera (L., 1771) Golofa unicolor (Bates, 1891)
CETONIINAE
Cetoniini
Euphoria steinheili Janson, 1878
Gymnetini
Desicasta lobata (Olivier, 1789) Gymnetis pantherina (Blanchard, 1843)
Desicasta purpurascens (Schoch, 1898) Gymnetis rufilateris (Illiger, 1800)
Gymnetis balzarica Janson, 1880 Gymnetis subpunctata Westwood, 1874 (5? G. varia-
Gymnetis coturnix Burmeister, 1842 (5? G. phasia- bilis Moser, 1921)
nus Burmeister, 1842) Hoplopyga liturata (Olivier, 1789)
Gymnetis mathani Pouillaude, 1913 Hoplopyga peruana Moser, 1912
Gymnetis holoserica Voet, 1778 (5? G. chancha- Marmarina maculosa (Olivier, 1789)
mayensis Pouillaude, 1913)
Gymnetis pardalis (Gory and Percheron, 1833)
(5? G. cupriventris Janson, 1880)
Cremastocheilini
Cyclidius lacordairei Thomson, 1860*
Genuchinus sp.
Trichiini
Golinca davisii Waterhouse, 1877*
Golinca ishiharai Nagai, 1994*
Acknowledgments
We are grateful to Caroline Chaboo (University of Kansas) for inviting us to
participate in the Beetles of Peru project and affording us the opportunity for
advancing scarab research in Peru. We thank Trond Larsen (Conservation
International, Arlington, VA) for sharing his records of Scarabaeinae from Peru.
This work was supported, in part, by a Biotic Surveys and Inventory Grant from the
National Science Foundation (DEB-0716899) to B. C. Ratcliffe and R. D. Cave. We
acknowledge NSF-EPSCoR #66928 (PI: CS Chaboo) and the University of Kansas
Department of Ecology and Evolutionary Biology-General Research Fund (PI: CS
Chaboo) for supporting the Beetles of Peru project.
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Clambidae, Eucinetidae, Scirtidae (Scirtoidea)
Author(s): Caroline S. Chaboo and William D. Shepard

terms_of_use.
critical research.
88(2), 2015, pp. 208210
Beetles (Coleoptera) of Peru: A Survey of the Families. Clambidae,

Eucinetidae, Scirtidae (Scirtoidea)
CAROLINE S. CHABOO1 AND WILLIAM D. SHEPARD2,3
Clambidae Fischer, 1821

Recognition: Clambids are easily recognized by their tiny oval-shaped bodies (12
mm), ability to roll into a ball, and expanded hind coxal plates. The head is deflexed
and the antenna has a 2-segmented club. The tarsal formula is 4-4-4 and the hind
wings are fringed with setae.
Habitat: Clambids have been found in decaying vegetable matter of many kinds
and they are known to feed on various fungi.
Notes: Blackwelder (19441957) did not report this family from Peru. Leschen
(2005a) is a good introduction to the diversity (5 gen/,150 species world-wide),
biology and morphology. Clambids are often collected at lights at night and from
leaf litter using Berlese funnels. Adults fly in the afternoon and at dusk and can be
collected using flight-intercept traps. The following list is derived from Endrody-
Younga (1998).
Checklist:
Clambus impressipennis Endrody-Younga, 1981 Clambus semicaecus Endrody-Younga, 1998
Clambus inca Endrody-Younga, 1998 Clambus signatus Endrody-Younga, 1981
Clambus mirandanus Endrody-Younga, 1998 Clambus uncus Endrody-Younga, 1998
Clambus peruanus Endrody-Younga, 1998
Eucinetidae Lacordaire, 1857

Recognition: This small family is easy to recognize by the teardrop-shaped habitus,
the mouthparts directed posteriorly in repose, and the large metacoxal plates capable
of mostly concealing the saltatorial hind legs. The tarsal formula is 5-5-5.
Habitat: Eucinetids are associated with various groups of fungi and the
microhabitats that support those fungi, such as rotting logs, leaf litter, humus, debris
piles, etc. Some fly readily.
Notes: This family has not been reported from Peru previously. Leschen (2005b) is
a handy recent overview of this very small family of nine genera and ,40 species. We
have seen specimens from Madre de Dios that key out to Eucinetus subcrispus Vit using
Vit (1990). However, that species is known only from Trinidad so we have reservations
about the determination. Specimens can be collected using flight-intercept traps and
1
2
3
Accepted 16 February 2014; Revised 16 August 2015
Berlese funnels, sifting litter and spraying fungal fruiting bodies with pyrethrin-based
insecticides.
Checklist:
Eucinetus undetermined species (M. L. Gimmel, personal observation)
Scirtidae Fleming, 1821

Recognition: Adult scirtids are recognized by their oval, 24 mm long, soft body,
their deflexed head, their conical procoxae and their short, broad pronotum. Many
adults have contrasting color patterns. Larvae have long antennae with numerous
antennomeres, a character unique among all beetle larvae.
Habitat: Scirtids are commonly called marsh beetles, because most adult scirtids
are found on vegetation around water. Larvae are mainly aquatic and can be found
in vegetation and debris in streams and ponds. Some larvae occur in phytotelmata
and tree holes (Lawrence, 2005).
Notes: Blackwelder (19441957) did not report this family from Peru. Lawrence
(2005) is a good introduction to this small family of about 30 genera and 800 species
world-wide. The following checklist of Peruvian Scirtidae was derived from Spangler
(1966), who reported the collection of four larvae representing an undetermined
species of Cyphon. Adult scirtids are mainly collected by sweeping riparian vegetation
and at lights. Many have enlarged femora and can jump upon disturbance.
Checklist:
Cyphon undetermined species (Spangler, 1966) (Identification made in 1966!). Recently Cyphon has been
replaced by the genus Contacyphon Gozis, 1886, but even that genus is in the process of being split into
several genera (see Zwick et al., 2013).
Acknowledgements
Evolutionary Biology-General Research Fund (PI: CS Chaboo). We are grateful to
Matthew Gimmel and two anonymous reviewers for help in improving the manuscript.
Literature Cited
Blackwelder, R. E. 19441957. Checklist of the Coleopterous insects of Mexico, Central America, the
West Indies, and South America. Parts 1 through 6. U. S. Natl. Mus. Bull. 185: ixii, 11492.
Endrody-Younga, S. 1998. The genus Clambus in Central and South America (Coleoptera: Eucinetoidea:
Clambidae). Annals of the Transvaal Museum 36:373403.
Lawrence, J. F. 2005. Scirtidae Fleming, 1821. In R. G. Beutel and R. A. B. Leschen (eds.), Coleoptera,
Beetles. Vol. 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga
partim), pp. 443450. Handbook of Zoology. Walter de Gruyter, Berlin and New York.
Leschen, R. A. B. 2005a. Clambidae Fischer, 1821. In R. G. Beutel and R. A. B. Leschen (eds.)
Coleoptera, Beetles. Vol. 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga,
Polyphaga partim), pp. 438443. Handbook of Zoology. Walter de Gruyter, Berlin and New York.
Leschen, R. A. B. 2005b. Eucinetidae Lacordaire, 1857. In R. G. Beutel and R. A. B. Leschen (eds.)
Coleoptera, Beetles.Vol. 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga,
Polyphaga partim), pp.433438. Handbook of Zoology. Walter de Gruyter, Berlin and New York.
Spangler, P. J. 1966. XIII Aquatic Coleoptera (Dytiscidae; Noteridae; Gyrinidae; Hydrophilidae;

Dascillidae; Helodidae; Psephenidae; Elmidae). In R. Patrick et al.(eds.) The Catherwood
Foundation Peruvian-Amazon Expedition. Limnological and Systematic Studies, pp, 377443.
Monographs of the Academy of Natural Sciences of Philadelphia 14:1495.
Vit, S. 1990. Revision des especes neotropicales du genre Eucinetus Germar (Coleoptera: Eucinetidae). Le
Naturaliste Canadien 117:103122.
Zwick, P., B. Klausnitzer, and R. Ruta. 2013. Contacyphon. Gozis, 1886 removed from synonymy
(Coleoptera. Scirtidae) to accommodate species so far combined with the invalid name Cyphon
Paykull, 1799. Entomologische Blatter und Coleoptera 109:337353.
Buprestidae
Author(s): Henry Hespenheide and Caroline S. Chaboo

terms_of_use.
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88(2), 2015, pp. 211214
Beetles (Coleoptera) of Peru: A Survey of the Families. Buprestidae

HENRY HESPENHEIDE1 AND CAROLINE S. CHABOO2,3
Buprestidae
Diversity in Peru: 4 subfamilies, 12 tribes, 29 genera, 110 species and 3 subspecies.
Recognition: Buprestidae are commonly called jewel beetles because of the
spectacular colors of the adults of many species. Most buprestids are small to
medium-sized beetles, although Euchroma gigantea is one of the largest Neotropical
insects (55 mm). Adults display two extremes of morphology and ecology: members
of one group are small (26 mm), rhomboidal, dorso-ventrally flattened, and mine
leaves as larvae; the rest of the family are borers in wood or soft tissues and are larger
(435 mm), elongate, and cylindrical or moderately flattened. Adults of many species
are strongly shining and brightly metallic in coloration, a few are more or less
covered by waxy yellow powder, and some have patterns formed by setae. Mimicry
occurs in adults of both leaf-miners (models include chrysomelids and coccinellids;
Hespenheide, 2014) and wood-borers (models include ants, flies and wasps;
Hespenheide, 1986, 2010, 2012). Larvae are typically flattened with a large
prothorax.
Habitat: Members of the family occur primarily at low- to mid-elevations, below
20003000 m. Buprestids are most common in primary forest canopies, at forest
edges, or in secondary growth, although a few species are found in open habitats on
weeds and sedges. Adult behavior is related to larval feeding type. Wood-boring
species are more numerous in areas of lower rainfall and a well-defined dry season
and are usually found running on the trunks and branches of recently dead trees or
branches, or feeding on leaves of their larval hosts. Adults are most active during the
sunny weather during midday, and they fly readily, making them difficult to capture.
Leaf-mining species are more common in areas with higher rainfall, and adults
usually feed on the leaves of the larval host. Larval mines are usually full-depth and
may be either serpentine or blotch-like in form, depending on the species. Except for
some of the larger wood-boring species, most buprestids of both ecological types are
apparently relatively narrowly host-specific and many appear to exist in very low
populations. Many are associated with woody dicots, especially legumes, although
leaf-miners are known from ferns and such monocots as sedges (Cyperaceae) and
palms (Arecaceae).
Pest species in Peru: Very few tropical species appear to be important
agriculturally. Leaf-mining species (Leiopleura spp.) are associated with cassava or
yuca (Manihot esculenta). The invasive Aphanisticus cochinchinae seminulum likely
occurs on sugar cane, but neither appears to have much effect on the plants.
Chrysobothris species have been reported as pests in plantations of woody plants.
1
Department of Ecology and Evolutionary Biology, University of California, Box 951606, Los Angeles,
California 90095-1606.
2
3
Accepted 02 December 2014; Revised 02 February 2015
Notes: The following checklist of Peruvian Buprestidae is taken from Bellamy

(2008), with additions by Moore and Manuel Dieguez (2008, 2010, 2014), Moore and
Lander (2010), and Bly (2013). Blackwelder (1944) recorded 81 species and 3
subspecies from Peru. Additional species records from Peru were obtained through
identification of material by HH from the expeditions of CSC (deposited in SEMC).
These represent either undescribed species or additional species records in unrevised
genera of Buprestidae, and so are simply designated sp. 1, sp. 2, etc. Although
only 129 species are listed here, there are probably many hundreds of species in the
fauna of Peru, many of which are undescribed.
Checklist:
Agrilinae: Coraebini
Dismorpha tenuis (Kirsch, 1873)
Agrilinae: Agrilini
Agrilus atahualpa Obenberger, 1935 Agrilus sp. 3
Agrilus augustus Obenberger, 1933 Agrilus sp. 4
Agrilus cordillerae Kirsch, 1873 Agrilus sp. 5
Agrilus huascar Obenberger, 1932 Agrilus sp. 6
Agrilus maculatus Gory & Laporte, 1837 Paragrilus nigritus Kerremans, 1899
Agrilus hilaris Waterhouse, 1889 Paragrilus peruvianus (Gory & Laporte, 1837)
Agrilus sp. 1 Paragrilus punctifrons Obenberger, 1924
Agrilus sp. 2
Agrilinae: Tracheini
Brachys carinicollis Kirsch, 1873 Pachyschelus virens (Kirsch, 1873)
Brachys occidentalis Obenberger, 1923 Pachyschelus viridescens (Kirsch, 1873)
Brachys sp. 1 Pachyschelus viridulus (Kirsch, 1873)
Brachys sp. 2 Euhylaeogena cognata (Kirsch, 1873)
Taphrocerus tenuis (Kirsch, 1873) Euhylaeogena compar (Kirsch, 1873)
Taphrocerus sp. 1 Euhylaeogena lata (Kirsch, 1873)
Taphrocerus sp. 2 Euhylaeogena nana (Kirsch, 1873)
Taphrocerus sp. 3 Euhylaeogena planifrons (Kirsch, 1873)
Taphrocerus sp. 4 Euhylaeogena sp. 1
Taphrocerus sp. 7 Lius bicolor (Kirsch, 1873)
Taphrocerus sp. 8 Lius emarginatus (Kirsch, 1873)
Taphrocerus depilis Kerremans, 1896 Lius observans (Kirsch, 1873)
Taphrocerus finitimus Obenberger, 1924 Callimicra coraeboides Kerremans, 1899
Pachyschelus aeneicollis (Kirsch, 1873) Callimicra ignobilis (Kirsch, 1873)
Pachyschelus alatus Kerremans, 1899 Callimicra occidentalis Obenberger, 1922
Pachyschelus austerus (Kirsch, 1873) Callimicra peruviana Kerremans, 1899
Pachyschelus brevis (Kirsch, 1873) Leiopleura aerate (Kirsch, 1873)
Pachyschelus sp. 1 cf collaris Gory & Laporte, 1840 Leiopleura cincta Kerremans, 1899
Pachyschelus sp. 1 cf nigricollis Waterhouse, 1889 Leiopleura colorata Kerremans, 1899
Pachyschelus erichsoni Saunders, 1871 Leiopleura inca Kerremans, 1899
Pachyschelus gemellus Kerremans, 1899 Leiopleura inornata (Kirsch, 1873)
Pachyschelus grassator Obenberger, 1925 Leiopleura limae Obenberger, 1936
Pachyschelus inca Kerremans, 1899 Leiopleura nigrocoerulea Obenberger, 1917
Pachyschelus jucundus (Kirsch, 1873) Leiopleura nobilis (Kirsch, 1873)
Pachyschelus kerremansi Obenberger, 1925 Leiopleura peruviae Obenberger, 1936
Pachyschelus occidentalis Kerremans, 1899 Leiopleura peruviana Kerremans, 1899
Pachyschelus peruvianus Kerremans, 1899 Leiopleura tristis (Kirsch, 1873)
Pachyschelus rambouseki Obenberger, 1925 Leiopleura sp. 1
Pachyschelus rotundatus Kerremans, 1889 Leiopleura sp. 2
Buprestinae: Buprestini
Buprestis rufipes Olivier, 1790
Buprestinae: Anthaxini
Bilyaxia concinna (Mannerheim, 1837)
Buprestinae: Chrysobothrini
Colobogaster bella Kirsch, 1873 Chrysobothris sp. 1
Colobogaster peruviana Obenberger, 1924 Chrysobothris sp. 2
Colobogaster weingaertneri Hoscheck 1931 Chrysobothris sp. 3
Chrysobothris banghaasi Thery, 1911 Chrysobothris sp. 4
Chrysobothris hypochloris Erichson, 1847 Chrysobothris sp. 5
Chrysobothris peruviae Obenberger, 1924
Buprestinae: Actenodini
Actenodes chalybaeitarsis (Chevrolat, 1834) Actenodes nobilis peruanicus Obenberger, 1932
Actenodes nobilis nobilis (Linnaeus, 1758) Actenodes sp. nr nobilis (Linnaeus, 1758)
Buprestinae: Stigmoderini
Conognatha rogersii Saunders, 1872
Lasionota espanoli (Cobos), 1958
Chrysochroinae: Paraleptodemini
Chrysesthes sp. 1 Pelecopselaphus blandus peruvianus Thery, 1911
Halecia deliciosa Kerremans, 1919 Pelecopselaphus blandus blandus (Fabricius, 1781)
Halecia simplex Kirsch, 1873 Euchroma giganteum inca Obenberger, 1928
Halecia parallela Saunders, 1874 Euchroma giganteum peruanum Obenberger, 1928
Halecia simplex Kirsch, 1873
Chrysochroinae: Dicercini
Ectinogonia (Kheiliella) bellamyi Moore & Dieguez, Pseudolampetis fasciata (Kerremans, 1919)
2014 Pseudolampetis cincta (Kerremans, 1897)
Pseudolampetis cincta (Kerremans, 1897) Pseudolampetis circumsulcata (Laporte & Gory,
Lampetis peruana (Obenberger, 1928) 1837)
Pseudolampetis bilineata (Latreille, 1813) Pseudolampetis plagiata (Kerremans, 1919)
Pseudolampetis circumsulcata (Laporte & Gory, Pseudolampetis soukupi Obenberger, 1939
1837)
Polycestinae: Polycestini
Polycesta bicolor Kerremans, 1897
Polycestinae: Haplostethini
Mastogenius peruvianus Fisher, 1949
Mastogenius sp. 1
Polycestinae: Perucolini
Perucola picturata Thery, 1925
Acknowledgements
publication. We thank Matthew Gimmel for help assembling some names and two
anonymous reviewers for comments that improved the manuscript.
Literature Cited
Bellamy, C. L. 1997. Buprestoidea. Version 01 January 1997 (under construction). URL: http://tolweb.
org/Buprestoidea/9082/1997.01.01. In The Tree of Life Web Project, URL: http://tolweb.org/ (Last
accessed 7 October 2014).
Bellamy, C. L. 2008. A World Catalogue and Bibliography of the Jewel Beetles (Coleoptera:
Buprestoidea). Vol. 2: Chrysochroinae: Sphenopterini through Buprestinae: Stigmoderini. Pensoft
Series Faunistica No. 77. Pensoft Publishers, Sofia.
Bellamy, C. L. 2014. The World of Jewel Beetles (Insecta: Coleoptera: Buprestidae). URL: http://
coleopsoc.org/buprestidae/ (Last accessed 7 October 2014).
Bly, S. 2013. A study on the Neotropical Anthaxiini (Coleoptera, Buprestidae, Buprestinae). ZooKeys
304:1747.
Blackwelder, R. E. 1946. Checklist of the Coleopterous insects of Mexico, Central America, the West
Indies, and South America part 4. Smithsonian Institution United States National Museum
Bulletin 185, United States Government Printing Office, Washington, DC. 551763 pp.
Hespenheide, H. A. 1986. Mimicry of ants of the genus Zacryptocerus (Hymenoptera: Formicidae). J. N. Y.
Entomol. Soc. 94:394408.
Hespenheide, H. A. 2010. New Agrilus Curtis (Coleoptera: Buprestidae) from Mexico and Costa Rica
mimicking parasitic wasps. Zootaxa 2545:3946.
Hespenheide, H. A. 2012. New Mexican and Central American species of Agrilus Curtis (Coleoptera:
Buprestidae) mimetic of flies. Zootaxa 3181:127.
Hespenheide, H. A. 2014. A reconsideration of Hylaeogena Obenberger, 1923 (Coleoptera: Buprestidae),
with descriptions of a new genus and new species from Mexico and Central America. Coleopts.
Bull. 68:2130.
Manley, G. V. 1986. Mastogenius Sol. (Coleoptera: Buprestidae) of Ecuador and Peru west of the Andean
Mountains with descriptions of five new species. Coleopts. Bull. 40:223231.
Moore, T., and T. Lander. 2010. Revision du Genre Conognatha. Edition Magellanes 24:172 pp.
Moore, T., and V. M. Manuel Dieguez. 2008. Aporte al conocimiento del genero Polycesta Dejean, 1833
en las Regiones Neotropicales y Andina, con descripcion especies nuevas (Coleoptera: Buprestidae:
Polycestini). Acta entomologica Chilena 32:722.
Moore, T., and V. M. Manuel Dieguez. 2010. Aporte al conocimiento del genero Dactylozodes Chevrolat,
1838 de Sudamerica (Coleoptera: Buprestidae). Revista peruana entomologica 46:914.
Moore, T., and V. M. Manuel Dieguez. 2014. A contribution to the knowledge of the Neotropical
Buprestidae: description of a new species of Ectinogonia Spinola (Coleoptera: Buprestidae:
Chrysochroinae: Dicercini). Coleopts. Bull. 68:4749.
Mauroniscidae Majer, 1994
Author(s): Adriean J. Mayor and Caroline S. Chaboo

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88(2), 2015, pp. 215216

Mauroniscidae Majer, 1994
ADRIEAN J. MAYOR1 AND CAROLINE S. CHABOO2,3

Recognition: Adults of the family Mauroniscidae are similar to melyrids of the
subfamily Dasytinae, but recognized by their small size, usually 34 mm; color, often
with elytra and legs pale; head and mouthparts of some narrowed and elongated;
antennae often with terminal antennomeres enlarged forming a loose club; and tarsal
claws not toothed and lacking membranous appendages. Adults are not sexually
dimorphic and the sexes are very difficult to distinguish. Larvae are unknown.
Habitat: Adults of Mauroniscidae typically occur in desert regions where they can
be found on flowers feeding on nectar, pollen, and occasionally other arthropods.
Personal observations (AJM) suggest adults are active primarily in the morning and
in the afternoon, and can be collected beating or sweeping flowers of desert plants in
the families Asteraceae (Puecephyllum), Bignoniaceae (Chilopsis), Chenopodiaceae
(Atriplex), Lamiaceae (Hyptis), Loasaceae (Eucnide, Mentzelia), and Rosaceae
(Fallugia). The larvae and thus biology of species of Mauroniscidae is completely
unknown.
Notes: The family Mauroniscidae was described by Majer (1994) based primarily
on characters of the antennomeres, mouthparts and tarsal claws (see above). Recent
molecular data (Bocakova et al., 2011) confirm placement of the family as a basal
clade within the melyrid lineage. The family is known only from the Americas
ranging from Chile and Argentina North to the southwestern United States where
some species occur as far east as Kansas. Blackwelder (1957) did not report members
of this recently described family from Peru. However, some species can be found
listed under the genera Dasytes (Dasytes titschacki Pic) and Sydates (Sydates
discipennis Pic) in Melyridae. The five genera and 27 species of Mauroniscidae were
monographed by Majer (1995), who recorded five species in two genera from Peru.
The main characters that will differentiate species of Mauroniscidae from other
families of the melyrid lineage in Peru, are the small size, and simple tarsal claws
lacking membranous (ungual) appendages. Most recently, Howell (1997) described
Mecomycter majeri from the Sonoran desert region of southern California. All
members of the family occur in desert or prairie regions, and there are probably
many new species to be found in the desert regions of Peru. Images of a few North
American species can be found on BugGuide at http://bugguide.net/node/view/
748441. The following checklist of validated names was derived from Majer (1995).
1
Department of Entomology and Plant Pathology, The University of Tennessee, Knoxville, Tennessee
37901-1071.
2
3
Corresponding author, E-mail: cschaboo@ku.edu
Checklist:
Amecomycter crassus Majer, 1995 Mauroniscus titschacki (Pic, 1954)
Mauroniscus discipennis (Pic, 1910) Mauroniscus weyrauchi Majer, 1995
Mauroniscus subfasciatus Majer, 1995
Acknowledgements
Gimmel for help in assembling species names and we thank two anonymous
reviewers for advice that improved the text.
Literature Cited
Blackwelder, R. E. 1957. Checklist of the Coleopterous Insects of Mexico, Central America, The West
Indies, and South America. Part 6. U.S. National Museum, Bulletin 185. Smithsonian Institution,
Washington, D.C. vii + 9271492 pp.
Bocakova, M., R. Constantin, and L. Bocak. 2011. Molecular phylogenetics of the melyrid lineage
(Coleoptera: Cleroidea). Cladistics 27:113.
BugGuide. http://bugguide.net/node/view/748441, hosted by Iowa State University, Iowa (Last accessed
20 April 2015).
Howell, C. D. 1997. Three new species of North American Dasytinae, with two new generic keys. Journal
of the Kansas Entomological Society 70:1117.
Majer, K. 1994. A review of the classification of the Melyridae and related families (Coleoptera,
Cleroidea). Entomologica Basiliensia 17:319390.
Majer, K. 1995. Revision of the family Mauroniscidae (Insecta: Coleoptera: Cleroidea). Entomologische
Abhandlungen aus dem Staatlichen Museum fur Tierkunde (Dresden) 57:5789.
Pic, M. 1910. Contribution a letude des Dasytides de lAmerique meridionale. LEchange, Revue
Linneenne hors-texte 26(303):14.
Pic, M. 1954. Malacodermata (Col.): Cantharidae, Malachiidae, Dasytidae, Drilidae, Lampyridae.
Beitrage zur Fauna Perus (Jena) 4:170175.
Nitidulidae
Author(s): Andrew R. Cline, Gareth S. Powell, and Paolo R. Audisio

terms_of_use.
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88(2), 2015, pp. 217220
Beetles (Coleoptera) of Peru: A Survey of the Families. Nitidulidae

ANDREW R. CLINE,1,4 GARETH S. POWELL,2 AND PAOLO R. AUDISIO3

Recognition: This family may be recognized by the following combination of adult
characters: maxilla lacking galea; antennae typically 11-segmented, with terminal
three antennomeres forming a club; antennal insertions moderately to broadly
separated; subocular antennal grooves on ventral surface of head present; procoxae
transverse with exposed trochantins; prosternum transverse with intercoxal process
extending between procoxal cavities, procoxae always separated; mesoventral
procoxal rests present; typically five visible abdominal ventrites present; metendo-
sternite stalk present; metasternum with discrimen present; and tarsal formula 5-5-5.
Habitat: Several recent comprehensive reviews of Nitidulidae biology are available
(Jelnek et al., 2010; Cline, 2005; Audisio et al., 2000; Audisio, 1993). Overall,
nitidulid beetles exhibit a diverse set of life history strategies, including: fungivory,
saprophagy, phytophagy, pollination activities, facultative predation, necrophagy,
frugivory, and inquilinism with social insects. Nitidulids may be collected using
a variety of directed (sweeping and beating vegetation/inflorescences, sifting leaf
litter, searching under bark and various fungal substrates, as well as in nests of social
Hymenoptera) and passive techniques (flight intercept traps, pitfall traps, malaise
traps, and mercury vapor or black lights at night). Large numbers of nitidulids can
typically be found in palm inflorescences (e.g., Mytrops and related genera); baited
fruit or molasses traps (Carpophilus, Colopterus, Brachypeplus, Amphicrossus,
Lobiopa, and Cryptarcha), and sifting leaf litter (Stelidota).
Notes: Nitidulidae have long been confused with numerous other families of
Cucujoidea. In particular, members of Cybocephalidae, Kateretidae, Monotomidae,
Cryptophagidae, Biphyllidae, Smicripidae, and Erotylidae have been attributed to
Nitidulidae. Recent and upcoming work on the New World Kateretidae (Cline and
Audisio, 2010, and unpubl. data) will help improve the diagnostic capabilities for
Nitidulidae. A recent treatment of Nitidulidae higher level phylogenetics formally
elevated Cybocephalidae to family status and provided an extensive list of adult and
larval characters to differentiate this group from Nitidulidae (Cline et al., 2014).
Invasive species such as Urophorus humeralis (F.), Epuraea luteola Erichson, and
Carpophilus hemipterus (L.) likely occur in the country as they have been transported
around the world in various agricultural commodities, in conjunction with several
other alien species more or less recently introduced into Peru (Carrasco, 1986;
Leschen and Marris, 2005; EPPO, 2010). However, there are no existing records for
1
California Department of Food & Agriculture, Plant Pest Diagnostics Center, 3294 Meadowview Rd.,
Sacramento, California, USA, 95832
2
Department of Entomology, Smith Hall, Purdue University, 901 West State Street, West Lafayette,
Indiana, USA, 47907
3
Department of Biology and Biotechnologies Charles Darwin, Sapienza Rome University, Via A.
Borelli 50, 00161, Rome, ITALY
4
Corresponding Author. E-mail: andrew.cline@cdfa.ca.gov
these species in Peru. The addition of Epuraea luteola to the fauna would increase the
number of genera and subfamilies (i.e., Epuraeinae) in the region.
Widespread species known throughout South America, although not currently known
from Peru, but should be found there, include: Brachypeplus mutilatus Erichson,
Colopterus amputatus Erichson, Colopterus latus Murray, Hebascus traili Sharp, Lobiopa
insularis Castelnau, and Teichostethus testaceus (Erichson) (Blackwelder, 1945). Genera
not represented in the checklist below, but with undetermined or new species (Cline and
Audisio, unpubl. data) in institutional collections include: Amphicrossus Erichson,
Prometopia Erichson, and Somatoxus Sharp. Including these records would subsequently
increase the known generic count. The record of Amphicrossus specimens would increase
the number of genera and subfamilies (i.e., Amphicrossinae) in the region.
Other genera with numerous undetermined or new species to Peru in collection
holdings include: Brachypeplus Erichson, Camptodes Erichson, Carpophilus Erich-
son, Colopterus Erichson, Mystrops Erichson, and others (Carrasco, 1986; Cline and
Audisio, unpubl. data). The Brachypeplus records would increase the number of
known genera in the region.
Checklist:
Carpophilinae
Carpophilus brevipennis (Blanchard, 1842) Carpophilus lugubris Murray, 1864
Carpophilus dimidiatus (Fabricius, 1792) Carpophilus truncatus Murray, 1864
Carpophilus ligneus Murray, 1864
Cillaeinae
Colopterus infimus Erichson, 1843 Halepopeplus conoteloides Reitter, 1876

Colopterus macropterus (Fabricius, 1801) Halepopeplus dimidiatus Kirsch, 1873
Colopterus truncatus (Randall, 1838) Halepopeplus kirschi Reitter, 1876
Colopterus varius Erichson, 1847 Halepopeplus morio Reitter, 1876
Conotelus rufipes Murray, 1864 Halepopeplus suturalis Reitter, 1876
Conotelus stenoides Murray, 1864
Cryptarchinae
Cryptarcha fuscipennis Kirsch, 1873
Nitidulinae
Anthepurops depressa Kirejtshuk, 1996 Camptodes turpis Kirsch, 1873

Anthocorcina subcalva Kirejtshuk & Jelnek, 2000 Camptodes umbripennis Reitter, 1873
Camptodes aerumnosa Reitter, 1876 Cychramus distinctus Grouvelle, 1912
Camptodes dichroa Kirsch, 1873 Cyllodes rotundata Reitter, 1876
Camptodes difficilis Kirsch, 1873 Eusphaerius lubricus Leschen & Carlton, 1996
Camptodes distincta Reitter, 1873 Hebascus vestitus Kirsch, 1873***
Camptodes irritans Kirsch, 1873 Lasiodactylus brunneus Perty, 1830
Camptodes kirschi Reitter, 1876 Lobiopa lineola Kirsch, 1873
Camptodes laeta Kirsch, 1873 Lobiopa peruviana Guerin Meneville, 1829
Camptodes lateralis Reitter, 1876 Lobiopa setulosa Erichson, 1847
Camptodes molesta Kirsch, 1873 Meoncerus punctatolineatus Grouvelle, 1898
Camptodes nitidicollis Reitter, 1873 Mystrops astrocaryi Kirejtshuk & Couturier, 2010
Camptodes obsoleta Kirsch, 1873 Mystrops atrata Kirejtshuk & Couturier, 2010
Camptodes opaca Kirsch, 1873 Mystrops beserrai Kirejtshuk & Couturier, 2010
Mystrops dalmasi (Grouvelle, 1902) Platychorodes adentatus Kirejtshuk & Couturier,

Mystrops debilis Erichson, 1843 2010
Mystrops discoidea Murray, 1864 Platychorodes plumicornis Reitter, 1884
Mystrops kahni Kirejtshuk & Couturier, 2010 Pocadiolycra peruensis Kirejtshuk & Leschen, 1998
Mystrops komissari Kirejtshuk & Couturier, 2010 Pocadius peruensis Cline, 2008
Mystrops lobanovi Kirejtshuk & Couturier, 2010 Psilotus bicolor Cline, 2004
Mystrops pectoralis Kirejtshuk, Couturier & Jel- Stelidota geminata (Say, 1825)
nek, 2010 Stelidota metabola Kirsch, 1873
Mystrops squamae Kirejtshuk & Couturier, 2010 Stelidota thoracica Kirsch, 1873
Pallodes gracilipes Kirsch, 1873
***As stated by Jelnek (1975), the record of Hebascus vestitus from Peru is
somewhat dubious. The label data states the specimen is from Brasilia and no
other specimens exist to corroborate a Peruvian distribution. The record is here
treated as suspect.
Acknowledgements
We acknowledge NSF-EPSCoR # 66928 (PI: CS Chaboo) for supporting the
Evolutionary Biology-General Research Fund (PI: CS Chaboo). We thank the
numerous curators and colleagues that provided specimens and/or records from
material in their holdings, and the anonymous reviewers for helping to improve the
manuscript.
Literature Cited
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Indies, and South America. Part 3. National Museum Bulletin 185. Smithsonian Institution,
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Cline, A. R. 2005. Revision of Pocadius Erichson (Coleotpera: Nitidulidae). Ph.D. Dissertation.
Department of Entomology, Louisiana State University. 384 pp.
Cline, A. R. 2008. Revision of the Sap Beetle Genus Pocadius Erichson, 1843 (Coleoptera: Nitidulidae:
Nitidulinae). Zootaxa 1799:1120.
Cline, A. R., and P. Audisio. 2010. Revision of the New World Short-Winged Flower Beetles (Coleoptera:
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Coleopterists Bulletin 64:173186.
Cline, A. R., and C. E. Carlton. 2004. Review of Lasiodactylus Perty (Coleoptera: Nitidulidae) with
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Nitidulidae: assessment of subfamilial and tribal classifications, and formalization of the family
Cybocephalidae (Coleoptera: Cucujoidea). Systematic Entomology DOI: 10.1111/syen.12084.
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187 EPPO Reporting Service no. 10 2010.
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R. A. B., R. G. Beutel, and J. F. Lawrence (eds.). Handbook of Zoology. Vol. IV:pp. 390407.
Arthropoda: Insecta. Part 38. Coleoptera, Beetles. 786 pp.
Kirejtshuk, G. A., and G. Couturier. 2009. Species of Mystropini (Coleoptera, Nitidulidae) associated
with inflorescence of palm Ceroxylon quindiuense (Karst.) H. Wendl. (Arecaceae) from Peru.
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Nitidulidae: Nitiidulinae). Folia Heyrovskyana 8:171192.
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with three new genera and notes on mycophagy. Annales Zoologici 48:253273.
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Erichson 1843 (Coleoptera: Nitidulidae). Tropical Zoology 7:209216.
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including a new species of Eusphaerius. Journal of Natural History 30:18611873.
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notes on Australian species. http://www.biosecurity.govt.nz/files/regs/exports/plants/carpophilus.
pdf [Accessed 20-8-2014]
Tenebrionidae
Author(s): Aaron D. Smith, Alfredo E. Mendoza, Gustavo E. Flores, and Rolf L.
Aalbu

terms_of_use.
critical research.
88(2), 2015, pp. 221228
Beetles (Coleoptera) of Peru: A Survey of the Families. Tenebrionidae

AARON D. SMITH,1,5 ALFREDO E. GIRALDO MENDOZA,2 GUSTAVO E. FLORES,3 AND
ROLF L. AALBU4
Diversity in Peru: 8 subfamilies, 37 tribes, 105 genera, and 353 species/subspecies.

Recognition: Adult tenebrionids, commonly called darkling beetles, are highly
variable in terms of color (from black or brown to a wide variety of bright or metallic
colors), size (from approximately 180 mm), and morphology (from flat wingless
species to elongate cylindrical winged taxa). However, with few exceptions
tenebrionids can be identified based on their: 5-5-4 tarsal formula (rarely 4-4-4 or
3-3-3); antennal insertions concealed under frons; five visible abdominal sternites,
with the basal three sternites connate and the apical two hinged; and 10- to 11-
segmented antennae (rarely 9-segmented). Detailed descriptions, including for
immature stages, are provided in Aalbu et al. (2002) and Matthews et al. (2010).
Geographic distribution and habitats: Tenebrionids are found in a wide variety of
habitats in Peru, from coastal dunes to tropical forests and through the Andes
mountain range, with many tribes and genera restricted to specific biogeographic
regions or specialized microhabitats.
- The coastal deserts of western Peru (,1000 m): Caenocrypticini (Caenocrypti-
coides), Edrotini (Kocakia), Elenophorini (Psammetichus), Epitragini (Parepitra-
gus, Hemasodes, Omopheres), Evaniosomini (Aryenis, Chorasmius, Evaniosomus,
Melaphorus), Physogasterini (Philorea, Physogasterinus), Praociini (Parapraocis),
Thinobatini (Cordibates), Opatrini (Blapstinus), Scotobiini (Ammophorus), and
Tenebrionini (Hipalmus). These genera include epigaeic detritivores and epiphytic
groups (Edrotini, Epitragini) associated with various vegetation types on sandy
soils.
- The Andes, including the Western Andean range (1000 m3800 m), interandean
valleys (1500 m3300 m) and the high plateaus (.3800 m): Nycteliini (Pilobalia,
Psectrascelis), Praociini (Praocis, Praocidia, Pilobaloderes, Platyholmus), Edrotini
(Hylithus), Evaniosomini (Vaniosus), Trilobocarini (Eremoecus). These genera are
epigaeic detritivores associated with grasslands and scrubs.
- Amazonia, including the Eastern Andean range (800 m3500 m) and the
Amazonian plain (,800 m): Goniaderini, Lagriini, Nilio, Phrenapatini, Penetini,
1
Department of Biological Sciences, Northern Arizona University, PO Box 5640, Flagstaff, Arizona
86011-5640, USA.
2
Universidad Nacional Agraria La Molina, Museo de Entomologa Klaus Raven Buller, Av. La
Molina s/n, Lima 12, Lima, Peru.
3
CONICET, Laboratorio de Entomologa, Instituto Argentino de Investigaciones de las Zonas Aridas
(IADIZA, CCT-CONICET Mendoza), Casilla de correo 507, 5500 Mendoza, Argentina.
4
Department of Entomology, California Academy of Sciences, 55 Music Concourse Dr., Golden Gate
Park, San Francisco, California, USA.
5
Corresponding author, E-mail: Aaron.Smith@nau.edu
Accepted 05 June 2014; Revised 09 March 2015
Acropteronini, Amarygmini, Bolitophagini, Tenebrionini (Zophobas), Toxicini,

Triboliini (Hypogena), Ulomini, Alleculini, Diaperini (Adelina, Cosmonota,
Cynaeus, Hypophlaeus, Liodema, Neomida, Platydema), Cnodalonini and
Stenochiini. These taxa are mostly arboreal, associated with fungus and dead
logs, in tropical forests.
- Some genera are widely distributed in the coastal deserts and the Andes. The
genera Prohylithus (Edrotini), Epitragopsis (Epitragini) and Achanius (Evanioso-
mini) are found in coastal desert and interandean valleys, while Scotobius
(Scotobiini) is found from coastal deserts to high plateaus.
- Other genera are found in specialized habitats. The genus Phaleria inhabits sandy
beaches and is frequently associated with algae deposits. The genera Esemephe
(Cossyphodini), Grammicus and Discopleurus (Stenosini) are myrmecophilous. The
genera Lepidocnemeplatia (Cnemeplatiini), Alphitobius (Alphitobiini), Tribolium
(Triboliini), Gnathocerus and Ulomoides (Diaperini) are invasive cosmopolitan species
associated with stored grains, poultry farms, and other anthropic environments.
Notes: The checklist was based initially on records from Gebiens Katalog der
Tenebrioniden (Gebien 1937, 19381942, 19421948) and Blackwelder (19441957),
with a review of the original descriptive publications up to 1948 and all known post-
1948 taxonomic literature dealing with Peruvian tenebrionids. It also contains newly
recorded genera and species for the country identified in collections. Higher
classification follows Bouchard et al. (2011).
There are currently 8 subfamilies, 37 tribes, 105 genera, and 353 darkling beetle
species or subspecies known from Peru. Four tribes, 13 genera, and 33 species are
newly recorded for the country. However, the tenebrionid fauna of Peru, as with
many other arthropod groups, is still poorly known. The total number of Peruvian
species will almost certainly increase greatly as more sampling is done. Though few
darkling beetle genera (,17%) are restricted to Peru, the majority (,62%) of species
and subspecies described from the country are endemic based on currently available
records.
A few species listed in catalogs as occurring in Peru were excluded from the
checklist, based on a lack of known specimens from within the modern-day
boundaries of the country, and other factors. Megelenophorus americanus ovipennis
Fairmaire was described from specimens collected in Tucuman (Argentina) and
occurs in the Chaco biogeographic province, which extends from northern
Argentina to southeastern Bolivia and southwestern Paraguay. However, the
region does not extend into Peru and no specimens have been confirmed for the
country. Praocis hirticollis Laporte was also excluded as no specimens have been
found and the type material is missing (Kulzer, 1958; Flores and Pizarro-Araya,
2010). It is likely a synonym of another Praocis species (Flores, unpubl. data) as
well. Kulzer (1956) mistakenly listed Pseudothinobatis ohausi Kulzer as occurring in
Peru in the original description, but Pena (1973) stated that the four species of this
genus are exclusively from Ecuador. Solier (1838) described both Scotobius gayi
and S. kirby from Peru and Chile, but both species inhabit only central Chile based
on collections data.
Checklist:
Alleculinae: Alleculini
Allecula ainaensis Borchmann 1941* Lobopoda quinqueimpressa Pic, 1936*
Allecula debilis Erichson, 1847* Lobopoda umbrosa Erichson, 1847*
Cteisa hirta Solier 1835 Lystronychus sp. New Record
Lobopoda claveri Pic, 1924* New Record Prostenus clavipes longicollis Borchmann, 1937*
Lobopoda diehli Borchmann 1941* Prostenus peruanus Borchmann, 1937*
Lobopoda impressa Erichson, 1847* Xystropus ainaensis Borchmann 1941
Lobopoda inculta Erichson, 1847* Xystropus blandus Erichson, 1847*
Lobopoda peruviana Pic, 1928 Xystropus pilosus Kirsch, 1873
Diaperinae: Crypticini
Gondwanocrypticus sp. New Record
Diaperinae: Diaperini
Adelina sp. New Record Platydema erotyloides Chevrolat, 1878
Cosmonota picta picta Gebien, 1928* Platydema histrionica Gemminger, 1870*
Cosmonota picta erythropus Gebien, 1928* Platydema guatemalensis Champion, 1886
Cosmonota peruana Kulzer, 1961* New Record
Gnathocerus cornutus (Fabricius, 1798) Platydema janus Laporte & Brulle, 1831*
Liodema maculatum (Fabricius, 1801) Platydema quindecimmaculatus Chevrolat, 1878
Liodema serricorne Bates, 1873 New Record
Neomida castanea (Bates, 1873) Platydema tibiale Chevrolat, 1878 New Record
Neomida cioides (Champion, 1886) Platydema transversum Laporte & Brulle, 1831
Neomida distans (Champion, 1886) New Record
Neomida hoffmanseggi (Laporte & Brulle, 1831) Platydema xanthostigmatum Triplehorn, 1994*
Neomida pogonocera Triplehorn, 1994 Ulomoides dermestoides (Chevrolat, 1878)
Diaperinae: Hypophlaeini
Corticeus notialis Bremer & Triplehorn, 1999
Corticeus rufipes (Fabricius, 1801) New Record
Diaperinae: Phaleriini
Phaleria gayi Laporte, 1840
Phaleria pacifica Champion, 1886
Phaleria subparallela Chevrolat, 1878*
Lagriinae: Belopini
Adelonia sp. New Record
Lagriinae: Goniaderini
Anaedus angusticollis Kirsch, 1873* Goniadera (Goniadera) impressa Erichson, 1843
Anaedus rufocinctus Pic, 1917* Goniadera (Goniadera) interrupta Bates, 1868*
Gamaxus hauxwelli Bates, 1868* Goniadera (Goniadera) novemcostata Fairmaire, 1889
Goniadera (Aemyone) simplex Fairmaire, 1889 Goniadera (Goniadera) obscuriceps Pic, 1913
Goniadera (Goniadera) alternata Champion, 1886 Goniadera (Goniadera) repanda (Fabricius, 1801)
Goniadera (Goniadera) ampliata Gebien, 1912 Goniadera oculata Champion 1886 New Record
Goniadera (Goniadera) ardoini Ferrer & Delatour, 2007 Paratenetus latipennis Pic, 1920*
Goniadera (Goniadera) barclayi Ferrer & Delatour, Phobelius metallicus Kirsch, 1873*
2007 Phymatestes bomansi Ferrer, 2009*
Goniadera (Goniadera) claveri Pic, 1913 Phymatestes monstrosus Gebien, 1912
Goniadera (Goniadera) dissipata Kirsch, 1866 Phymatestes oedipus Gebien, 1928*
Goniadera (Goniadera) distinctipennis Pic, 1913 Phymatestes opacus Gebien, 1912*
Goniadera (Goniadera) girardi Ferrer & Delatour, Phymatestes rosemariae Ferrer & Moragues, 2000
2007
Lagriinae: Lagriini
Colparthrum bicinctum Champion, 1917* Statira coxalis Borchmann, 1921*
Colparthrum elegantulum Borchmann, 1916* Statira cupreoviridis Champion, 1917*
Colparthrum kolbei Borchmann, 1916* Statira distigma Champion, 1917*
Colparthrum nigrum Borchmann, 1916* Statira filicornis Champion, 1917*
Colparthrum spinicauda Champion, 1917* Statira laccata Borchmann, 1921*
Gebienia atricolor Pic, 1936* Statira maculipennis Borchmann, 1936*
Gebienia fissicollis Borchmann, 1921* Statira nigroapicalis Pic, 1912*
Gebienia impressicollis Borchmann, 1921* Statira nitidicollis Borchmann, 1921*
Gebienia rimulosa Borchmann, 1921* Statira pallidipes Pic, 1917*
Lagriola denticulata Kirsch, 1873* Statira peruana Champion, 1917*
Lagriola operosa Kirsch, 1873* Statira propinqua Borchmann, 1921
Statira aliena Borchmann, 1921* Statira relucens Borchmann, 1921*
Statira anthicoides anthicoides Kirsch, 1873* Statira semicatenata Borchmann, 1921
Statira anthicoides staudingeri Pic, 1912* Statira semicuprea Champion, 1917*
Statira caelata Erichson, 1847* Statira setosella Borchmann, 1921*
Statira callangana Borchmann, 1921* Statira vigintipunctata Champion, 1917*
Statira calophaenoides Champion, 1917* Statira viridinotata Pic, 1912*
Statira casaensis Borchmann 1936 Thoracostira sculpta Kirsch, 1873*
Statira chloroptera Champion, 1917* Uroplatopsis peruviana Pic, 1911*
Statira consobrina Borchmann, 1921* Uroplatopsis vitticollis Borchmann, 1921*
Nilioninae: Nilionini
Nilio borchmanni Mader, 1936 Nilio peruvianus Thomson, 1860
Nilio lafertei Thomson, 1860 New Record Nilio testaceus Thomson, 1861 New Record
Nilio latipenne Pic, 1911* Nilio violaceus Mader, 1936*
Nilio margaritaceus Brethes, 1920* Nilio vulpinus Mader, 1936
Nilio marginellus Erichson, 1847
Phrenapatinae: Phrenapatini
Phrenapates dux Gebien, 1911
Phrenapatinae: Penetini
Molion cervus Bates, 1873*
Peneta sp. New Record
Pimeliinae: Cossyphodini
Esemephe tumi Steiner, 1980*
Pimeliinae: Caenocrypticini
Caenocrypticoides penai Kaszab, 1969*
Caenocrypticoides peruanus Endrodi-Younga, 1996*
Pimeliinae: Cnemeplatiini
Lepidocnemeplatia sericea (Horn, 1870) New Record
Lepidocnemeplatia laticollis (Champion, 1884) New Record
Pimeliinae: Edrotini
Hylithus alpinus simplex Kaszab, 1964 Kocakia opaca (Kaszab, 1981)*
Hylithus curtus Kaszab, 1964 Prohylithus barbatus Kaszab, 1964*
Hylithus humilis Erichson, 1847 Prohylithus kulzeri Kaszab, 1964*
Hylithus peruensis peruensis Kaszab, 1964* Prohylithus peruanus Kaszab, 1981*
Hylithus peruensis complicans Kaszab, 1964*
Pimeliinae: Elenophorini
Psammetichus carinatus Solier, 1838 Psammetichus dissimilis Pena, 1994*
Psammetichus costatus Guerin-Meneville, 1831 Psammetichus gracilis Erichson, 1834
Psammetichus kulzeri Pena, 1973*
Pimeliinae: Epitragini
Epitragopsis antennatus Freude, 1981* Omopheres difficilis peruensis Freude, 1967*
Epitragopsis batesi bothrotiformis Freude, 1968 Parepitragus ater ater Kulzer, 1958
Epitragopsis batesi peruensis Freude, 1968* Parepitragus ater rossi Freude, 1967*
Epitragopsis diremptus Karsch, 1881 Parepitragus ater marcuzzi Freude, 1967
Epitragopsis olivaceus olivaceus Erichson, 1847 Parepitragus macrophtalmus Marcuzzi, 1961*
Epitragopsis olivaceus onensis Marcuzzi, 1961 Parepitragus pulverulentus pulverulentus Erichson,
Epitragopsis rotundicollis Marcuzzi, 1961 1847
New Record Parepitragus pulverulentus denticeps Gebien, 1928
Epitragus sallaei Champion, 1884 Parepitragus solieri Casey, 1907 New Record
Hemasodes peruensis Freude, 1967* Phytophylus helopioides Guerin-Meneville, 1831*
Pimeliinae: Evaniosomini
Achanius (Achanius) angusticollis Kulzer, 1956* Achanius (Ambigatus) rhinosomoides Kulzer, 1950*
Achanius (Achanius) anthicoides Erichson, 1847* Aryenis haagi Bates, 1872*
Achanius (Achanius) castanescens (Fairmaire, Chorasmius procerus (Erichson, 1847)*
1902)* Evaniosomus crassicornis Erichson, 1847*
Achanius (Achanius) minutus Kulzer, 1950* Evaniosomus declivis Erichson, 1847*
Achanius (Achanius) obscurus Kulzer, 1950* Evaniosomus orbygnianus Guerin-Meneville, 1834*
Achanius (Achanius) piceofuscus (Fairmaire, 1878)* Melaphorus elegans Kulzer, 1956
Achanius (Achanius) peruensis Kulzer, 1950* Melaphorus reichei Guerin-Meneville, 1834*
Vaniosus paradoxus Kulzer, 1956*
Pimeliinae: Nycteliini
Pilobalia alboscripta Fairmaire, 1903* Pilobalia ornata Kulzer, 1958*
Pilobalia baeri Fairmaire, 1902* Pilobalia philippi Kulzer, 1958*
Pilobalia becki Kulzer, 1954* Pilobalia ruficollis Kulzer, 1954*
Pilobalia blancasi Kulzer, 1958* Pilobalia soror Kulzer, 1958*
Pilobalia crassicornis Kulzer, 1954* Pilobalia subnuda Kulzer, 1954*
Pilobalia crassicosta Kulzer, 1954* Pilobalia tenella Kulzer, 1954*
Pilobalia decorata decorata Erichson, 1834 Pilobalia tristis Kulzer, 1954*
Pilobalia decorata titschaki Kulzer, 1954* Pilobalia unicolor Kulzer, 1954*
Pilobalia dorsoplicata Kulzer, 1954* Pilobalia voogdi Kulzer, 1954*
Pilobalia freyi Kulzer, 1954* Pilobalia weyrauchi Kulzer, 1954*
Pilobalia gracilipes Kulzer, 1954* Psectrascelis aequalis Fairmaire, 1876
Pilobalia haagi Fairmaire, 1878* Psectrascelis laevigata laevigata (Erichson, 1834)
Pilobalia lineata Kulzer, 1958* Psectrascelis laevigata rufipes Kulzer, 1954*
Pilobalia loffleri Kulzer, 1955* Psectrascelis marginipennis Kulzer, 1954*
Pilobalia michelbacheri Kulzer, 1954* Psectrascelis politicollis Fairmaire, 1876*
Pilobalia oberthuri Kulzer, 1954* Psectrascelis subplanata Pena, 1994*
Pimeliinae: Physogasterini
Philorea acunai Pena, 1980* Philorea peruana (Erichson, 1847)*
Philorea escomeli Lesne, 1911 Philorea picipes (Erichson, 1834)
Philorea koepckei Kulzer, 1956* Philorea pilosula Kulzer, 1956
Philorea leechi Kulzer, 1956* Philorea rossi Kulzer, 1956*
Philorea michelbacheri Kulzer, 1956 Philorea setipennis Lesne, 1911
Philorea mucronata Lesne, 1911* Philorea stangei Pena, 1980*
Philorea opaca Pena, 1980* Philorea weyrauchi Kulzer, 1956*
Philorea penai Kulzer, 1966* Physogasterinus lanuginosus Kaszab, 1981*
Pimeliinae: Praociini
Parapraocis fumaria (Kulzer, 1966)* Praocis (Filotarsus) forsteri Kulzer, 1958
Parapraocis rossi (Kulzer, 1958)* New Record
Parapraocis vagecostata (Fairmaire, 1902)* Praocis (Filotarsus) grossa Kulzer, 1958*
Pilobaloderes gebieni Kulzer, 1958* Praocis (Filotarsus) obesa Kulzer, 1958
Platyholmus brevis Kulzer, 1952* Praocis (Filotarsus) peltata Erichson, 1847
Platyholmus canus Kulzer, 1952* Praocis (Filotarsus) peruana Fairmaire, 1902*
Platyholmus nigrinus Kulzer, 1952* Praocis (Filotarsus) titschaki Kulzer, 1958*
Platyholmus peruanus Kulzer, 1952* Praocis (Filotarsus) weyrauchi Kulzer, 1958*
Platyholmus subglaber Kulzer, 1952 Praocis (Orthogonoderes) variolosa variolosa
Platyholmus variolosus Kulzer, 1958* Erichson, 1834*
Praocidia acuticollis Kulzer, 1958* Praocis (Orthogonoderes) variolosa laxepunctata
Praocidia nervosa (Fairmaire, 1902)* Kulzer, 1958*
Praocis (Filotarsus) brevicornis Kulzer, 1958* Praocis (Praocida) montana Kulzer, 1958
Pimeliinae: Stenosini
Discopleurus peruanus Kulzer, 1956*
Grammicus antennatus Kulzer, 1956*
Grammicus chilensis Waterhouse, 1845 New Record
Pimeliinae: Thinobatini
Cordibates fuscus Kulzer, 1958* Cordibates nitidus Pena, 1974*
Cordibates michelbacheri Freude, 1959* Cordibates opacus Pena, 1974*
Cordibates minor Pena, 1974* Cordibates pilosus Pena, 1974*
Pimeliinae: Trilobocarini
Eremoecus cordicollis Kirsch, 1873
Stenochiinae: Cnodalonini
Blapida bicolor Gebien, 1919 Epicalla matertera Gebien, 1928*
Blapida peruensis Gebien, 1919* Isicerdes spp. New Record
Camaria guttipennis Gebien, 1919* Mylaris gigas (Linnaeus, 1787)
Camaria incerta Pic, 1930* Mylaris procerus (Champion, 1885) New Record
Camaria singularis Pic, 1919* Nuptis sp. New Record
Choastes sp. New Record Othryoneus barclayi Ferrer 2009
Cyrtosoma amazonicum Marcuzzi, 1976* Paroeatus nitidus Gebien, 1928
Cyrtosoma angulicollis Marcuzzi, 1992* Sycophantes peruana Gebien, 1928*
Cyrtosoma coracina Gebien, 1928* Taphrosoma dohrni Kirsch, 1866
Cyrtosoma peruvianum Marcuzzi, 1976* Tenebriocamaria atra Pic, 1919*
Cyrtosoma poggii Marcuzzi, 2002*
Stenochiinae: Stenochiini
Cuphotes cincta cincta Olivier, 1795 Strongylium chalcodes Erichson, 1847*
Cuphotes cincta bimaculata Pic, 1918* Strongylium cyanicorne Erichson, 1847*
Cuphotes erichsoni Thomson, 1859* Strongylium denticolle Sharp 1891
Falsostrongylium subcostulatum Pic, 1915* Strongylium haemorrhoidale Fabricius, 1792
Poecilesthus curvipes Maklin, 1878* Strongylium pavonii Erichson, 1847*
Poecilesthus eximius Maklin, 1878* Strongylium peruvianum Pic, 1918*
Poecilesthus quadrisignatus Maklin, 1878* Strongylium procerum Erichson, 1847*
Poecilesthus testaceitarsis Pic, 1930* Strongylium rubrithorax Brethes, 1920*
Poecilesthus tumidus Maklin, 1878* Strongylium ruficorne Kirsch, 1873*
Strongylacanthus peruvianus Brethes, 1925* Strongylium rutilans Erichson, 1847*
Strongylium axillare Kirsch, 1873* Strongylium subimpressipenne Pic, 1935*
Strongylium azureum (Germar 1824) Strongylium trinudulatum Pic, 1918*
Stenochiinae: Talanini
Talanus sp. New Record
Tenebrioninae: Acropteronini
Acropteron pallicrus Fairmaire, 1902*
Tenebrioninae: Alphitobiini
Alphitobius diaperinus (Panzer, 1797)
Tenebrioninae: Amarygmini
Cymatothes undata Fabricius, 1792
Tenebrioninae: Bolitophagini
Rhipidandrus peruvianus Laporte, 1840*
Tenebrioninae: Opatrini
Ammodonus granosus Fall, 1912 New Record Blapstinus holosericius (Laporte, 1840)
Blapstinus cisteloides Erichson, 1847* Trichoton posthumum Gebien, 1928 New Record
Blapstinus helopioides Erichson, 1847* Trichoton sordidum LeConte, 1851 New Record
Tenebrioninae: Scotobiini
Ammophorus blairi Van Dyke, 1953* Scotobius exaratus Erichson, 1847*
Ammophorus costatus Guerin-Meneville, 1834 Scotobius fallax Kulzer, 1966*
Ammophorus peruvianus Guerin-Meneville, 1831 Scotobius leechi Kulzer, 1955*
Ammophorus rubripes Solier, 1838 Scotobius michelbacheri Kulzer, 1955*
Ammophorus spinolae Solier, 1838* Scotobius planatus Erichson, 1834
Pumiliofossorum moche Silvestro & Flores 2015* Scotobius planicosta Guerin-Meneville, 1834
Pumiliofossorum sechurae Giraldo & Flores 2015* Scotobius rossi Kulzer, 1955*
Scotobius atacamensis Germain, 1855 Scotobius vulgaris Guerin-Meneville, 1834
Scotobius costatus Guerin-Meneville, 1834 Scotobius weyrauchi Kulzer, 1955*
Scotobius crassus Kulzer, 1955 Scotobius zischkai Kulzer, 1955
Tenebrioninae: Tenebrionini
Hipalmus costatus (Guerin-Meneville, 1831)* Zophobas (Macrozophobas) tibialis (Kraatz, 1880)
Rhinandrus peruanus Kraatz, 1880* Zophobas (Zophobas) atratus (Fabricius, 1775)
Zophobas (Macrozophobas) bifasciatus Erichson, Zophobas (Zophobas) tridentatus Kraatz, 1880
1847 Zophobas (Zophobas) opacus (Sahlberg, 1823)
Zophobas (Macrozophobas) haagi Kraatz, 1880 Zophobas (Zophobas) lugubris Boheman, 1858
Zophobas (Macrozophobas) maculicollis Kirsch, 1866
Tenebrioninae: Toxicini
Ozolais divisa Pascoe, 1883 New Record Wattius vestitus Champion, 1886
Ozolais scruposa Pascoe, 1866 New Record Wattius n. sp.
Tenebrioninae: Triboliini
Hypogena biimpresa (Latreille, 1813) New Record Hypogena cat Steiner, 2005*
Hypogena triceratops Steiner, 2005* Tribolium castaneum (Herbst, 1797)
Tenebrioninae: Ulomini
Alegoria caviceps Gebien, 1928* Apteruleda acutangula Gebien, 1928*
Alegoria dilatata Laporte, 1840 Apteruleda uncipes Gebien, 1928*
Alegoria parallelogramma Gebien, 1928* Uloma retusa (Fabricius, 1801)
Antimachus sp. New Record Uloma subcylindrica Gebien, 1928*
Acknowledgements
Beetles of Peru project, the NSF ARTS program (DEB-1258154) for support of
Aaron Smiths research, and CONCIET (Argentina) for support of Gustavo E.
Floress research. The University of Kansas Department of Ecology and
Evolutionary Biology-General Research Fund (PI: CS Chaboo) is funding this
publication. We also thank Matthew Gimmel and two anonymous reviewers for
their help.
Literature Cited
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(eds.). American Beetles. Vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea, pp. 463509.
CRC Press, Boca Raton, xiv + 861 pp.
Blackwelder, R. E. 19441957. Checklist of the Coleopterous insects of Mexico, Central America, the
West Indies, and South America. USNM Bulletin, 185:ixii, 1492.
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Brethes J. 1920. Insectes du Perou. Anales de la Sociedad Cientfica Argentina, 89:2754.
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de la Societe entomologique de France. 148151.
Flores, G. E., and J. Pizarro-Araya. 2010. Two new synonymies in the genus Praocis (Coleoptera:
Tenebrionidae). Zootaxa 2386:6568.
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Gebien, H. 19381942. Katalog der Tenebrioniden. Teil II. Mitteilungen der Munchener Entomolo-
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Guerin-Meneville, F.-E. 1831. Crustaces, arachnides et insectes. Plate 4. In L. I. Duperrey (ed.), Voyage
autour du monde, execute par ordre du Roi, sur la corvette de la majeste, La Coquille, pendant les
annees 1822, 1823, 1824 et 1825, sous le ministere et conformement aux instructions de S. E. M. le
marquis de Clermont-Tonnerre, ministre de la marine. Zoologie. Atlas. A. Bertrand, Paris.
Kirsch T. 1873. Beitrage zur Kenntnis der Peruanischen Kaferfauna. Berliner entomologische Zeitschrift,
17: 339418. Berlin.
Kulzer, H. 1956. Neue Tenebrioniden aus Sudamerika. (15. Beitrag zur Kenntnis der Tenebrioniden).
Entomologische Arbeiten aus dem Museum Georg Frey 7:895964.
Kulzer, H. 1958. Monographie der sudamerikanischen Tribus Praocini (Col.). (16 Beitrag zur Kenntnis
der Tenebrioniden). Entomologische Arbeiten aus dem Museum Georg Frey 9:1105.
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Berlin/New York, Chapter 11.14, pp. 574659
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Coccinellidae
Author(s): Guillermo Gonzlez

terms_of_use.
critical research.
88(2), 2015, pp. 229236
Beetles (Coleoptera) of Peru: A Survey of the Families. Coccinellidae

GUILLERMO GONZALEZ
Nocedal 6455, La Reina, Santiago, Chile
E-mail: willogonzalez@yahoo.com
Diversity in Peru: 64 genera, 329 species, 199 endemics (60%).

Recognition: Coccinellids are popularly known as ladybugs or ladybird beetles.
The adults are minute to medium sized (112 mm). The body is oblong or semicircular,
flat underneath and normally convex dorsally, some depressed. The tarsal formula is
pseudotetramerous or trimerous. The antennae have 811 antennomeres and a 15
club. Pilosity may be present or absent. Body colors are frequently red or orange with
black or white; in some groups the color is yellow, brown or piceous, rarely metallic
green or blue. A post-coxal line is present on the first abdominal ventrite and the penis
is modified into a sipho. Detailed descriptions of the family can be founded in
Vandenberg (2002) and Slipinski and Tomaszewska (2010).
Habits and Habitats: Coccinellids have a wide range of food preference. Some
species are plant feeders (Epilachninae) and a few feed exclusively on fungi
(Halyziini). The majority, however, are predators of adults and immature stages of
hemipteran from the suborder Sternorrhyncha (Aphididioidea, Aleyrodoidea,
Psyllidae, Margarodidae, etc). Exceptions from this general predatory pattern might
occur, the most relevant being the specialization on tetranychid mites by members of
the tribe Stethorini. Given their predatory habits, several coccinellids are used in the
biological control of agricultural pest. Some plant feeders, on the other hand, might
reach pest status. Coccinellids develop quickly, in about a month in spring or
summer; the adults live about six months. They are commonly diurnal species.
Notes: The checklist below covers until 2014. The classification used follows Bouchard
et al. (2011) with minor changes in tribe structure. The last list available considered only
about 100 species (Blackwelder, 1945). The 2/3 of the currently recognized species were
described in the last 50 years, especially through the works of Robert D. Gordon from
1969 to today. The bibliography contains all the taxonomic works not included in
Blackwelder (1945) or in the bibliography of Gonzalez (2010). An updated bibliography
including taxonomy, phylogeny, biology and agriculture-related jobs, as well as a key to
the genera of Coccinellidae in South America, and individual pages about the majority of
the species cited from Peru can be found at the website: www.coccinellidae.cl.
Some species have been erroneously assigned to Peru in previous publications:
Delphastus pusillus was cited by Miro and Castillo (2010) (misidentification), but this
species has been removed from the Peruvian fauna by Gordon (1994).
Pentilia egena Mulsant cited by Gonzalez, 2010 (error in countries list).
Psyllobora lenta Mulsant cited by Miro and Castillo (2010) (misidentification).
Scymnobius bilucernarius Mulsant, 1850, cited for Gordon and Gonzalez (2002)
(error in countries list in key, confirmed with R.D. Gordon, personal communication).
Accepted 21 April 2014; Revised 25 November 2014

Serratitibia sarah Gordon and Canepari (2013) in the original description, where the
type locality (Tabatinga) was mistakenly assigned to Peru instead of Brazil
(confirmed with Robert D. Gordon, personal communication).
Zagloba beaumonti Casey cited by Gonzalez (2013b) (error in countries list).
In the present work, I provide eleven new records for coccinellids in Peru:
Delphastus huahuai Gordon, 1994: PERU, Madre de Dios, Tahuamanu, Iberia,

292 m. 10I2010. leg. J. Miro/Ctricos (Museo de Entomologa de la
Universidad Nacional de Tumbes, Tumbes, Peru).
Pseudoazya boliviana Gordon, 1980: PERU, Madre de Dios, Tambopata, Las Piedras,
Alegra, 5VIII2012, leg. J. Miro/N: 8860120 E: 486620, Ctricos: Naranjo
(Museo de Entomologa de la Universidad Nacional de Tumbes, Tumbes, Peru).
Stethorus grossepunctatus Gordon and Chapin, 1983: PERU, Depto. [Departa-
mento] Piura, Sol Sol, Chulucanas, 27VII2009, leg. A. Garces (Coleccion
Manuel Dieguez, Santiago, Chile).
Scymnobius ecuadoricus Gordon and Gonzalez, 2002: PERU, Provincia de Tumbes,
Corrales, Los Cedros, 27VIII2011, leg. Ruth Ros (Museo de Entomologa de
la Universidad Nacional de Tumbes, Tumbes, Peru).
Cryptognatha gemellata Mulsant, 1850: PERU, Amazonas, Pedro Ruiz, 1500 m,
8II2009, leg. R. Westerduijn. Vegetation near stream (Coleccion Museo de
Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima,
Peru).
Hyperaspis matronata Mulsant, 1853: Peru, Madre de Dios, Tahuamanu, Siringayoc,
3-IX-2011, leg. Jimmy Miro/Ctricos, Limon sutil (Museo de Entomologa de la
Universidad Nacional de Tumbes, Tumbes, Peru). Cited previously from Peru by
Gonzalez (2013b) without label data.
Cycloneda ecuadorica (Timberlake, 1943): PERU, [Ancash] Caraz, 28IX [19]96, [leg.]
Karina Vilca (Universidad Nacional Santiago Antunez de Mayolo, Ancash, Peru).
Olla roatanensis Vandenberg, 1992: [PERU, Tumbes], Puerto Pizarro, 13IX
[19]97, [leg.] P. Castillo (Museo de Entomologa de la Universidad Nacional de
Tumbes, Tumbes, Peru).
Epilachna nigrovittata Crotch, 1874: PERU, Amazonas, Pedro Ruiz, 1500 m. 8II
2009, leg. R. Westerduijn. Vegetation near stream (Museo de Entomologa Klaus
Raven Buller, Universidad Nacional Agraria La Molina, Lima, Peru).
Epilachna holmgreni (Weise, 1926): PERU, Cusco Dept. [Departamento],
Wayquecha Field Station, Canopy Trail near road, 13.1856uS 71.58877uW,
2990m, 13-V-2011, [leg.] D.J. Bennet, sweeping, PER-11-DJB-011 (KUNHM).
Eremochilus peregrinus Weise, 1912: PERU: Cusco Dept. Villa Carmen field station
12.89497uS 71.40364uW 520m 22.V.2011 D.J. Bennett misc. hand collecting
PER-11-DJB-025(KUNHM).
Checklist:
Microweiseinae Leng, 1920
Microweiseini Leng, 1920
Coccidophilus citricola Brethes, 1905

Coccidophilus lozadai Gonzalez, 2013*
Coccidophilus occidentalis Gonzalez, 2013
Serangiini Pope, 1962
Delphastus anthracinus Gordon, 1970 Delphastus huahuai Gordon, 1994 new record
Delphastus berryi Gordon, 1994* Delphastus quinculus Gordon, 1994
Coccinellinae Latreille, 1807

Cephaloscymnini Gordon, 1985
Prodilis maculata Weise, 1902*
Coccidulini Mulsant, 1846

Eupalea venusta Weise, 1899
Mimoscymnus rossi Gordon, 2002*
Rhyzobius lophanthae (Blaisdell, 1892)
Poriini Mulsant, 1850
Poria rubens Weise, 1899*

Chnoodini Mulsant, 1850
Chnoodes abendrothi Kirsch, 1876* Chnoodes terminalis Mulsant, 1850

Chnoodes decemmaculata Mader, 1957* Dioria zonata Kirsch, 1876*
Chnoodes dorsalis Kirsch, 1876* Exoplectra ruberrima Erichson, 1847*
Chnoodes maculamantis Gonzalez, 2013* Exoplectra spatularis Gonzalez, 2013*
Chnoodes separata Mader, 1957 Gordonita anomala Gonzalez, 2013*
Chnoodes splendidus Gonzalez, 2013* Incurvus lesnei (Sicard, 1912)
Oryssomini Gordon, 1974

Gordonoryssomus mirnae Almeida and Santos, 2014*
Azyni Mulsant, 1850
Azya satipoi Gordon, 1980* Azya orbigera ecuadorica Gordon, 1980

Azya scutata Mulsant, 1850 Azya weyrauchi Gordon, 1980*
Azya orbigera Mulsant, 1850 Pseudoazya boliviana Gordon, 1980 new record
Noviini Mulsant, 1846

Anovia peruviana Gordon, 1972*
Anovia punica Gordon, 1972
Rodolia cardinalis (Mulsant, 1850)
Ortaliini Mulsant, 1850
Zenoria discoidalis (Kirsch, 1876)* Zenoria peruviana Gordon, 1972*
Zenoria dozieri Gordon, 1972* Zenoria purpurea Gordon, 1972*
Zenoria lativerpa Gonzalez and Honour, 2012* Zenoria sylvatica Gonzalez and Honour, 2012*
Zenoria papryzyckii Gordon, 1971* Zenoria variabilis Gordon, 1972
Stethorini Dobzhansky, 1924
Parastethorus histrio (Chazeau and Fursch, 1974) Stethorus peruvianus Gonzalez et al., 2008*
Stethorus grossepunctatus Gordon and Chapin, Stethorus tridens Gordon, 1982
1983 new record
Scymnini Mulsant, 1846
Nephaspis acuta Gonzalez, 2009* Scymnobius ecuadoricus Gordon and Gonzalez,
Nephaspis aquarius Gordon, 1996 2002 new record
Nephaspis isabelae Gonzalez, 2009* Scymnobius galapagoensis (Waterhouse, 1845)
Scymnobius triangularis Gordon and Gonzalez, Scymnus mesomelas Kirsch, 1876*(1)

2002 Scymnus notatus Kirsch, 1876*(1)
Scymnus cerinotum Gordon, 2000* Scymnus paprzyckii Gordon, 2000*
Scymnus curviger Kirsch, 1876*(1) Scymnus peruanus Weise, 1929*
Scymnus demerarensis Gordon, 2000 Scymnus reyi Kirsch, 1876*(1)
Scymnus discimacula Kirsch, 1876*(1) Scymnus rubicundus Erichson, 1847
Scymnus hamatus Gordon, 2000 Scymnus simillinus Gordon, 2000
Scymnus quadrimaculatus Kirsch, 1876*(1) Scymnus spanglerorum Gordon, 2000
Scymnus labiatus Kirsch, 1876*(1) Scymnus vulneratus Kirsch, 1876*
Scymnus loewii Mulsant, 1850
Diomini Gordon, 1999

Diomus angela Gordon, 1999 Diomus jerome Gordon, 1999*
Diomus anthony Gordon, 1999 Diomus juliana Gordon, 1999*
Diomus bruno Gordon, 1999* Diomus leonard Gordon, 1999
Diomus castilloi Gonzalez and Honour, 2011* Diomus macarius Gonzalez and Honour, 2011*
Diomus chrysanthus Gordon, 1999* Diomus martha Gordon, 1999*
Diomus chrysogonus Gordon, 1999 Diomus melchiades Gordon, 1999*
Diomus cyril Gordon, 1999* Diomus protase Gordon, 1999*
Diomus damasus Gordon, 1999* Diomus secunda Gordon, 1999
Diomus daria Gordon, 1999* Diomus seminulus Mulsant, 1850
Diomus eleutherius Gordon, 1999* Diomus slipinskii Gonzalez and Honour, 2011*
Diomus eudes Gordon, 1999* Diomus thomas Gordon, 1999*
Diomus eusebius Gordon, 1999* Diomus thoracicus Gordon, 1999
Diomus frances Gordon, 1999* Diomus tucumanus Weise, 1906
Diomus hippolytus Gordon, 1999* Diomus westwerduijni Gonzalez and Honour, 2011*
Diomus hyacinthus Gonzalez and Honour, 2011* Diomus william Gordon, 1999
Diomus innocentius Gonzalez and Honour, 2011* Diomus xenon Gonzalez and Honour, 2011*
Scymnillini Casey, 1899
Zagloba mimica Gonzalez and Aguilera, 2009*

Zilus miroi Gonzalez and Aguilera, 2009*
Cryptognathini Mulsant, 1850
Cryptognatha auriculata Mulsant, 1850 Pentilia cincta Kirsch, 1876*

Cryptognatha gemellata Mulsant, 1850 new record Pentilia dispar Kirsch, 1876*
Curticornis bicolor Gordon, 1971* Pentilia minuta Kirsch, 1876*
Curticornis satipensis Gordon, 1971* Pentilia specularis Kirsch, 1876*
Hyperaspidini Mulsant, 1846

Clypeaspis trilineata Mulsant, 1850 Hyperaspis prolata Gordon and Canepari, 2008
Diazonema fallax Weise, 1926* Hyperaspis proserpinae Mulsant, 1850*(1)
Diazonema pubescens Weise, 1926 Hyperaspis satipoensis Gordon and Canepari,
Hyperaspidius trimaculatus Linnaeus, 1767 (1) 2008*
Hyperaspis arida Gordon and Canepari, 2008* Hyperaspis vetusta Weise, 1902*(1)
Hyperaspis cingulata Korschefsky, 1931* Menoscelis saginata Mulsant, 1850
Hyperaspis esmeraldas Gordon and Gonzalez, 2011 Peruaspis hypocrita Gordon and Canepari, 2008*
Hyperaspis festiva Mulsant, 1850 Peruaspis paprzyckii Gordon and Canepari, 2008*
Hyperaspis matronata Mulsant, 1853 Tenuisvalvae bisquinquepustulata Fabricius, 1801
Hyperaspis onerata Mulsant, 1850 Tenuisvalvae bromelicola Sicard, 1925
Brachiacanthini Mulsant, 1850
Brachiacantha april Gordon and Canepari, 2014 Brachiacantha blandula (Weise), 1902
Brachiacantha bistripustulata (Fabricius, 1801) Brachiacantha bruchi Weise, 1906
Brachiacantha buckleyi Crotch, 1874 Serratitibia barbara Gordon and Canepari, 2013*
Brachiacantha darlene Gordon and Canepari, 2014 Serratitibia beverly Gordon and Canepari, 2013*
Brachiacantha groendali (Mulsant), 1850 Serratitibia bonnie Gordon and Canepari, 2013*
Brachiacantha hazel Gordon and Canepari, 2014* Serratitibia dennise Gordon and Canepari, 2013*
Brachiacantha jamie Gordon and Canepari, 2014* Serratitibia donna Gordon and Canepari, 2013
Brachiacantha loricata (Mulsant), 1850 Serratitibia doris Gordon and Canepari, 2013*
Brachiacantha octopustulata (F.) 1801 Serratitibia fraudulenta (Kirsch, 1876)
Cyrea blandula (Weise, 1902)* Serratitibia heather Gordon and Canepari, 2013*
Cyrea jocosa (Mulsant, 1850)* Serratitibia judy Gordon and Canepari, 2013*
Cyrea propria (Kirsch, 1876)* Serratitibia julie Gordon and Canepari, 2013*
Cyrea renifera (Kirsch, 1876)* Serratitibia katherine Gordon and Canepari, 2013*
Cyrea staudingeri (Weise, 1901)* Serratitibia kimberly Gordon and Canepari, 2013*
Cyrea thriacantha (Mulsant, 1850) Serratitibia laura Gordon and Canepari, 2013*
Dilatitibialis carolinae (Crotch, 1874) Serratitibia loreto Gordon and Canepari, 2013
Dilatitibialis connie Canepari and Gordon, 2013 Serratitibia mary Gordon and Canepari, 2013*
Dilatitibialis edith Canepari and Gordon, 2013* Serratitibia melissa Gordon and Canepari, 2013*
Dilatitibialis ellen Canepari and Gordon, 2013* Serratitibia paprzycki Gordon and Canepari, 2013*
Dilatitibialis florence Canepari and Gordon, 2013* Serratitibia quincemil Gordon and Canepari, 2013*
Dilatitibialis luteola (Mulsant, 1850) Serratitibia regularis (Erichson, 1847)*
Dilatitibialis mulsanti (Kirsch, 1876) Serratitibia satipoensis Gordon and Canepari,
Dilatitibialis semicincta (Weise, 1899) 2013*
Dilatitibialis shannon Canepari and Gordon, 2013 Serratitibia susan Gordon and Canepari, 2013*
Dilatitibialis thelma Canepari and Gordon, 2013* Serratitibia teresa Gordon and Canepari, 2013*
Serratitibia abendrothi (Kirsch, 1876)* Serratitibia uncinata (Mulsant, 1853)
Chilocorini Mulsant, 1846
Curinus coeruleus Mulsant, 1850 Zagreus decempunctatus (Weise, 1893)

Exochomus orbiculus Weise, 1893*(1) Zagreus hexasticta (Crotch, 1874)*
Harpasus quadrifolium Gonzalez et al., 2008
Coccinellini Latreille, 1807

Anatis lebasi (Mulsant, 1850) Eriopis heliophila Mulsant, 1850
Cirocolla conspicillata (Mulsant, 1850) Eriopis huancavelicae Bustamante et al., 2009*
Coleomegilla maculata limensis (Philippi and Eriopis lawalawani Bustamante et al., 2007*
Philippi, 1864) Eriopis minima Hofmann, 1970
Coleomegilla occulta Gonzalez, 2014b Eriopis nobilis Mader, 1958*
Cycloneda ancoralis (Germar, 1824) Eriopis peruviana Hofmann, 1970*
Cycloneda andresi Oroz et al., 2009* Eriopis punicola Hofmann, 1970
Cycloneda arcula (Erichson, 1847) Eriopis sebastiani Bustamante, 2005*
Cycloneda ecuadorica (Timberlake, 1943) new record Harmonia axyridis (Pallas, 1772)
Cycloneda ebenina (Mulsant, 1866)* Hippodamia convergens Guerin-Meneville,1836
Cycloneda fryii Crotch, 1874 Neda patula Erichson, 1847
Cycloneda marcapatae Oroz et al. 2009* Neda boliviana Weise, 1898
Cycloneda reclusa Weise, 1902* Neda ochracea Erichson, 1847
Cycloneda sanguinea (Linnaeus, 1763) Neda cardinalis Erichson, 1847*
Eriopis alticola Hofmann, 1970* Neda ostrina Erichson, 1847
Eriopis canrash Bustamante et al., 2007* Neocalvia blanchardi Mulsant, 1850.
Eriopis concordia Gonzalez, 2014 Olla roatanensis Vandenberg, 1992* new record
Eriopis connexa (Germar, 1824) Paraneda pallidula guticollis (Mulsant, 1850)
Halyziini Mulsant, 1846
Oxytella longula Weise, 1902* Psyllobora huancayensis Almeida, 1991*

Psyllobora abancayana Almeida, 1991* Psyllobora lutescens Crotch, 1874
Psyllobora confluens Fabricius, 1801 Psyllobora peruana Weise, 1902*
Discotomini Mulsant, 1850

Pristonema coccinea Erichson, 1847*
Epilachnini Mulsant, 1846
Adira nucula (Weise, 1902)* Epilachna pastica (Weise, 1902)

Epilachna aureola Gordon, 1975 Epilachna patricia Mulsant, 1850
Epilachna aureopilosa Gordon, 1975* Epilachna peltata Erichson, 1847
Epilachna basalis (Weise, 1898)* Epilachna pemptea Gordon, 1975*
Epilachna bisbivittata Gordon, 1975* Epilachna persimilis Crotch, 1874
Epilachna bonplandi Mulsant, 1850 Epilachna peruviana Crotch, 1874*
Epilachna cacica (Guerin-Meneville, 1836) Epilachna propinqua (Weise, 1898)*
Epilachna callangae Gordon, 1975* Epilachna pseudolepida Gordon, 1986*
Epilachna ciliata Gordon, 1986* Epilachna pseudostriata Gordon, 1975*
Epilachna confixa Gordon, 1975* Epilachna quirozensis Gordon, 1975*
Epilachna consimilis Gordon, 1975* Epilachna satipensis Gordon, 1975*
Epilachna convergens Crotch, 1874 Epilachna schunkei Gordon, 1975*
Epilachna cuscoi Gordon, 1975* Epilachna sellata Weise, 1895
Epilachna cushmani Gordon, 1975* Epilachna sexmaculata Kirsch, 1876*
Epilachna discoidea Erichson, 1847* Epilachna simulans Gordon, 1975*
Epilachna discolor Erichson, 1847 Epilachna staudingeri (Weise, 1902)
Epilachna dives Erichson, 1847* Epilachna strictanotata Gordon, 1975*
Epilachna divisa (Weise, 1899)* Epilachna striola (Weise, 1902)*
Epilachna divisoides Gordon, 1975* Epilachna sztolcmani Jadwiszczak and
Wegrzynowicz, 2003*
Epilachna dorsigera Erichson, 1847*
Epilachna transverselineata (Mader, 1958)
Epilachna emerita Gordon, 1975*
Epilachna velata Erichson, 1847
Epilachna esemephata Gordon, 1986*
Epilachna viridilineata Crotch, 1874
Epilachna eusema (Weise, 1904)
Epilachna viridinitens Crotch, 1874*
Epilachna fausta Erichson, 1847
Epilachna vittigera Crotch, 1874
Epilachna fenestrata Erichson, 1847
Epilachna v-pallidum Blanchard, 1846
Epilachna fenestroides Gordon, 1975*
Epilachna woytkowskii Gordon, 1975
Epilachna flavofasciata (Laporte, 1840)
Toxotoma andicola Weise, 1899*
Epilachna furtiva Gordon, 1975*
Toxotoma chapini Gordon, 1975*
Epilachna fuscopilosa (Weise, 1902)* Toxotoma cuzcoensis Gordon, 1975*
Epilachna geometrica (Weise, 1899)* Toxotoma disparans Gordon, 1975*
Epilachna honesta (Weise, 1899)* Toxotoma huanucoi Gordon, 1975*
Epilachna ignobilis (Weise, 1902)* Toxotoma imitator Gordon, 1975*
Epilachna incaorum Gordon, 1975* Toxotoma leechi Gordon, 1975*
Epilachna languida (Weise, 1899)* Toxotoma longicrura Gordon, 1975*
Epilachna lepida Erichson, 1847* Toxotoma mimetica Gordon, 1975*
Epilachna mandibularis Gordon, 1975* Toxotoma nunenmacheri Gordon, 1975*
Epilachna monovittata Gordon, 1975 Toxotoma opulenta (Weise, 1902)*
Epilachna mutabilis Crotch, 1874 Toxotoma orbicula Gordon, 1975*
Epilachna nigrovittata Crotch, 1874 new record Toxotoma pilifera (Weise, 1902)
Epilachna obliqua Gordon, 1975* Toxotoma pulchra (Weise, 1902)*
Epilachna olmosi Gordon, 1975* Toxotoma rosae, Gordon, 1975*
Epilachna ostensa (Weise, 1902) Toxotoma rugulosa Weise, 1901*
Epilachna ostensoides Gordon, 1975 Toxotoma soukupi Gordon, 1975*
Epilachna oviforma Gordon, 1975* Toxotoma townsendi Gordon, 1975*
Epilachna pachiteensis (Weise, 1926)* Toxotoma tridentata Gordon, 1975*
Epilachna paenulata (Germar, 1824) Toxotoma venusta (Erichson, 1847)*
Epilachna parastriata Gordon, 1975* Toxotoma weuyrauchi Gordon, 1975*
Madaini Gordon, 1875

Eremochilini Gordon and Vandenberg, 1987 Lorma paprzyckii Gordon, 1975*
Eremochilus peregrinus Weise, 1912 Mada amazona (Weise, 1926)
Mada circumflua (Mulsant, 1850) Mada insolitaphallus Gordon, 1975*

Mada elegans Gordon, 1975* Mada nexophallus Gordon, 1975*
(1)
Species whose types have not been located in the revisions to the corresponding
genera or that are located in genera not present in South America, and whose generic
location is uncertain.
Acknowledgements
I thank Abdhiel Bustamante and Anahi Oroz (Universidad Nacional San Antonio
Abad, Cusco, Peru), Jimmy Miro and Pedro Castillo (Museo de Entomologa de la
Universidad Nacional de Tumbes, Tumbes, Peru), Luis Figueroa and Angelico
Asenjo (Museo de Historia Natural de la Universidad Nacional Mayor de San
Marcos, Lima, Peru), Clorinda Vergara (Museo de Entomologa Klaus Raven
Buller, Universidad Nacional Agraria La Molina, Lima, Peru), Karina Vilca
(Universidad Nacional Santiago Antunez de Mayolo, Ancash, Peru), Pedro Lozada
(Servicio Nacional de Sanidad Agraria, Peru), Juan Enrique Barriga-Tunon (Curico,
Chile), Natalia Vandenberg (USDA Systematic Entomology Laboratory Staff,
Washington DC, USA), and all specialists and collectors who sent me materials
from their collections or collections that they represent. Special thanks to Manuel
Dieguez (Santiago, Chile) who managed multiple loans from Peruvian institutions
to the author. Also, I thank Adriano Giorgi (Pernambuco, Brazil) for his smart
revision of the manuscript, and to the anonymous reviewer who indicated many
corrections to the checklist. I acknowledge NSF-EPSCoR #66928 (PI: CS Chaboo)
for supporting the Beetles of Peru and the University of Kansas Department of
Ecology and Evolutionary Biology-General Research Fund (PI: CS Chaboo) for
funding this publication.
Literature Cited
Almeida, L. M., and P. B. Santos. 2014. Synopsis of Oryssomini Gordon (Coleoptera: Coccinellidae) from
the Neotropical region with new species of Oryssomus Mulsant, Pseudoryssomus Gordon and
Gordonoryssomus Almeida and Lima. Zootaxa 3846:042068.
Bouchard, P., Y. Bousquet, A. E. Davies, M. A. Alonso-Zarazaga, J. F. Lawrence, C. H. C. Lyal, A. F.
in Coleoptera (Insecta). ZooKeys 88:1972.
Indies, and South America, Part. 3. United States National Museum Bulletin 185:343550.
Canepari, C., R. D. Gordon, and G. A. Hanley. 2013. South American Coccinellidae (Coleoptera), Part XV:
systematic revision of Dilatitibialis Duverger (Coccidulinae; Hyperaspidini). Insecta Mundi 0312:191
Gonzalez, G. 2007. Los Coccinellidae de Peru [online]. URL: http://www.coccinellidae.cl.PaginasWeb
Peru/Paginas/InicioPeru.php. (accessed 17 May 2014).
Gonzalez, G. 2010. Actualizacion de la bibliografa y nuevos registros en Coccinellidae de America del Sur
(Insecta: Coleoptera). Boletn de la Sociedad Entomologica Aragonesa (S.E.A) 47:245256.
Gonzalez, G. 2012. Revision de los generos Coccidophilus Brethes y Microweisea Cockerell (Coleoptera:
Coccinellidae: Microweiseinae) en America del Sur. Boletn de la Sociedad Entomologica
Aragonesa (S.E.A) 51:6188.
Gonzalez, G. 2013a. Gordonitan. gen. y otros aportes al conocimiento de los Chnoodini de America del
Sur (Coleoptera Coccinellidae) Boletn de la Sociedad Entomologica Aragonesa (S.E.A) 53:6379.
Gonzalez, G. 2013b. Lista y distribucion de especies de Coccinellidae (Insecta: Coleoptera) presentes en
Paraguay. Boletn del Museo Nacional de Historia Natural de Paraguay 17:162.
Gonzalez, G. 2014a. Especies nuevas del genero Eriopis Mulsant (Coleoptera: Coccinellidae) del norte de
Chile. Boletn de la Sociedad Entomologica Aragonesa (S.E.A) 54:6172
Gonzalez, G. 2014b. Una nueva especie del genero Coleomegilla Timberlake (Coleoptera: Coccinellidae)
de America del Sur. Boletn de la Sociedad Entomologica Aragonesa (S.E.A) 54:109112.
Gonzalez, G., and R. Honour. 2011. Especies nuevas del genero Diomus Mulsant (Coleoptera,
Coccinellidae) de America del Sur. Boletn de la Sociedad Entomologica Aragonesa (S.E.A)
49:114.
Gonzalez, G., and R. Honour. 2012. Tres nuevas especies del genero Zenoria (Coleoptera: Coccinellidae)
con diseno similar aZ. discoidalis (Kirsch) Boletn de la Sociedad Entomologica Aragonesa (S.E.A)
50:175181.
Gonzalez G & T. Kondo, 2014. Geographical distribution and phenotypic variation of Anovia punica
Gordon (Coleoptera: Coccinellidae: Noviini), a predatory ladybeetle of fluted scales (Hemiptera:
Coccoidea: Monophlebidae). Insecta Mundi 0398 (2014):16.
Gordon, R. D. 1986. Additions to the Peruvian fauna of the plant feeding genus Epilachna Chevrolat.
Revista Colombiana de Entomologia 12:35.
Gordon, R. D., C. Canepari, and G. A. Hanley. 2013. South American Coccinellidae (Coleoptera), Part
XII: New name for Cyra Mulsant, review of Brachiacanthini genera, and systematic revision of
Cleothera Mulsant, Hinda Mulsant and Serratitibia Gordon and Canepari, new genus. Insecta
Mundi 0278:1150.
Gordon, R. D., C. Canepari, and G. A. Hanley. 2014. South American Coccinellidae (Coleoptera), Part XVI:
systematic revision of Brachiacantha Dejean (Coccinellinae: Hyperaspidini). Insecta Mundi 0390:176.
Gordon, R. D., and G. Gonzalez. 2002. South American Coccinellidae (Coleoptera). Part IX: a systematic
revision of Scymnobius Casey (Scymninae: Scymnini). Frustula Entomologica 25:5785.
Gordon, R. D., and G. Gonzalez. 2011. Additions to the Hyperaspis Chevrolat (Coleoptera: Coccinellidae)
fauna of South American, descriptions of nine new species, and recognition of Hyperaspis pectoralis
Crotch as a valid species. Insecta Mundi 160:120.
Jadwiszczak, A., and P. Wegrzynowicz. 2003. World catalogue of Coccinellidae. Part I Epilachninae.
Mantis, Olsztyn, Poland. 264 pp.
Miro, J., and P. Castillo. 2010. Especies de mariquitas (Coleoptera: Coccinellidae) en los frutales de
Tumbes. Revista Peruana de Entomologa 46:17.
Slipinski, A., and W. Tomaszewska. Coccinellidae Latreille, 1802. In Leschen, R. A. B., R. G. Beutel, and
J. F. Lawrence (eds.). Handbook of Zoology, Vol. 2, Coleoptera, pp. 454472. Berlin/New York:
Walter de Gruyter GmbH and Co. KG. XIII + 786 pp.
Vandenberg, N. J. 2002. Family 93. Coccinellidae Latreille 1807. In Arnett, R. H., Jr., M. C. Thomas, P.
E. Skelley, and J. H. Frank (eds.). American Beetles. Vol. 2. Polyphaga: Scarabaeoidea through
Curculionoidea, pp. 371389. CRC Press LLC, Boca Raton, FL, xiv + 861 pp.
Trogossitidae
Author(s): Ji Kolib

terms_of_use.
critical research.
88(2), 2015, pp. 237239
Beetles (Coleoptera) of Peru: A Survey of the Families. Trogossitidae

JIRI KOLIBAC
Moravian Museum, Department of Entomology, Hviezdoslavova 29a, 627 00 Brno,
Czech Republic
e-mail: jkolibac@mzm.cz
Diversity in Peru: 1 subfamily, 1 tribe, 2 genera, 4 species.

Recognition: This small family has about 600 species world-wide. Adult. Body wide
and flattened in Lophocaterinae, flat or conglobate in Peltinae and Rentonium-group,
elongate and cylindrical in Trogossitinae. Body mostly bare or sparsely pubescent but
sometimes also with tufts of setae, scales, dense pubescence or long hairs. Anterior
margin of frons straight to very deeply emarginate, with two horns at sides.
Antennal insertions partly covered by edge of frons or visible in dorsal view. Eyes not
emarginate or posteriorly emarginate. Antennae 811-segmented, with conspicuous
13-segmented club or with widened distal antennomeres that may be distinctly
asymmetrical in Trogossitinae. Mandible bidentate, rarely unidentate (Corticotomus).
Male submentum with tuft of setae (ctenidium) in some Trogossitinae. Pronotum
usually transverse in Peltinae and Lophocaterinae; elongate in some Trogossitinae
(Nemozoma, Airora). Lateral carina almost always present, reduced only in some
Trogossitinae (Corticotomus). Procoxa transverse. Coxal cavities internally open and
externally closed (Trogossitinae) or widely open (Peltinae, Lophocaterinae). Elytra
usually regularly punctate, with or without carinae. Metacoxae extending laterally to
meet elytral epipleura. Wings usually present. Radial cell as long as wide or shorter
than wide, sometimes very reduced or absent. Medial field with two to four veins,
wedge cell and no medial fleck. Trochanters triangular. Femora sometimes clavate.
Tibiae often with row of spines along outer side; apex of tibia with row of spines and
two spurs (one of them hooked) or only with hooked spur or spurs reduced. Tarsi
5-5-5; basitarsus sometimes partially fused with tarsomere 2 and tarsal pattern
seemingly 4-4-4 or 4-4-5; tarsomeres 14 never with membranous lobes; terminal
tarsomere usually as long as combined length of tarsomeres 14; claws large,
without denticles; empodium bisetose, strongly projecting. Abdomen with five or
six ventrites. Aedeagus cucujiform, usually inverted. Tegmen usually with anterior
ventral strut and two opposing dorsal struts, and composed of two or three parts.
Penis with two anterior struts. Ovipositor lightly sclerotized, except for baculi,
moderate in size with sparsely pubescent coxites and styli.
Larva: Body elongate, cylindrical. Color white or pale, but sclerotized areas
distinctly pigmented (head capsule, thoracic and abdominal terga, and urogomphi).
Median endocarina usually present, usually extending between frontal arms. Frontal
arms V-shaped, straight (Trogossitinae, Lophocaterinae) or curved (Peltinae). Five
stemmata usually present and arranged in a pattern with two anteriorly and three
in a posterior row. Mandibles bidentate; mola absent (Trogossitinae, part of
Lophocaterinae) or present (Peltinae, part of Lophocaterinae. Prothorax usually
with one large dorsal sclerite. Meso- and metathorax usually with pair of sclerites
Accepted 29 April 2015; Revised 10 September 2015

dorsally and one weakly sclerotised, pale plate ventrally or ventral sclerites absent.
Urogomphi usually well-developed, hook-shaped or nearly straight; strongly
sclerotized and pigmented; often with spines or secondary processes (Kolibac, 2013).
Habitat: The larvae and adults of Trogossitidae are either mycophagous (Peltinae,
Lophocaterinae), predatory (Trogossitinae, Lophocaterinae) or anthophilous and
partially feeding on pollen grains (some Lophocaterinae). Most trogossitines and
lophocaterines are predacious with both larvae and adults living under bark, in
galleries of xylophagous insects or on logs and branches where hunt for insects.
However, some species are also mycophagous and one unidentified Peruvian
Temnoscheila sp. was found as larvae and adults feeding on an Ascomycete.
Tenebroides mauritanicus L. is a pest species feeding on stored grains, similarly as
Lophocateres pusillus Klug. Larvae and adults of cylindrical species (Nemozoma,
Corticotomus, Airora), live in galleries of, for example, Scolytinae and Bostrichidae.
The members of the Rentonium-group can be extracted from forest litter or rotten
wood. Detailed biological data are mostly unknown except for the two above-
mentioned pests (see the worldwide web; Kolibac and Leschen, 2010; Kolibac, 2014).
Notes: The following checklist was derived from Blackwelder (1945). No new
reports are known from Peru according to the recent world review by Kolibac
(2013). The morphology, phylogeny, distribution and classification of the family
have been revised by Kolibac (2005, 2006, 2008) and Kolibac and Leschen (2010).
The latest key to Trogossitidae as well as descriptions of new Central American taxa
have been given by Lawrence et al. (2014).
The trogossitid fauna of Peru is apparently unsufficiently known. The species-rich
genera Temnoscheila Westwood and Tenebroides Piller et Mitterpacher are certainly
more frequently represented in the country than by the few species listed below; records
of more genera can be expected. Considering findings in neighboring countries,
members of the following genera could be also distributed in Peru: Airora Reitter,
Calanthosoma Reitter, Nemozoma Latreille, Corticotomus Sharp (all Trogossitinae)
and Leptonyxa Reitter (Lophocaterinae). Members of the Rentonium-group cannot be
excluded. Two more cosmopolitan pests of stored products (e.g. food, grain) surely
occur in Perus Pacific ports: Tenebroides mauritanicus L. (commonly called the cadelle
beetle; Trogossitinae) and Lophocateres pusillus Klug (commonly called the Siamese
Grain beetle; Lophocaterinae). A lack taxonomic specialists suggests that world
diversity of Trogossitidae is under-estimated; the University of Kansas collection
(SEMC) has more 120 undetermined specimens from various parts of Peru.
Checklist:
Subfamily Trogossitinae: Tribe Trogossitini

Temnoscheila chalcea Kirsch, 1873 Temnoscheila rugulosa Kirsch, 1873
Temnoscheila peruviana Leveille, 1907 Tenebroides dilatatus Erichson, 1847
Acknowledgements
Evolutionary Biology-General Research Fund (PI: CS Chaboo). Jir Kolibac is
supported by a long-term conceptual development program for research institutions

by the Ministry of Culture of the Czech Republic (ref. MK000094862).
Literature Cited
Indies, and South America. Parts 1 through 6. U. S. National Museum Bulletin 185:ixii, 11492.
Kolibac, J. 2005. A review of the Trogossitidae. Part 1: morphology of the genera (Coleoptera, Cleroidea).
Entomologica Basiliensia et Collectionis Frey 27:39159.
Kolibac, J. 2006. A review of the Trogossitidae. Part 2: Larval morphology, phylogeny and taxonomy
(Coleoptera, Cleroidea). Entomologica Basiliensia et Collectionis Frey 28:105153.
Kolibac, J. 2008. Morphology, taxonomy and phylogeny of Phloiophilus edwardsi Stephens, 1830
(Coleoptera, Cleroidea). Entomologica Basiliensia et Collectionis Frey 30:105133.
Kolibac, J. 2013. Trogossitidae: a review of the beetle family, with a catalogue and keys. ZooKeys
366:1194. doi:10.3897/zookeys.366.6172.
Kolibac, J. 2014. Nemozoma gymnosternalis sp. nov., a new anomalous species of Trogossitidae from
Brazil. Zootaxa 3815:417427. doi:10.11646/zootaxa.3815.3.7.
Kolibac, J., and R. A. B. Leschen. 2010. Trogossitidae Fabricius, 1801. In R. Leschen, R. G. Beutel, and
J. F. Lawrence (eds.), Handbuch der Zoologie/Handbook of Zoology. Band 4: Arthropoda,
2. Halfte: Insecta. Part 39: Coleoptera, Beetles Vol. 2, pp. 241247. W. de Gruyter, Berlin-New
York, 812 pp.
Lawrence, J. F., R. A. B. Leschen, and A. Slipinski. 2014. Antillipeltis, a new genus of Antillean
Trogossitidae (Coleoptera: Cleroidea) with a key to the Cleroidea. Zootaxa 3794:435454.
Phengodidae LeConte, 1861
Author(s): Caroline S. Chaboo, Matthew L. Gimmel, and Milada Bocakova

terms_of_use.
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88(2), 2015, pp. 240242
Beetles (Coleoptera) of Peru: A Survey of the Families. Phengodidae

LeConte, 1861
CAROLINE S. CHABOO,1,3 MATTHEW L. GIMMEL,2 AND MILADA BOCAKOVA2

Recognition: Phengodidae are commonly called glow-worms because the larviform
females and larvae have bioluminescent organs in transverse bands along the body.
They glow yellow-green using eleven paired photic organs of 2nd thoracic through
9th abdominal segments; some genera have head organs that glow red. Some adult
males are also bioluminescent, and at least one species has bioluminescent eggs. The
adult male body length is 3.325 mm; larviform females are generally larger than
males. The antenna has 1012 antennomeres in adult males, and may be filiform,
serrate, bipectinate or plumose. Adult males have truncate or dehiscent elytra.
Phengodid larvae are slender, mostly well-sclerotized, and rounded in cross-
section (compared to the flattened larvae of Lampyridae and Lycidae), resembling
those of click beetles.
Habitat: These may be nocturnal, active at nights. Winged males may be attracted
at light traps. Females may be collected from loose bark while larvae are usually
found under leaf litter or rotting logs. The larviform females and larvae are specialist
predators of millipedes and other litter arthropods.
Notes: The American family Phengodidae is now considered as separate from
Asiatic Rhagophthalmidae, although sister group relationships of the families were
supported by both morphological evidence (Lawrence et al., 1995), and compre-
hensive molecular phylogenies (Bocakova et al., 2007; Kundrata et al., 2014).
Previous studies suggested rhagophthalmids as a clade of fireflies (Crowson, 1955;
Suzuki, 1997).
The following checklist is derived from Pic in Titschack (1954; as Drilidae),
Zaragoza-Caballero (1984), and specimens housed in SEMC. The family was
monographed by Wittmer (1976), but a number of new genera have been added
since. Lawrence et al. (2000) reported the minute Penicillophorus (24 mm) as
occurring in Colombia and Peru. The railroad worm, Phrixothrix, occurs in Peru
but it is unclear if it is a pest (see Costa et al., 1999). An interactive key to the world
fauna is available online (Lawrence et al., 2000); see Zaragoza-Caballero (2010) for
an updated key to the genera of the large subfamily Mastinocerinae.
Taximastinocerus is reported as new genus record for Peru based on 19 specimens
(probably multiple species) collected by Chaboo in 20102011 with flight intercept
traps and Malaise traps at the CICRA Biological Field Station, Madre de Dios,
Peru.
1
2
Department of Biology, Faculty of Education, Palacky University, Purkrabska 2, CZ-77140 Olomouc,
Czech Republic.
3
Accepted 23 July 2014; Revised 24 March 2015
Wittmer (1963) described the genus Oxymastinocerus Wittmer, 1963 and

designated Phrixothrix peruanus Wittmer as the type species (p. 96). As Phrixothrix
peruanus Wittmer, 1963 was proposed in the same arcticle (p. 78), Wittmer (1963)
hardly intended to designate this species as the type species of Oxymastinocerus.
Instead, Wittmer (1963) probably intended to designate as the type species of
Oxymastinocerus the first species placed in the new genus, i.e., Oxymastinocerus
peruanus (Wittmer) and stated that he originally placed the species in the genus
Mastinocerus (Ent. Arb. Museum Frey 7, 1956, p. 225). As Wittmer (1956) described
Mastinocerus peruvianus (nec peruanus), it seems to be another lapsus calami.
Therefore, now we take the advantage of the article 67.6 of ICZN (1999) and
consider Wittmers (1963) type species designation as the use of incorrect spelling.
Consequently, Mastinocerus peruvianus Wittmer, 1956 is considered to be the type
species of the genus Oxymastinocerus.
Checklist:
Mastinocerinae
Brasilocerus opacus (Pic, 1937) Phrixothrix obscurus Pic, 1915
Mastinocerus (Paramastinocerus) peruanus Wittmer, Phrixothrix peruanus Wittmer, 1963
1976 Stenophrixothrix pallens (Berg, 1885)
Oxymastinocerus peruvianus (Wittmer, 1956) Taximastinocerus undetermined species
Phengodinae
Pseudophengodes cincinnata (Erichson, 1847) Pseudophengodes floccosa (Erichson, 1847)
Pseudophengodes flavicollis (Leach, 1824) Pseudophengodes pulchella (Guerin-Meneville, 1843)
Penicillophorini
Penicillophorus undetermined species
Acknowledgements
We acknowledge NSF-EPSCoR #66928 (PI: CS Chaboo) for supporting the Peru
beetle project and the University of Kansas Department of Ecology and
publication. This study was also supported by grants of the European Social Fund
and the Ministry of Education of the Czech Republic (CZ. 1.07/2.3.00/30.0004 to
Matthew L. Gimmel and CZ.1.07/2.3.00/20.0166 to Milada Bocakova).
Literature Cited
Bocakova, M., L. Bocak, T. Hunt, M. Teravainen, and A. P. Vogler. 2007. Molecular phylogenetics of
Elateriformia (Coleoptera): evolution of bioluminescence and neoteny. Cladistics 23:477496.
Constantin, R. 2014. Contribution a la connaissance des Phengodidae de Guyane et description de huit
especes nouvelles (Coleoptera, Elateroidea).: Coleopteres de Guyane. Tome VIII. Le Coleopteriste
(Supplement), 86104.
Costa, C., S. A. Vanin, S. A. Casari, and V. R. Viviani. 1999. Larvae of Neotropical Coleoptera. XXVII.
Phrixothrix hirtus: immatures, neotenic female, adult male and bionomic data (Phengodinae,
Phengodidae, Coleoptera). Iheringia, Serie Zoologica 86:928.
Crowson, R. A. 1955. The natural classification of the families of Coleoptera. Nathaniel Lloyd & Co.,
London.
Kundrata, R., M. Bocakova, and L. Bocak. 2014. The comprehensive phylogeny of the superfamily
Elateroidea (Coleoptera: Elateriformia). Molecular Phylogenetics and Evolution 76:162171.
Lawrence, J. F., A. M. Hastings, M. J. Dallwitz, T. A. Paine, and E. J. Zurcher. 2000 and onwards.
Elateriformia (Coleoptera): descriptions, illustrations, identification, and information retrieval for
families and subfamilies. Version: 9th October 2005. http://delta-intkey.com [Accessed 2 June 2014].
Lawrence, J. F., N. B. Nikitsky, and A. G. Kirejtshuk. 1995. Phylogenetic position of Decliniidae
(Coleoptera: Scirtoidea) and comments on the classification of Elateriformia (sensu lato). In
Lawrence, J. F., J. Pakaluk, and S. A. Slipinski (eds.). Biology, phylogeny, and classification of
Coleoptera. Muzeum i Institut Zoologii PAN, Warszawa, pp. 375410.
Suzuki, H. 1997. Molecular phylogenetic studies of Japanese fireflies and their mating systems
(Coleoptera: Cantharoidea). TMU Bulletin of Natural History 3:153.
Titschack, E. 1954. Beitrage zur fauna perus. Band iv Wissenschaftliche Bearbeitungen, 1954, Hamburger
Sudperu-Expedition 1936. Verlag von Gustav Fischer, Jena, 4:386 pp.
Wittmer, W. 1956. 19. Beitrag zur Kenntnis der neotropischen malacodermata (Col.). Entomologische
Arbeiten aus dem Museum G. Frey 7:217226.
Wittmer, W. 1963. Zur Kenntnis der Phengodidae (Coleoptera) (25. Beitrag zur Kenntnis der
neotropischen Malacodermata). Mitteilungen der Schweizerischen Entomologischen Gesselschaft
36:7399.
Wittmer, W. 1976. Arbeiten zu einer Revision der Familie Phengodidae (Coleoptera). Entomologische
Arbeiten aus dem Museum G. Frey 27:415524.
Zaragoza-Caballero, S. 1984. Catalogo de la familia Phengodidae (Coleoptera). Anales del Instituto de
Biologa, Universidad Nacional Autonoma de Mexico, Serie Zoologa 55:307324.
Zaragoza-Caballero, S. 2010. Paraphrixothrix ecuadoranus, a new genus and species of Phengodidae
(Coleoptera: Mastinocerinae) from Ecuador. Pan-Pacific Entomologist 86:105110.
Beetles (Coleoptera) of Peru: A Survey of the Families. Lycidae
Laporte, 1836
Author(s): Milada Bocakova, Matthew L. Gimmel, and Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 243247
Beetles (Coleoptera) of Peru: A Survey of the Families. Lycidae

Laporte, 1836
MILADA BOCAKOVA,1 MATTHEW L. GIMMEL,1 AND CAROLINE S. CHABOO2,3
Diversity in Peru: 1 subfamily, 6 tribes, 23 genera, and 126 species.

Recognition: Lycidae, or net-winged beetles, have adults mostly about 428 mm
long and often brightly colored (Bocak and Bocakova, 2008). Their aposematic
colors, tendency to aggregate, and spontaneous bleeding are attributed to their
chemical defenses; some may be involved in mimicry complexes. The six tribes of
Peruvian lycids have mostly yellow and dark brown coloration. One of the most
frequent of the various color patterns is the cross pattern (elongate dark brown cross
on yellow to red background), or yellow coloration forming transverse or
longitudinal stripes.
The Lycidae are soft-bodied as they have thin, weakly sclerotized cuticle. The
adult body is dorsoventrally flattened and head sometimes elongate forming
a rostrum in floricolous taxa. The pronotum often bears costae forming areolae and
the flexible elytra are reinforced by longitudinal costae and distinct reticulate cells
thus, their common name as net-winged beetles. Among related soft-bodied elateroid
beetles, the Lycidae can be distinguished by elongate trochanters with femoral
articulation distally, while the trochanters of Lampyridae and Cantharidae are small
with femoral articulation oblique.
Other morphological features (Bocak and Bocakova, 1990; Bocakova, 2003) of
lycid adults are: head prognathous to hypognathous, partly covered by pronotum,
antennae 11-segmented, filiform to flabellate, mandibles arcuate, pronotum oblong
to trapezoidal, scutellum rectangular, slightly emarginate distally. The elytra are
mostly parallel, sometimes widened posteriorly (Calopteron), each elytron with 39
longitudinal costae, legs compressed, trochanters elongate to triangular, tarsal
segments 34 lobed, abdomen with 7 (males) and 8 (females) visible sterna, male
genitalia mostly trilobate, parameres shortened (as long as K2/3 of phallus in
Calopterini, Neolycus), or absent (Plateros), female genitalia with coxites mostly
separate, paraproctal baculi mostly rod-like, sometimes widened and fused basally,
styli setose.
Lycid larvae are flattened, with the middle part of abdomen sometimes widened
(Lycini). Their body is weakly sclerotized, similar to the adults; the head is
prognathous, partly retracted into pronotum, and sometimes elongate (Lycini).
Stemmata are absent or reduced to one stemma on each side of the head. The
antenna is 2-segmented (while 3-segmented in larvae of Lampyridae), mandibles
falciform, entire mandible cleft longitudinally; sometimes with paired urogomphi
(Bocak and Matsuda, 2003).
1
Department of Biology, Faculty of Education, Palacky University, Purkrabska 2, CZ-77140 Olomouc,
Czech Republic.
2
Division of Entomology, Biodiversity Institute,1501 Crestline Drive, Suite 140, University of Kansas,
Lawrence, Kansas 66045, USA, e-mail: cschaboo@ku.edu
3
Accepted 22 April 2014; Revised 14 October 2014
The head of Calochromini and Lycini is anteriorly elongated forming

a rostrum, while that of other tribes is short. Lycini have median longitudinal
carina in the anterior half of the pronotum; each elytron bears 4 longitudinal
costae and irregular reticulate cells. Conversely, the pronotum of Calochromini
has only lateral folds (pronotal carinae absent), and each elytron has 9
longitudinal costae. Erotini adults have a closed median areola on the
pronotum.
Most Peruvian lycids belong in the tribe Calopterini with distal portion of phallus
dorsoventrally flattened, female terminal sternum having short spiculum ventrale,
and paraproctal baculi rod-like, fused basally, forming median bridge. The
Eurrhacini (separated from Calopterini recently by DNA data) have male terminal
sternum basally elongate (i.e., attachment of tergum 9 almost in median portion) and
phallobase distorted clockwise.
The Platerodini have antennal tubercles inconspicuous, pronotum with median
longitudinal line channeled at base and carinae absent, each elytron with nine
longitudinal costae (primary and secondary costae not distinguished, of similar
strength), reticulate cells rather oval, and paraproctal baculi basally widened.
Habitat: Adults are found typically in rainforests on vegetation, namely on the
bottom side of the leaves and mostly do not feed in the adult stage. Because of the
thin cuticle and limited ability to avoid desiccation they preferentially fly in
morning and evening hours avoiding hot weather. Some Calopterini (e.g.,
Calopteron, Idiopteron), Calochrominae, and Lycinae are often aposematically
colored and are usually collected in dry forests, savannah, and rural areas where
the adults may feed on nectar. Larvae are found usually in shaded habitats in
dead wood, under bark or in leaf litter where they feed on fermented juices (Bocak
and Matsuda, 2003). Larvae of some lycid genera (e.g., Calopteron, Calochromus,
Neolycus) live on tree trunks in sunny habitats, or even in charred trunks on burnt
forest clearings. Calopteron and Neolycus larvae have often been found
aggregated.
Notes: We follow the classification of Bocak and Bocakova (2008) which is based
on the latest morphological (Bocakova, 2005) and molecular phylogenies of Lycidae
(Bocak et al., 2008). Sources of names for the checklist are updated from published
catalogues (Kleine, 1933; Blackwelder, 1945). Additional records and nomenclatural
acts are documented in Kleine (1941, 1942), Bocakova (2001, 2003), Nascimento and
Bocakova (2009), Barancikova et al. (2010), Kazantsev (2011), Bocakova et al.
(2012), and Bocakova and Nascimento (2013).
Idiopteron homonymum Blackwelder, 1945 syn. n. is considered to be a younger
objective synonym of Idiopteron callanganum Pic, 1932. We transfer the following
taxa from the genus Calleros to Plateros: Plateros bogotensis grandescapularis (Pic,
1934) comb. n., P. clermonti (Pic, 1934) comb. n., P. conjunctus (Pic, 1934) comb. n.,
P. diosensis (Pic, 1922) comb. n., P. diversecostatus (Pic, 1922) comb. n., P.
marcapatanus (Pic, 1934) comb. n., P. minimus (Pic, 1931) comb. n., P. peruvianus
(Pic, 1922) comb. n., P. postseparatus (Pic, 1930) comb. n., P. punoensis (Pic, 1934)
comb. n., P. quadrinotatus (Pic, 1922) comb. n., P. reductenotatus (Pic, 1934) comb. n.,
P. scapularis (Pic, 1934) comb. n., P. subfasciatus (Pic, 1930) comb. n., P. subfasciatus
kraatzi (Pic, 1934) comb. n., P. terminalicornis (Pic, 1931) comb. n., and. P.
unifasciatus (Pic, 1934) comb. n. We also transfer Leptoceletes subtilis (Kleine, 1941)
comb. n. from the genus Celetes.
Checklist:
Subfamily Lycinae
Lycini
Neolycus sp. *Peru: Villa Rica, Pasco prov., 8. xii. 2013 [2, UPOL]
Calochromini
Lygistopterus peruvianus Kleine, 1941
Calopterini
Brasilycus peruanus Bocakova et Nascimento, 2013 Flabellolycinella croceus Kleine, 1941
Caenia mirifica Kirsch, 1873 Idiopteron breveapicale Pic, 1922
Caenia peruviana Kirsch, 1873 Idiopteron brevenotatum Pic, 1922
Calopteron affine Lucas, 1857 Idiopteron callanganum Pic, 1932
Calopteron atrithorax Pic, 1923 (5Idiopteron homonymum Blackwelder, 1945)
Calopteron basale Kirsch, 1865 syn. n.
Calopteron confrater Taschenberg, 1874 Idiopteron nigricolle Kleine, 1941
Calopteron consulare Erichson, 1847 Idiopteron peruvianum Pic, 1933
Calopteron costatulum Kirsch, 1865 Idiopteron signatum Pic, 1922
Calopteron cyaneum Erichson, 1847 Idiopteron subapicale Pic, 1922
Calopteron dichroum Kirsch, 1865 Idiopteron subapicale elevatum Pic, 1933
Calopteron diffusicoloratum Kleine, 1942 Leptoceletes subtilis (Kleine, 1941) comb. n.
Calopteron fallax Kirsch, 1865 Lycomorphon angusticolle (Pic, 1922)
Calopteron lebasi Bourgeois, 1878 Lycomorphon elongaticolle (Pic, 1922)
Calopteron melanoxanthum Kirsch, 1870 Lycomorphon diversicolle (Pic, 1931)
Calopteron montiphilum Kleine, 1942 Lycomorphon irregularis (Pic, 1931)
Calopteron parens Bourgeois, 1899 Lycomorphon opacum (Pic, 1926)
Calopteron parens minor Pic, 1931 Mesopteron acrocirrhum Kirsch, 1873
Calopteron phaleratum Blanchard, 1843 Mesopteron amoenum Kirsch, 1865
Calopteron quadraticolle Taschenberg, 1874 Mesopteron carinatum Pic, 1922
Calopteron reticulatum Fabricius, 1775 Mesopteron crassicornis Kleine, 1941
Calopteron serratum Linnaeus, 1758 Mesopteron diosanum Pic, 1928
Calopteron serratum abdominale Lucas, 1857 Mesopteron infirmum Kirsch, 1873
Calopteron subcruciatum Bourgeois, 1878 Mesopteron inhumerale Pic, 1922
Calopteron subparallelum Pic, 1938 Mesopteron minutum Pic, 1922
Calopteron terminatum Latreille, 1833 Mesopteron peruvianum Pic, 1922
Calopteron terminatum nigricorne Latreille, 1833 Mesopteron plurinotatum Pic, 1922
Calopteron torquatum Erichson, 1847 Mesopteron prolongaticolle Pic, 1939
Calopteron torquatum albofasciatum Bourgeois, Mesopteron reductum Pic, 1922
1905 Mesopteron scitum Pic, 1922
Ceratopriomorphus piceus (Pic, 1922) Mesopteron staudingeri Pic, 1922
Cyrtopteron acroleucum Kirsch, 1873 Mesopteron subflabellicorne Pic, 1939
Cyrtopteron fasciatum Pic, 1922 Mesopteron titschacki Kleine, 1941
Cyrtopteron muhlenbecki Bourgeois, 1879 Mesopteron unifasciatum Pic, 1922
Cyrtopteron obliquum Pic, 1922 Metapteron atronotatum Pic, 1922
Falsocaenia apicicornis (Pic, 1922) Metapteron diffusum Kleine, 1941
Falsocaenia longehumeralis (Pic, 1922) Metapteron humerale Kirsch, 1873
(5Falsocaenia callangana (Pic, 1922)) Metapteron observans Kleine, 1941
Falsocaenia grisea Kleine, 1941 Metapteron suturale Latreille, 1833
Eurrhacini
Emplectus angustatus Pic, 1922 Emplectus foveolatus Pic, 1923
Emplectus apicicornis Pic, 1923 Emplectus minor Pic, 1923
Emplectus bicoloricornis Pic, 1923 Emplectus platythorax Kleine, 1941
Emplectus bimaculatus Erichson, 1847 Emplectus peruvianus Pic, 1923
Emplectus callanganus Pic, 1923 Emplectus trinotatus Pic, 1923
Emplectus fatigans Kirsch, 1873 Emplectus vicinus Pic, 1923
Linoptes atronotatus Pic, 1922 Haplobothris longeprolongata Pic, 1938

Lycoplateros mimicus Pic, 1922 Haplobothris peruviana Pic, 1911
Haplobothris apicicornis Pic, 1931
Erotini
Eros crocatus Erichson, 1847
Eros decoratus Erichson, 1847
Eros patruelis Kirsch, 1873
Platerodini
Cavoplateros ater Pic, 1922 Plateros mediotestaceus Pic, 1923
Cavoplateros vicinus Pic, 1922 Plateros minimus (Pic, 1931) comb. n.
Picomicrolycus pilosus (Pic, 1934) Plateros mosolovi Kazantsev, 2011
Plateros amazonensis Kazantsev, 2011 Plateros nautaensis Kazantsev, 2011
Plateros atricolor Pic, 1923 Plateros parvulus Kirsch, 1873
Plateros basicornis Pic, 1925 Plateros peruvianus (Pic, 1922) comb. n.
Plateros bimaculatus Kirsch, 1873 Plateros postseparatus (Pic, 1930) comb. n.
Plateros bogotensis grandescapularis Plateros punoensis (Pic, 1934) comb. n.
(Pic, 1934) comb. n. Plateros quadrinotatus (Pic, 1922) comb. n.
Plateros clermonti (Pic, 1934) comb. n. Plateros reductenotatus (Pic, 1934) comb. n.
Plateros conjunctus (Pic, 1934) comb. n. Plateros scapularis (Pic, 1934) comb. n.
Plateros crocatus Pic, 1922 Plateros semilimbatus Pic, 1922
Plateros diosensis (Pic, 1922) comb. n. Plateros speciosus Kazantsev, 2011
Plateros diversecostatus (Pic, 1922) comb. n. Plateros subapicalis Pic, 1922
Plateros duomaculatus Kazantsev, 2011 Plateros subfasciatus (Pic, 1930) comb. n.
(5Calleros bimaculatus Pic, 1922) Plateros subfasciatus kraatzi (Pic, 1934) comb. n.
Plateros elongatipennis Pic, 1923 Plateros subhumeralis Pic, 1923
Plateros grisescens Pic, 1923 Plateros terminalicornis (Pic, 1931) comb. n.
Plateros inlineatus Pic, 1923 Plateros uhmanni Kleine, 1941
Plateros limbatipennis Pic, 1923 Plateros unifasciatus (Pic, 1934) comb. n.
Plateros longipennis Pic, 1923 Plateros zakharovi Kazantsev, 2011
Plateros marcapatanus (Pic, 1934) comb. n. Teroplas petrovi Kazantsev, 2011
Acknowledgements
Beetles of Peru project and the University of Kansas Department of Ecology
and Evolutionary Biology-General Research Fund (PI: CS Chaboo). This study
was also supported by grants of Palacky University Olomouc, Czech Republic
(IGA_PdF_2015_029 to Milada Bocakova), the European Social Fund and the
Ministry of Education of the Czech Republic (CZ.1.07/2.3.00/20.0166 to Milada
Bocakova and CZ. 1.07/2.3.00/20.0004 to Matthew L. Gimmel).
Literature Cited
Barancikova, B., E. A. Nascimento, and M. Bocakova. 2010. Review of the genus Ceratopriomorphus Pic,
1922 (Coleoptera: Lycidae). Zootaxa 2683:6165.
Bocak, L., and M. Bocakova. 1990. Revision of the suprageneric classification of the family Lycidae
(Coleoptera). Polskie Pismo Entomologiczne 59:623676.
Bocak, L., and M. Bocakova. 2008. Phylogeny and classification of the Family Lycidae (Insecta:
Coleoptera). Annales Zoologici 58(4):695720.
Bocak, L., M. Bocakova, T. Hunt, and A. P. Vogler. 2008. Multiple ancient origins of neoteny in Lycidae
(Coleoptera): consequences for ecology and macroevolution. Proceedings of the Royal Society B,
275, 20152023.
Bocak, L., and K. Matsuda. 2003. Review of the immature stages of the family Lycidae (Insecta:
Coleoptera). Journal of Natural History 37:14631507.
Bocakova, M. 2001. Revision and phylogenetic analysis of the subfamily Platerodinae (Coleoptera,
Lycidae). European Journal of Entomology 98:5385.
Bocakova, M. 2003. Revision of the tribe Calopterini (Coleoptera, Lycidae). Studies on Neotropical
Fauna and Environment 38(3):207234.
Bocakova, M. 2005. Phylogeny and classification of the tribe Calopterini (Coleoptera, Lycidae). Insect
Systematics and Evolution 35(4):347447.
Bocakova, M., B. Baciakova, and E. A. Nascimento. 2012. Revision of the genus Falsocaenia (Coleoptera:
Lycidae), Zootaxa 3478:282296.
Bocakova, M., and E. A. Nascimento. 2013. A new species of Brasilycus (Coleoptera: Lycidae) from Peru.
Zootaxa 3637(5):597599.
Kazantsev, S. V. 2011. New and little known taxa of Platerotini, with a note on biogeography of the tribe
(Coleoptera: Lycidae). Russian Entomological Journal 20(2):151187.
Kleine, R. 1933. Lycidae. 145 pp. In Junk, W., and Schenkling, S. (eds.), Coleopterorum Catalogus, Pars
128, W. Junk, Berlin.
Kleine, R. 1941. Lycidae (Col.). In Titschack E. (editor), Beitrage zur Fauna Perus, Band 1. Gustav Fisher,
Jena, 295306. [Reprinted as Kleine, R. (1951) Lycidae (Col.). In Titschack E (ed.), Beitrage zur
Fauna Perus. Band 2. Gustav Fischer, Jena, 283294].
Kleine, R. 1942. Neue Lyciden des aethiopischen und neotropischen Faunengebietes (Col.). Mitteilungen
der Muenchner Entomologischen Gesellschaft 32:149162.
Nascimento, E. A., and M. Bocakova. 2009. A revision of the genus Lycomorphon (Coleoptera: Lycidae).
Zootaxa 2132:4052.
Lampyridae
Author(s): M. A. Branham

terms_of_use.
critical research.
88(2), 2015, pp. 248250
Beetles (Coleoptera) of Peru: A Survey of the Families. Lampyridae

M. A. BRANHAM
Department of Entomology and Nematology, University of Florida,
Gainesville, Florida 326110620, USA
e-mail: marcbran@ufl.edu

Recognition: The adults have the head completely or partially concealed from
above by pronotum. Antennae filiform to serrate, bipectinate, flabellate, biflabellate,
or rarely capitate. Mandibles are falcate, without a mola. The trochanter is obliquely
joined to the femur. The tibial spurs are usually indistinct or absent. The claws are
paired and simple or toothed or bifid, without setae near base. Light organs
sometimes present on the adults of one or both sexes and are usually located on
ventrites 5, 6, and 7, which correspond to segments VI, VII, and VIII (Branham,
2010). Bioluminescent signals in adults are used to locate mates. Non-luminous adults
often possess large non-filiform antennae and use pheromones to locate mates.
Habitat: Lampyrids are found in a diverse array of habitats and across most
elevations. While higher species diversity usually corresponds to regions with high
humidity, lampyrids are also known from arid regions. All lampyrids are predacious
as larvae, often specializing on either snails or earthworms. While the adults of most
species are not known to feed, some evidence exists for adults opportunistically
feeding on nectar. It is well known that the adult females of the Photuris
pennsylvanica-versicolor species group in North America practice aggressive mimicry.
These femmes fatales lure in and feed on heterospecific males by mimicking the
flashed responses of heterospecific females (Barber, 1951; Lloyd, 1965). While
aggressive mimicry is not known to occur in Peruvian Photuris species, Viviani (1996)
,http://entnemdept.ufl.edu/Lloyd/firefly/ffcomp1-2.pdf . reported it in Bicellonycha
ornaticollis (Blanchard) in Brazil. Bicellonycha ornaticollis is known to occur in Peru.
Notes: The following checklist of Peruvian Lampyridae was derived from a survey
of the Peruvian fauna by Kirsch (1873, 1874), a checklist by Blackwelder (1945), the
catalog of Lampyridae by McDermott (1966), and specimens housed in the USNM
collection. The author is not aware of any regional works, checklists and
identification keys to the Peruvian fauna of Lampyridae other than those listed
above. The Peruvian lampyrid fauna is undoubtedly under sampled as the
Neotropics is probably the most species diverse region for this family.
Checklist:
Amydetinae
Amydetes vigorsi Westwood, 1830 Psilocladus atromaculatus Pic, 1930
Dodacles interfusa E. Olivier, 1912 Psilocladus haemorrhoidalis E. Olivier, 1909
Dryptelytra cayennensis Laporte, 1833 Psilocladus peruvianus (Kirsch, 1874)
Dryptelytra longipennis Pic, 1927 Psilocladus pici E. Olivier, 1907
Ethra conserta (E. Olivier, 1909) Vesta cincticollis Blanchard, 1846
Ethra testaceicollis (Pic, 1929) Vesta melanura (Laporte, 1833)
Psilocladus apicalis (Kirsch, 1865) Magnoculus jocosus (E. Olivier, 1909)
Psilocladus atricornis Pic, 1925 Magnoculus olivieri (Pic, 1930)

Lampyrinae
Aspisoma aelianum (Gorham, 1884) Lucidota reductitincta Pic, 1943
Aspisoma angustum Kirsch, 1873 Lucidota saepta E. Olivier, 1908
Aspisoma bisignatum (Motschulsky, 1854) Lucidota severa (E. Olivier, 1911)
Aspisoma grossum Erichson, 1847 Lucidota submaculata (Pic, 1940)
Aspisoma nitens (DeGeer, 1774) Lucidota viridescens McDermott, 1965
Aspisoma sinuaticolle Kirsch, 1873 Lucio dimidiatum E. Olivier, 1885
Calyptocephalus opimus E. Olivier, 1909 Lucio diversum E. Olivier, 1912
Calyptocephalus picturatus E. Olivier, 1911 Lucio oblectata E. Olivier, 1911
Dilychnia guttula (Fabricius, 1801) Lucio obscurum E. Olivier, 1885
Lucidota albocincta (Pic, 1940) Macrolampis acicularis (E. Olivier, 1907)
Lucidota apicipennis (Pic, 1927) Macrolampis olivieri Pic, 1930
Lucidota aurantiaca (Pic, 1929) Petalacmis praeclarus E. Olivier, 1908
Lucidota baeri (Pic, 1927) Phaenolis infausta (E. Olivier, 1909)
Lucidota basalis Pic, 1930 Phaenolis olivieri Pic, 1924
Lucidota bicolor Kirsch, 1873 Photinoides mystrionophorus McDermott, 1963
Lucidota binotata (Pic, 1929) Photinus cinctellus (Motschulsky, 1854)
Lucidota consors E. Olivier, 1909 Photinus erichsoni Gemminger, 1870
Lucidota dissimilis E. Olivier, 1909 Photinus flaveolus E. Olivier, 1909
Lucidota eucera E. Olivier, 1907 Photinus fuscicornis Kirsch, 1873
Lucidota heterocera E. Olivier, 1910 Photinus interruptus (Erichson, 1847)
Lucidota immemor E. Olivier, 1909 Photinus lavali Pic, 1940
Lucidota interrupta Pic, 1927 Photinus maculicollis Kirsch, 1873
Lucidota laticornis Pic, 1943 Photinus marcapatanus Pic, 1941
Lucidota lecontei (Kirsch, 1873) Photinus minutissimus Pic, 1940
Lucidota luteohumeralis Pic, 1940 Photinus otuzeosus Pic, 1942
Lucidota macrescens E. Olivier, 1909 Photinus reductimarginalis Pic, 1941
Lucidota marginicollis Lucas, 1857 Photinus subinterruptus Pic, 1941
Lucidota ornate E. Olivier, 1909 Photinus tricolorithorax Pic, 1940
Lucidota pallipes (Pic, 1940) Photinus venustulus Erichson, 1847
Lucidota parallela (Pic, 1926) Photinus viduus Erichson, 1847
Lucidota peruviana (Pic, 1927) Platylampis explanata Pic, 1943
Lucidota probata E. Olivier, 1909 Platylampis madresa Pic, 1943
Photurinae
Bicellonycha ornaticollis (Blanchard, 1846) Photuris signatipennis E. Olivier, 1911
Bicellonycha rospigliosi Brethes, 1920 Photuris unicolor E. Olivier, 1909
Photuris apicalis Pic, 1926 Photuris unifasciata Pic, 1932
Photuris cincticollis Erichson, 1847 Pyrogaster testaceithorax Pic, 1928
Photuris inapicalis Pic, 1926
Acknowledgements
I acknowledge NSF-EPSCoR #66928 (PI: CS Chaboo) for supporting the Beetles
of Peru project and the University of Kansas Department of Ecology and
publication. I thank Caroline Chaboo for inviting me to collaborate on the Beetles of
Peru project and two anonymous reviewers for their careful reading of the manuscript.
Literature Cited
Barber, H. P. 1951. North American fireflies of the genus Photuris. Smithsonian Institutions
Miscellaneous Collections 117:158.
Blackwelder, R. E. 1945. Checklist of the coleopterous insects of Mexico, Central America, the West Indies,
and South America, part 3. Bulletin of the United States National Museum 185:343550.
Branham, M. A. 2010. Lampyridae. In Beutel, R. G., and R. A. B. Leschen (eds.), Handbuch der
Zoologie, Band IV Arthropoda: Insecta, Teilband 39, Evolution and Systematics, pp. 141147.
Walter de Gruyter, Berlin; xiii+786 pp.
Kirsch, T. F. W. 1874. Beitrage zur Kenntniss der Peruanischen Kaferfuana auf Dr. Abendroths
Sammlungen basirt. Berliner Entomologische Zeitschrift1873 (1874), 17:338418 (Zweites Stuck.)
Lloyd, J. E. 1965. Aggressive mimicry in Photuris: firefly femmes fatales. Science 149:653654.
McDermott, F. A. 1966. Lampyridae. In Steel, W. O. (ed.), Coleopterorum Catalogus Supplementa. Pars
9 (editio secunda). W. Junk, SGravenhage; 149 pp.
Viviani, V. 1996. Occurrence of aggressive mimicry in a Brazilian Bicellonychia firefly (Lampyridae:
Photurinae). http://entnemdept.ufl.edu/Lloyd/firefly/ffcomp1-2.pdf (Last downloaded 3 September
2014).
Cucujidae, Laemophloeidae, Silvanidae, Passandridae
(Cucujoidea)
Author(s): Michael C. Thomas and Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 251257

Cucujidae, Laemophloeidae, Silvanidae, Passandridae (Cucujoidea)
MICHAEL C. THOMAS1,3 AND CAROLINE S. CHABOO2
Recognition: The cucujid group of lower cucujoid families was for a long time
considered to be one large family, composed of the four following families, plus
several tenebrionoid genera (e.g., Hemipeplus of the Mycteridae and Inopeplus of the
Salpingidae) that share the dorsoventrally flattened body form. By the mid-20th
Century the composite nature of the family was beginning to be widely recognized
and by the end of the century the dismantling of it had been completed, although the
position of the Old World family Propalticidae is still under discussion.
Most members of these four families share a dorsoventrally compressed body
form, feed on fungi, and are associated with a substrate of dead plant material (bark,
wood, dead leaves, leaf litter). One family, Passandridae, has secondarily evolved to
be larval ectoparasitoids of other wood-inhabiting insects, and several genera of that
family and of the Laemophloeidae have evolved a subcylindrical body form and are
found in the galleries of wood-boring Coleoptera (Thomas, 1993).
Cucujidae Latreille, 1802

Diversity in Peru: 1 genus, 3 species. The world fauna consists of 57 species in 4
genera; 1 genus, 3 species species are known from South America.
Recognition: These are commonly called flat bark beetles because of the
dorsoventral compression of the adult body (Thomas and Leschen, 2010b). In addition
to the greatly flattened body, adult South American cucujids are characterized by their
bright color (orange and black, or red and metallic blue); possession of rather short
antennae without a distinct club; head larger than pronotum with prominent temples;
large, prominent mandibles (very large and bizarre in male Palaestes Perty, 1830);
small, transverse pronotum with irregular lateral margin and sharply, anteriorly
projecting front angles; open procoxal cavities with exposed trochantin; elytra with or
without stria; tarsal formula 5-5-4 in males, 5-5-5 in females; male genitalia curved,
without a flagellum, and resting on its side in abdomen. Length 1015 mm.
Habitat: Adults and larvae live under bark, where they appear to be predaceous.
Sharp (1899: 505) wrote of Palaestes: Mr. Champion informs me that these insects
are chiefly found between the thin crevices of freshly split sappy timber, and they are
often to be seen on the wing in forest clearings.
Notes: The following list of Cucujidae of Peru is derived from Blackwelder (1945).
Blackwelder (1945) listed Peru for the distribution of Palaestes nigriceps
Waterhouse, 1880, but this was a mistake since the type locality is cited as
Chiguinda, which is in Ecuador.
1
Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services,
P.O. Box 147100, Gainesville, Florida 32614-7100, USA, e-mail: michael.thomas@freshfromflorida.com
2
Lawrence, Kansas, 66045, USA; E-mail: cschaboo@ku.edu
3
Accepted 5 January 2015; Revised 5 January 2015
There have not been any previous faunal studies of the Peruvian cucujid fauna,
and only Palaestes tenuicornis Waterhouse, 1880 has been recorded from Peru.
Additionally, an undescribed species apparently representing a new genus is known
from Peru and as of this writing was being studied by John Marris and Adam
Slipinski (pers. commun.). The two species below have been recorded from South
America, but not yet from Peru.
Checklist:
Palaestes freyreissi (Heyden, 1827)
Palaestes nigriceps Waterhouse, 1880
Laemophloeidae Ganglbauer, 1899

Diversity in Peru: 4 genera, 6 species. About 470 species are known worldwide;
about 75 species in 10 genera from South America.
Recognition: These are very small beetles with the adults usually being
dorsoventrally compressed and with a characteristic system of submarginal ridges
and grooves on the head and pronotum and longitudinal impressed areas (cells)
on the elytra (Thomas and Leschen, 2010c). Head prognathous with usually
prominent mandibles (the head is prolonged anteriorly to form a distinct rostrum
in several Neotropical genera [i.e., Rhinomalus Gemminger and Harold, 1870;
Rhinophloeus Sharp, 1899; Metaxyphloeus Thomas, 1984]); antennae variable,
short with a distinct club to very long and filiform, in some genera the male scape
may be grotesquely modified; head bordered laterally with a longitudinal groove
or ridge which often continues around to form a posterior border; pronotum
usually narrower posteriorly, bordered sublaterally with longitudinal grooves or
ridges, the lateral margin may be explanate and irregular in a few genera, procoxal
cavities open to closed; elytra often with humeral carina and 13 longitudinal
impressed cells; tarsal formula usually 5-5-4 in males; 5-5-5 in females; male
genitalia inverted and with parameres fused to basal piece. Length 1.05.0 mm,
usually 1.52.5 mm.
Habitat: Larvae and adults of most species feed on ascomycete and related fungi
and are typically found under bark. A few species that are pests of stored grains
(Cryptolestes ferrugineus (Stephens, 1831) and C. pusillus (Schonherr, 1817) are
nearly worldwide in distribution and will probably be found in stored products in
Peru. Specimens may be found under bark of dead wood of fairly sound logs. Soft,
wet, rotten logs rarely yield laemophloeids. As the adults are strongly attracted to
light at night, ultraviolet light traps often catch large numbers of laemophloeids,
especially when operated in areas with dead wood nearby. The samples should be
examined carefully in alcohol under a microscope. Laemophloeids may be collected
at light sheets but are so small they are easily overlooked or disturbed by the arrival
of larger insects. Occasionally, they may be taken by beating dead branches over
a beating sheet, or by Berlesing litter. Several genera (e.g., Dysmerus Casey, 1884;
Narthecius LeConte, 1861; Leptophloeus Casey, 1916) are subcylindrical in cross-
section and typically are found in the galleries of scolytine curculionids, where they
may be predaceous. Members of these genera are rather poorly represented in
collections. Of the older literature, Kessel (1926), and Grouvelle (1896) provide
useful information and should be consulted.
Notes: The following checklist was developed from Thomas (1984, 1988, 2009,
2012, 2013, 2014). There have not been any previous faunal studies of the Peruvian
laemophloeid fauna, and only the following six species have been recorded from
Peru.
Checklist:
Cryptolestes ampiyacus Thomas, 1988 Placonotus pallentipennis (Grouvelle, 1876)
Dysmerus caseyi (Grouvelle, 1898) Laemophloeus corporeflavus Thomas, 2014
Dysmerus skelleyi Thomas, 2009 Laemophloeus incisus Sharp, 1899
The 51 described species below have been recorded from South America, but not yet
from Peru:
Cryptolestes calabozus Thomas, 1988 Laemophloeus megacephalus Grouvelle, 1876
Cryptolestes capillulus Thomas, 1988 Laemophloeus planaclavatus Thomas, 2014
Cryptolestes obesus Thomas, 2003 Laemophloeus sexarticulatus Kessel, 1926
Cryptolestes robinclarkei Thomas, 2004 Laemophloeus suturalis Reitter, 1876
Cryptolestes spatulifer Thomas, 1988 Laemophloeus taurus Thomas, 2014
Cryptolestes spectabilis Thomas, 1988 Metaxyphloeus germaini (Grouvelle, 1896)
Cryptolestes trinidadensis Thomas, 1988 Metaxyphloeus zeus Thomas, 1984
Cryptolestes uncicornis (Reitter, 1876) Odontophloeus crybetes Thomas, 1984
Dysmerus boliviensis Thomas, 2009 Odontophloeus kesseli (Hetschko, 1928)
Dysmerus curvicornis Thomas, 2009 Odontophloeus quadridentatus (Champion, 1913)
Phloeolaemus curtus (Grouvelle, 1876)
Dysmerus genaspinosus Thomas, 2009
Phloeolaemus impressus (Grouvelle, 186)
Dysmerus impolitus Thomas, 2009
Phloeolaemus quinquearticulatus (Grouvelle, 1896)
Dysmerus monstrosus Thomas, 2009
Phloeolaemus reitteri (Grouvelle, 1877)
Dysmerus politus Thomas, 2009
Placonotus politissimus (Wollaston, 1867)
Dysmerus rondoniensis Thomas, 2009
Placonotus pseudomodestus Thomas, 1984
Dysmerus symphilus Thomas, 2009 Rhabdophloeus constrictus Kessel, 1926
Dysmerus trinidadensis Thomas, 2009 Rhabdophloeus difficilis Kessel, 1926
Laemophloeus buenavista Thomas, 2013 Rhabdophloeus procul Kessel, 1926
Laemophloeus capitisculptus Thomas, 2014 Rhabdophloeus strictus Kessel, 1926
Laemophloeus concinnus Thomas, 2013 Rhinomalus fulvicollis Grouvelle, 1896
Laemophloeus dozieri Thomas, 2014 Rhinomalus ruficollis Grouvelle, 1896
Laemophloeus germaini Grouvelle, 1896 Rhinophloeus elegans (Grouvelle, 1896)
Laemophloeus lecontei Grouvelle, 1876 Rhinophloeus facetus (Grouvelle, 1896)
Laemophloeus macrognathus Reitter, 1876 Rhinophloeus productus (Grouvelle, 1876)
Laemophloeus mathani Grouvelle, 1889 Rhinophloeus salpingoides (Grouvelle, 1876)
Another 22 species of uncertain generic placement also have been described from
South America. Several other genera (Charaphloeus Casey, 1916; Lathropus
Erichsom, 1845; Lepidophloeus Thomas, 1984; Leptophloeus Casey, 1916; Narthecius
LeConte, 1861; Parandrita LeConte and Horn, 1880) are known from undescribed
South American species or have distributions suggesting they occur there.
Passandridae Erichson, 1845

Diversity in Peru: 2 genera, 2 species. Worldwide, there are 9 genera and 104
species; 5 genera and 25 species have been recorded from South America.
Recognition: The adults are small to medium-sized and dorsoventrally compressed
or subcylindrical, with thick, moniliform antennae and deep longitudinal grooves on
the elytra. This family has the common name parasitic flat bark beetles because the
known larvae are ectoparasitoides of the immature stages of other beetles and some
Hymenoptera (Dimmock, 1884; Fiske, 1905). Head prognathous with prominent

mandibles, bordered laterally and sometimes basally with a deep groove; pronotum
narrowed basally, anterior coxal cavities open; elytra usually with a humeral carina or
costa and longitudinal grooves; tarsi formula 5-5-5 in both sexes; male genitalia
inverted with articulated parameres and a flagellum. Size 420 mm.
Habitat: These beetles are found where their prey is found, and may be abundant
where Cerambycidae and other wood-boring beetles are found in large numbers.
They occur on trees and under bark, and are attracted to light, so that ultraviolet
light trapping may collect them in numbers.
Notes: The following list is derived from Burckhardt & Slipinski (2003).
Blackwelder (1946) did not report the family from Peru. There have not been any
previous faunal studies of the Peruvian passandrid fauna, and only the following two
species have been recorded from Peru.
Checklist:
Catogenus lebasii Guerin-Meneville, 1844
Taphroscelidia postica (Grouvelle, 1916)
The 25 described species below have been recorded from South America, but not yet
from Peru:
Catogenus acutangulus Reitter, 1878 Taphroscelidia atra (Grouvelle, 1916)
Catogenus asper Slipinski, 1989 Taphroscelidia atratula (Grouvelle, 1916)
Catogenus castaneus Perty, 1834 Taphroscelidia contorta Burckhardt & Slipinski, 1991
Catogenus decoratus Newman, 1839 Taphroscelidia dentata Burckhardt & Slipinski, 1991
Catogenus depressus Slipinski, 1989 Taphroscelidia filum (Reitter, 1876)
Catogenus gracilicornis Slipinski, 1989 Taphroscelidia gounellei (Grouvelle, 1916)
Catogenus grouvellei Slipinski, 1989 Taphroscelidia humeralis (Grouvelle, 1916)
Catogenus longicornis Grouvelle, 1874 Taphroscelidia nigra Burckhardt & Slipinski, 1991
Catogenus thomasi Slipinski, 1989 Taphroscelidia semicastanea (Reitter, 1876)
Passandra fasciata (Gray, 1832) Taphroscelidia sharpi (Grouvelle, 1916)
Passandrella visenda Grouvelle, 1916 Taphroscelidia tenuissima (Reitter, 1876)
Scalidiopsis youngi Burckhardt & Slipinski, 1991
Silvanidae Kirby, 1837

Recognition: These are small, brown, pubescent, flattened beetles, often with
anterolateral angles of the pronotum toothed or lobed or the lateral margins dentate
or denticulate (Thomas and Leschen, 2010a). The antenna may be elongate and
filiform, without a conspicuous club, or shorter with a distinct club. Head
prognathous, mandibles usually not prominent; eyes large, often with a well-marked
temple; pronotum narrowed basally, anterior coxal cavities broadly closed in almost
all South American genera; elytral striate, without humeral carinae; tarsal formula
5-5-5 in both sexes; male genitalia inverted or not, parameres articulated or not. Size
1.513.0 mm, mostly 25 mm.
The taxonomy of the Neotropical members of this family has been long neglected.
Species of the largest Neotropical genera (Telephanus Erichson, 1845; Ahasverus
Gozis, 1881; Nausibius Redtenbacher, 1858) are virtually unidentifiable with the
existing literature. Halstead is revising the latter two genera, both of which may
double or triple in numbers of species (D.G.H. Halstead, pers. commun.).
Habitat: Adults and larvae appear to eat fungus, but frequently they are not found
under bark. The tribe Telephanini, which contains the bulk of the Neotropical
species, are usually found on pendant, withered leaves, often those of Musaceae and
Heliconiaceae. Species of Coccidotrophus Schwarz and Barber, 1921 and Eunausibius
Grouvelle, 1912 tend mealybugs (Hemiptera: Pseudococcidae) in the leaf petioles of
the ant plant (Tachygalia sp.) and feed on honeydew (Wheeler, 1921). Members of
the genus Nepharis are ant inquilines in Australia (Lea, 1904), and it would not be
surprising to find social insect inquilines in South America; there are several
undescribed silvanines whose anatomical modifications suggest such a lifestyle. At
least a few silvanines seem to be facultative predators. Most silvanines are attracted
to light, and ultraviolet light trapping is an efficient method to survey for them.
Telephanines may often be collected in numbers by beating branches with masses of
dead pendant leaves.
Several species of Silvanidae are notorious stored products pests, especially two
species of the Old World genus Oryzaephilus Ganglbauer, 1899, O. surinamensis
(Linnaeus, 1758), and O. mercator (Fauvel, 1889). Neither has been recorded from
Peru but are certain to be found there. Other silvanid stored products pests include
Ahasverus advena (Waltl, 1834), Nausibius clavicornis (Kugelann, 1858), and
Cathartus quadricollis (Guerin-Meneville, 1854), all of which are of New World
origin. Eight species of Old World silvanids are known to be established in South
America: Cryptamorpha desjardinsi (Guerin-Meneville, 1844), Monanus concinnulus
(Walker, 1858), Monanus sp., Psammoecus trimaculatus Motschulsky, 1858,
Silvanoprus scuticollis (Walker, 1859), S. frater (Grouvelle, 1904), Silvanus difficilis
Halstead, 1973, and S. proximus Grouvelle, 1904. Most are of Asian origin. None is
known to be of economic importance.
There have not been any previous faunal studies of the Peruvian silvanid fauna,
and only one species has been recorded from Peru.
Checklist:
Monanus concinnulus (Walker, 1858)
Note: Halstead (1993) documented only one species for Peruan exotic species
from a Lonchocarpus root (the source of cube resin; Fabaceae).
The 79 species below have been recorded from South America, but not yet from
Peru. Four of these genera (Australohyliota Thomas, 2004; Australophanus Thomas,
2008; Brontoliota Thomas, 2004; Microhyliota Thomas, 2004) are probably restricted
to the Chilean fauna.
Ahasverus advena (Waltl, 1834) Coccidotrophus cordiae Barber, 1928

Ahasverus cryptophagoides (Reitter, 1878) Coccidotrophus socialis Schwarz and Barber, 1921
Ahasverus excisus (Reitter, 1876) Cryptamorpha desjardinsii (Guerin-Meneville, 1844)
Ahasverus nausibioides Grouvelle, 1912 Eunausibius elongatus (Grouvelle, 1896)
Ahasverus subopacus Grouvelle, 1912 Eunausibius lophius Parsons, 1974
Australohyliota chilensis (Blanchard, 1851) Eunausibius salutaris Parsons, 1974
Australophanus redtenbacheri (Reitter, 1876) Eunausibius tenebrionoides (Grouvelle, 1896)
Brontoliota lawrencei Thomas, 2010 Eunausibius wheeleri Schwarz and Barber, 1921
Cathartosilvanus aitkenae Halstead, 1993 Euplatamus buqueti (Grouvelle, 1877)
Cathartosilvanus opaculus (LeConte, 1854) Euplatamus castaneus (Grouvelle, 1881)
Cathartosilvanus vulgaris (Grouvelle, 1878) Euplatamus deyrollei (Grouvelle, 1876)
Cathartus quadricollis (Guerin-Meneville, 1854) Euplatamus humeralis (Reitter, 1877)
Euplatamus richteri (Reitter, 1878) Telephanus dubitalis Grouvelle, 1889

Euplatamus schaumi (Grouvelle, 1876) Telephanus dubius Grouvelle, 1880
Microhyliota integricollis (Fairmaire, 1860) Telephanus elongatus Grouvelle, 1889
Monanus (Monanus) sp. Telephanus fallax Grouvelle, 1880
Nausibius clavicornis (Kugelann, 1858) Telephanus haroldi (Schauffuss, 1876)
Nausibius gigas Grouvelle, 1896 Telephanus humerosus Reitter, 1874
Nausibius ingens Grouvelle, 1896 Telephanus insignis Grouvelle, 1880
Nausibius sahlbergi Grouvelle, 1896 Telephanus melanchlorus Nevermann, 1931
Nausibius sinuatus Grouvelle, 1896 Telephanus melanocephalus Grouvelle, 1889
Nausibius wagneri Grouvelle, 1912 Telephanus micans Grouvelle, 1880
Oryzaephilus mercator (Fauvel, 1889) Telephanus niger Reitter, 1874
Oryzaephilus surinamensis (Linnaeus, 1758) Telephanus obscurus Grouvelle, 1881
Psammoecus trimaculatus Motschulsky, 1858 Telephanus ornatus Reitter, 1874
Silvaninus nitidus (Grouvelle, 1889) Telephanus ovalis Grouvelle, 1889
Silvanoprus frater (Grouvelle, 1904) Telephanus paradoxus Reitter, 1874
Silvanoprus scuticollis (Walker, 1859) Telephanus parallelus Grouvelle, 1889
Silvanops angulicollis (Reitter, 1878) Telephanus pilicornis Reitter, 1874
Silvanops columbinus (Grouvelle, 1881) Telephanus procerulus Reitter, 1874
Silvanosoma striatum Brethes, 1922 Telephanus pulchellus Reitter, 1878
Silvanus difficilis Halstead, 1973 Telephanus reductus Schauffuss, 1876
Silvanus proximus Grouvelle, 1904 Telephanus sahlbergi Grouvelle, 1896
Synobius lobatus (Grouvelle, 1896) Telephanus schwarzi Nevermann, 1931
Telephanus acuminatus Grouvelle, 1877 Telephanus similis Grouvelle, 1889
Telephanus argentatus Reitter, 1874 Telephanus subpubescens Grouvelle, 1899
Telephanus bruchi Grouvelle, 1904 Telephanus terminatus Grouvelle, 1880
Telephanus decoratus Grouvelle, 1880 Telephanus wahlbergi Grouvelle, 1896
Telephanus dilutus Reitter, 1874
Acknowledgements
Literature Cited
Blackwelder, R. E. 1946. Checklist of the coleopterous insects of Mexico, Central America, The West
Indies, and South America. Part 4. U.S. National Museum, Bulletin 185. Smithsonian Institution,
Washington, D.C., 763 pp.
Burckhardt, D., and S. A. Slipinski. 2003. Phylogeny and taxonomy of the world Passandridae. In
Cuccodoro, G., and R. A. B. Leschen (eds.). Systematics of Coleoptera: papers celebrating the
retirement of Ivan Lobl. Memoirs on Entomology International, volume 17:753883. Gainesville,
Florida: Associated Publishers, 955 pp.
Dimmock, G. 1884. Notes on Catogenus rufus. Psyche 1884:341342.
Fiske, W. F. 1905. Catogenus rufus, a coleopterous parasite. Proceedings of the Entomological Society of
Washington 7:9092.
Grouvelle, A. 1896. Nitidulides, Colydiides, Cucujides et Parnides recoltes par M. E. Gounelle au Bresil et
autres Clavicornes nouveaux dAmerique. Annales de la Societe Entomologique de France 65:
187209.
Halstead, D. G. H. 1993. Keys for the identification of beetles associated with stored productsII.
Laemophloeidae, Passandridae and Silvanidae. Journal of Stored Products Research 29:99197.
Kessel, F. 1926. Synopse geral do genero Laemophloeus Cast. (Col.) com a descripcao de algumas novas
especies Sul-Americanas. Archivos do Museu Nacional, Rio de Janeiro. 26:5993.
Sharp, D. 1899. Cucujidae. In Biologia Centrali Americana, Col. 2(1):449563.
Thomas, M. C., and R. A. B. Leschen. 2010a. Silvanidae Kirby, 1837. In Leschen, R. A. B., R. G. Beutel,
and J. F. Lawrence (eds.). Coleoptera, Beetles. Vol. 2: Morphology and Systematics (Elateroidea,
Bostrichiformia, Cucujiformia partim), pp. 346350. Handbook of Zoology. Walter de Gruyter,

Berlin.
Thomas, M. C., and R. A. B. Leschen. 2010b. Cucujidae Latreille, 1802. In Leschen, R. A. B., R. G.
Beutel, and J. F. Lawrence (eds.). Coleoptera, Beetles. Vol. 2: Morphology and Systematics
(Elateroidea, Bostrichiformia, Cucujiformia partim), pp. 350354. Handbook of Zoology. Walter
de Gruyter, Berlin.
Thomas, M. C., and R. A. B. Leschen. 2010c. Laemophloeidae, Ganglbauer, 1899. In Leschen, R. A. B.,
R. G. Beutel, and J. F. Lawrence (eds.). Coleoptera, Beetles. Vol. 2: Morphology and Systematics
(Elateroidea, Bostrichiformia, Cucujiformia partim), pp. 376380. Handbook of Zoology. Walter
de Gruyter, Berlin.
Thomas, M. C. 1984. A revision of the New World species of Placonotus Macleay (Coleoptera: Cucujidae:
Laemophloeinae). Occasional Papers of the Florida State Collection of Arthropods 3:ivii, 128.
Thomas, M. C. 1988. A revision of the New World species of Cryptolestes Ganglbauer (Coleoptera:
Cucujidae: Laemophloeinae). Insecta Mundi 2:4365.
Thomas, M. C. 1993. The flat bark beetles of Florida (Laemophloeidae, Passandridae, Silvanidae).
Arthropods of Florida and Neighboring Land Areas 15:iviii and 193.
Thomas, M. C. 2009. A review of the genus Dysmerus Casey (Coleoptera: Laemophloeidae). Insecta
Mundi 0074:130.
Thomas, M. C. 2012. A preliminary checklist of the flat bark beetles of the world (Cucujidae (s. str.),
Laemophloeidae, Passandridae, Silvanida). Available online: ,http://www.fsca-dpi.org/Coleoptera/
Mike/chklist.htm.. Last accessed: 8 June 2014.
Thomas, M. C. 2013. A review of New World Laemophloeus Dejean (Coleoptera: Laemophloeidae): 1.
Species with antennal club of more than three antennomeres. Insecta Mundi 0294:123.
Thomas, M. C. 2014. A review of New World Laemophloeus Dejean (Coleoptera: Laemophloeidae): 2.
Neotropical species with antennal club of three antennomeres. Insecta Mundi 0363:138.
Wheeler, W. M. 1921. A study of some social beetles in British Guiana and of their relations to the ant-
plant Tachigalia. Zoologica (New York) 3:35126.
Lymexylidae Fleming, 1821
Author(s): Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 258259

Lymexylidae Fleming, 1821
CAROLINE S. CHABOO
Division of Entomology, Natural History Museum and Department of Ecology and
Evolutionary Biology, 1501 Crestline Drive, Suite 140, University of Kansas,
Lawrence, Kansas 66049, USA
e-mail: cschaboo@ku.edu
Diversity in Peru. 2 subfamilies, 2 genera, 2 species.

Recognition. Both adults and larvae are elongate and narrow. Adults have the
elytra either very short with the hind wings and abdomen mostly exposed
(Atractocerinae) or the elytra nearly complete (Melittommatinae). The vertex has
an externally evident epicranial pit. Males have a complex multi-branched maxillary
palp organ originating from palpomere 3. Larvae have a rounded head, a dorsally
humped prothorax, and (in Peruvian subfamilies) abdominal tergite 9 modified,
sclerotized, and either bulbous with dense asperities or cylindrical and truncate with
large teeth (Wheeler, 1986).
Habitat. These wood-boring beetles are commonly called ship-timber beetles
because some species are pests in wooden structures (e.g., houses, ships). The larvae
bore into wood (living or dead), where they consume fungi (e.g., Ascoidea spp.,
Endomyces hylecoeti) brought in via fungus-pouches in the female genitalia. The
larva of Atractocerus brasiliensis Lepeletier & Audinet-Serville, 1825 and the larva of
a Melittomma species are described (Costa et al., 1988; Casari and Possidonio
Teixeira, 2011). I have collected specimens in Peru at both Malaise and UV light
traps.
Notes. The following checklist of Peruvian Lymexylidae was derived from Vaurie
(1956) and Wheeler (1986). Blackwelder (1945) and lists only Melittommopsis
abdominale (Pic, 1936) for Peru. Wheeler (1986) provided a review of the family at
the genus level.
Checklist
Atractocerinae
Atractocerus brasiliensis Lepeletier & Audinet-Serville, 1825
Melittommatinae
Melittommopsis abdominale (Pic, 1936)
Acknowledgements
Evolutionary Biology-General Research Fund (PI: CS Chaboo). I also thank
Matthew Gimmel for his help.
Accepted 23 July 2014; Revised 25 August 2015
Literature Cited
Blackwelder, R. E. 1945. Checklist of the coleopterous insects of Mexico, Central America, the West Indies,
and South America. Parts 1 through 6. United States National Museum Bulletin 185(part 3): iiiiv,
ixii, 343550.
Casari, S. A. and E. Possidonia Teixeira. 2011. Larva of Atractocerus brasiliensis (Lepeletier & Audinet-
Serville, 1825) (Lymexylidae, Atractocerinae). Papeis Avulsos de Zoologia 51(12):197205.
Vaurie, P. 1956. Atractocerus brasiliensis in Cuba. The Coleopterists Bulletin 10:86.
Wheeler, Q. D. 1986. Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the
American Museum of Natural History 183:113210.
Beetles (Coleoptera) of Peru: A Survey of the Families. Cleridae
Latreille, 1802
Author(s): Alan Burke and Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 260266

Cleridae Latreille, 1802
ALAN BURKE1 AND CAROLINE S. CHABOO2,3
Diversity: 7 subfamilies, 27 genera, 92 species.

Introduction: The most recent catalog of the Cleridae enlists 3366 species (Corporaal,
1950). This number is certainly outdated, as several species have been described and
catalogued since. Until 1960, many author extensively worked with the clerid fauna of
South America, including Peru. Among these, we can mention authors such as Johan C.
Fabricius, Maurice Pic, Sigmund Schenkling, Edward A. Chapin and Johann B.
Corporaal. Later on, surveys of the Cleridae in South America diminished considerably.
It was not until recently that, people like Weston Opitz and Jaime Solervicens resumed
work in this region. Thanks to the efforts of these scientists, it is possible to assert that,
currently, the clerid fauna of Peru records 92 species classified into 27 genera and 7
subfamilies. Much work remains to be done in this exceptionally diverse Andean
country, and many species are yet to be described and classified. This work has the sole
purpose of giving a brief description of what the clerids are, how they are classified, and
finally, provide a checklist of the species inhabiting the Peruvian territory.
General recognition: Members of the Cleridae, commonly referred to as checkered
beetles, comprise a group of beetles of moderate dimensions, ranging in size from less
than 5 mm to more than 30 mm. They are moderately broad to elongate, almost always
covered with erect and sub-erect setae. Coloration ranges from conspicuously bright to
dull or almost black. The eyes are of variable size, may be bulging laterally or not, slightly
to strongly emarginate. The antennae are greatly variable in shape and length. The elytra
almost always fully cover the abdomen, and rarely up to two abdominal segments may be
exposed. The tibiae are typically armed with two spurs. The tarsi possess pulvilli or tarsal
pads which are heavily vested. The abdomen has six visible segments, the last abdominal
segment of males may be conspicuously modified from that of females or not.
Diagnosis: Clerid members can be recognized by the presence of the following
combination of characters: possession of a postgular plate or postgular projection
(Opitz, 2010b), this projection is absent in remaining families within Cleroidea. The
lateral portion of the lacinia is enlarged, ranging in size from greatly expanded to
a lacinial folding. Finally, the protarsus is ventrally dilated, either in the form of
tarsal pulvilli or setiferous extensions. The size and number of tarsomeres bearing
pulvilli or setiferous extensions can vary among genera.
Habitat: Little is known about the biology and ecology of the Cleridae. As a general
rule, both larvae and adults are primarily predaceous of other insects; however, a small
number of species are anthophillic (feeding on pollen). The ecological preferences of
these beetles are markedly variable. Some species are associated with flowers and
grasses, among these, various member of the genus Trichodes Herbst; others are found
1
Department of Entomology, 123 Waters Hall, Kansas State University, Manhattan, Kansas, 66506
4004, USA, e-mail: burkea@ksu.edu
2
Division of Entomology, 1501 Crestline Drive, Suite 140, University of Kansas, Lawrence, Kansas,
66045, USA, e-mail: cschaboo@ku.edu
3
Accepted 28 August 2014; Revised 9 March 2015
feeding on the bark and foliage of various tree species, as in the genus Thanasimus
Latreille; many are found in the nests of Isoptera and Hymenoptera species, such as
Lecontella Wolcott and Chapin; and some are carrion visitors, like Korinetes Herbst.
One species, Necrobia rufipes (DeGeer), commonly known as the red-legged ham
beetle, is an important pest of various stored meat products.
Classification and distribution: In recent years, the higher classification of the
Cleridae has undergone considerable fluctuation. Presently, the most accepted
classification of the Cleridae encompasses 13 subfamilies (Crowson, 1964; Kolibac,
1992; Opitz, 2010b; Gunther et al., 2013) distributed in all continents, except the
Antarctica. From these, seven subfamilies are represented in Peru. Subfamily
diagnoses and habitat notes, together with checklists of the clerid species inhabiting
the Peruvian territory are detailed below.
Subfamily Clerinae Latreille, 1802

Diagnosis: Clerinae is the most speciose subfamily within the Cleridae. They can
be distinguished from other checkered beetles if they have the following combination
of characters: tarsal formula of 5-5-5; the fourth tarsomere is not reduced; the
protarsomeres are not compacted; the tarsal pulvilli are well developed; and the
procoxal cavities are almost always open, if closed, the prointercoxal process does
not reach the pronotal projections.
Habitat: Members of this subfamily have a worldwide distribution. Diversity of
genera and species is particularly high in the tropical and subtropical regions.
Clerinae larvae and adults are generally predators of insects infesting wood, most
notably on bark and ambrosia beetles (Scolytinae and Platypodinae); other species
are known to inhabit flowers and grasses, very likely preying on various insects
visiting those habitats; and few species feed on pollen. Adults of many species within
the group prey predominantly on the outer portion of the bark, whereas larvae feed
inside the tunnels and chambers constructed by bark and wood-boring beetles.
Distribution and species checklist: Currently, more than 1500 species classified in
109 genera have been described worldwide, but many taxa remain new to science.
Twenty nine species in six genera are found in Peru:
Checklist:
Aphelocerus immarginatus (Chevrolat, 1874) Enoclerus nodulifer Gorham, 1927
Axina fasciata Kirsch, 1873 Enoclerus peruvianus Pic, 1936
Ctenaxina brunnea (Schenkling, 1906) Enoclerus pusio (Schenkling, 1906)
Enoclerus anceps (Gorham, 1882) Enoclerus pusio ab. laetipes (Schenkling, 1915)
Enoclerus apicatus (Schenkling, 1906) Enoclerus rufimanus (Schenkling, 1900)
Enoclerus arrowi (Schenkling, 1907) Enoclerus rufofemoratus (Schenkling, 1915)
Enoclerus axillaris (Erichson, 1847) Enoclerus triplagiatus (Blanchard, 1843)
Enoclerus bellus (Schenkling, 1898) Enoclerus venator (Chevrolat, 1843)
Enoclerus cinctus (Schenkling, 1915) Perilypus acus Ekis, 1977
Enoclerus cinereopilosus (Blanchard, 1843) Perilypus crassus Ekis, 1977
Enoclerus deliciolus (Gorham, 1876) Priocera abdominalis (Blanchard, 1843)
Enoclerus faber (Chevrolat, 1874) Priocera cylindrica Thomas, 1940
Enoclerus fasciatus (Schenkling, 1900) Priocera femoralis Kirsch, 1865
Enoclerus faustus (Schenkling, 1900) Priocera miersiana White, 1849
Enoclerus lugubris (Erichson, 1847) Priocera quadrinotata Schenkling, 1900
Enoclerus mutabilis (Chevrolat, 1874) Priocera spinosa (Fabricius, 1801)
Subfamily Enopliinae Gistel, 1856

Diagnosis: Specimens in this subfamily can be reliably diagnosed based on the
following combination of characters: presence of a reduced fourth tarsomere;
absence of pronotal trichobothria (sensory setae arising from cupuliform base); and
the pronotal dorsolateral carina reaches the pronotal hem.
Habitat: Very little is known about the biology and ecology of this subfamily.
Various species in the subfamily are known to display aposematic coloration, others
are probably mimicking various lycid beetles. Some members are thought to be
anthophilic.
Distribution and species checklist: A small subfamily of checkered beetles with 66
described species distributed in 15 genera. Members of the Enopliinae are
cosmopolitan in distribution. The greatest diversity of species is found in the
tropical regions of Africa. Only one species is known from Peru.
Checklist:
Pyticara flavicollis Gorham, 1877
Subfamily Epiphloeinae Kuwert 1893

Diagnosis: Diagnostic characters to distinguish the Epiphloeinae from other
checkered beetles are the next: reduced fourth tarsomere, antennal insertion
conspicuously separated from eye emargination, two pairs of trichobotria located
on the pronotal disc, and the tarsal pulvillar formula can be 3-3-2 or 3-3-1.
Habitat: Larvae and adults of the Epiphloeinae are strongly associated with
woodland environments of temperate to tropical forest stands. Member of this
subfamily feed on a wide variety of wood-boring, bark and ambrosia beetles.
Various species can be brightly colored, very likely mimicking aposematic insects.
Larvae of these checkered beetles are found inside tunnels and chambers of
various beetle species associated with the decaying of forest stands, feeding on
immature stages. Adults are active consumers of other beetles inhabiting woody
environments.
Distribution and species checklist: Members of this subfamily are restricted to the
New World. The group is particularly species-rich in the tropical regions of the
Americas, although they can be found from Canada to Argentina. The subfamily is
composed of 267 described species classified in 25 genera. Presently, 29 species in
10 genera are found in Peru; nevertheless, many remain to be described. Various
species inhabiting surrounding countries are very likely found in the Peruvian
territory too.
Checklist:
Amboakis micula Opitz, 2006 Ichnea marginella (Klug, 1842)
Ellipotoma tenuiformis Spinola, 1844 Ichnea plumbea Gorham, 1877
Epiphloeus duodecimmaculatus Klug, 1842 Ichnea procera Schenkling, 1900
Ichnea aequinoctialis Spinola, 1844 Ichnea roseicollis Kuwert, 1894
Ichnea callanga Opitz, 2010 Iontoclerus humeralis (Klug, 1842)
Ichnea divisa (Chevrolat, 1843) Iontoclerus sericeus (Klug, 1842)
Ichnea frenata (Erichson, 1847) Madoniella cracentis Opitz, 2011
Ichnea gregata Opitz, 2010 Megaphloeus animosus (Wolcott, 1927)
Ichnea helvolicollis Corporaal, 1927 Megaphloeus mucoreus (Klug, 1842)
Ichnea incerta Gorham, 1877 Megaphloeus setulosus (Thomson, 1860)
Megaphloeus sexplagiatus (Kuwert, 1893) Plocamocera confrater Kuwert, 1893

Megaphloeus terzonatus (Gorham, 1877) Plocamocera sericella Spinola, 1844
Megaphloeus variegatus (Klug, 1842) Plocamocera sesquipedalis Opitz, 2004
Megaphloeus velutinus (Gorham, 1877) Pyticeroides arrogans Kuwert, 1894
Plocamocera castanea Opitz, 2004 Stegnoclava fumigata (Gorham, 1877)
Subfamily Korynetinae Laporte 1836

Diagnosis: Members of this subfamily can be distinguished from other checkered
beetles by the reduction of the fourth tarsomere; a fully developed pronotal
commissure, the pronotal commissure reaches the pronotal hem; a complete
dorsolateral ridge; and a reduced antennal club.
Habitat: Species in this subfamily are well known due to their relationship with the
decomposition of carcasses of various vertebrates. For instance, the cosmopolitan
Necorbia rufipes (DeGeer) has been studied extensively for its role in the
disintegration process of rotten flesh, feeding on various body parts, including
bones and skin. Adults and immature stages of this species are also know to be an
important pest of various smoked fish, dried meat and cheese products. After
reaching maturity, larvae will move from stored products or carcasses and develop
a pupal chamber where, after 1216 days, adults will emerge. Adults are also know
to feed on other saprophytic insects, such as blow flies, dermestid beetles and chees
flies. Biology of remaining genera is less known.
Distribution and checklist: The subfamily is composed of 76 species in 7 genera.
They have a worldwide distribution. The genus Necrobia Olivier is widely
distributed. Various genera of minor size are restricted to the south hemisphere,
with concentration of species in Oceania and South America.
Checklist:
Necrobia rufipes (DeGeer, 1775)
Subfamily Neorthopleurinae Opitz, 2009

Diagnosis: The diagnostic characters to identify members of this subfamily are:
a complete pronotal dorsolateral ridge, which fully surrounds the pronotum;
a reduced gula; a complete and bilobed gular process; the gular sutures are divergent;
an incomplete anterolateral pronotal extension; the pronotal commissure is well
developed; the dorsolateral pronotal carina does not meet the pronotal hem; and the
ovipositor is longer than the abdomen (Opitz, 2010b).
Habitat: Like many checkered beetles, members of this group, both adults and
larvae, prey in lignicolous environments. Enlarged ommatidia on various species
may serve as an indication of nocturnal activity. A number of individuals in the
genus Allochotes Westwood are known to mimic coccinellid beetles. The larveae of
many species are found inside tunnels of wood-boring beetles. Adults are known
to feed on various cerambycid, buprestid and curculionid species (Champlain,
1920).
Distribution and species checklist: Currently, 258 species distributed in 22 genera
have been described. These checkered beetles have a worldwide distribution.
Concentration of species occurs in tropical regions of Africa and Asia. Only two
species have been documented from Peru.
Checklist:
Neorthopleura tricolor Barr, 1976
Loedelia mexicana (Gahan, 1910)
Subfamily Peloniinae Opitz 2010

Diagnosis: Species in this subfamily are readily diagnosed if they possess a reduced
fourth tarsomere; pronotal trichobotria are absent; possession of an enlarged
antennal club, the club is at least as large as remaining antennomeres, but almost
always much longer than the length of preceding antennomeres combined; and the
dorsolateral carina always meets the pronotal hem at the posterolateral angles of the
pronotum.
Habitat: Some species of this subfamily display characteristic stout legs. Opitz
(2010b) indicates that, due to the condition of the legs, these beetles are not
particularly swift hunters. Various species in this group mimic aposematic species of
insects that have slow motion and are found inhabiting broad-leaved vegetation. A
number of species resemble members of the leaf beetle subfamily Galerucinae. The
reason of this similarity may be related to the fact that certain galerucine species
display aposematic colorations.
Distribution and species checklist: The subfamily is widely distributed with
remarkable abundance of species in Central and South America. Two hundred and
thirty four species within 31 genera are recognized. Twenty species classified in six
genera are found in Peru.
Checklist:
Apolopha eucharis Opitz, 1998 Cregya peruviana Pic, 1952
Apolopha reichei Spinola, 1844 Cregya sexnotata (Klug, 1842)
Apolopha suturalis (Klug, 1842) Cregya sexnotata var. quadrinotata (Schenkling,
Corinthiscus lividus Schenkling, 1906 1903)
Corinthiscus lobaticollis Lesne, 1909 Pelonium amoenum Guerin, 1838
Corinthiscus riveti Lesne, 1909 Pelonium lividum (Erichson, 1847)
Corinthiscus sexpunctatus Kirsch, 1873 Pelonium lobaticolle Lesne, 1909
Corinthiscus spectabilis Peracchi, 1960 Pelonium pilosum (Klug, 1842)
Cregya conformis (Chevrolat, 1876) Pelonium riveti Lesne, 1909
Cregya divisa (Gorham, 1903) Pelonium sexpunctatum Kirsch, 1873
Cregya flavomarginata (Chevrolat, 1874) Pelonium viridipenne (Kirby, 1818)
Cregya frontalis (Kuwert, 1894) Pilosirus brunoi Opitz, 1997
Cregya gemina (Schenkling, 1900) Platynoptera peruviana Pic, 1950
Cregya irrorata (Gorham, 1877) Platynoptera peruviana var. lineatithorax
Cregya klugi Gorham, 1949 Pic, 1956
Subfamily Tillinae Leach 1815

Diagnosis: The Tillinae can be readily identified from remaining checkered beetles
if they have the procoxal cavities closed posteriorly and interiorly, that means, the
intercoxal process joins the pronotal projections, and the procryptosternum is
complete. Additionally, the anterior portion of each metacoxal cavity bears a ridge
or carina.
Habitat: The Tillinae are generalist predators of various groups of insects.
A number species in the genus Cymatodera Gray have been observed feeding on
wood-boring beetles, solitary and gall wasps, the larvae of numerous lepidopterous
species, aphids, and scales; many of these insects are considered sanitary pests of
moderate to minor importance. Tillinids are commonly associated to lignicolous
environments. Coarsely faceted eyes in several species may be an indication of
nocturnal activity. A comprehensive review of the predatory activity of the genus
Lecontella Wolcott and Chapin is given by Mawdsley (2002). Rifkind (2006)
mentions that some Cymatodera species are capable of stridulating. This behavior
may be the result of an auditory Batesian mimetic strategy.
Distribution and checklist: After Clerinae, this is the second largest subfamily of
checkered beetles. The group has a global distribution. Species richness is found in
the sub-temperate to sub-tropical, thorny and scrub forests of North America, and
the tropical regions of Africa and Madagascar. Currently, approximately 585 species
distributed into 67 genera have been described.
Checklist:
Callotillus elegans (Erichson, 1847)
Note: The checklists given here derived from Corporaal (1942), Corporaal (1950),
Pic (1952, 1955, 1956), Peracchi (1960), Barr (1976a, b), Ekis (1977), Opitz (1997, 1998,
2004, 2005, 2006, 2007, 2008a, b, 2010a, 2011a, b) and Solervicens (2005). Corporaal
(1942) listed several unconfirmed genera of likely occurrence in Peru; nevertheless, for
the sake of accuracy these species are not listed here. For phylogenetic information
and a detailed analysis of the suprageneric classification of the Cleridae, please refer to
Crowson (1964), Kolibac (1992), Opitz (2010b), and Gunter et al. (2013).
Acknowledgements
Gimmel for help assembling species names.
Literature Cited
Barr, W. F. 1976a. Taxonomy of the new clerid genus Neorthopleura (Coleoptera). Melanderia 24:114.
Barr, W. F. 1976b. Descriptions and taxonomic notes of Enoclerus and some allied genera (Coleoptera:
Cleridae). Melanderia 24:1735.
Champlain, A. B. 1920. Part II. Notes of the seasonal history and biology of North American Cleridae.
Proceedings of the United States National Museum 57:575624.
Corporaal, J. B. 1942. Cleridae (Col.). Beitrage zur Fauna Perus 2(13):135147.
Corporaal, J. B. 1950. Cleridae. In Hincks, W. D. (ed.). Coleopterorum Catalogus Suplementa, Pars 23
(edition seconda), J. Junk; Gravenhage, 373 pp.
Crowson, R. A. 1964. A review of the classification of Cleridae (Coleoptera) with description of two new
genera of Peltidae and of several new larval types. Transaction of the Royal Entomological Society
of London 116(12):275327.
Ekis, G. 1977. Classification, phylogeny, and zoogeography of the genus Perilypus (Coleoptera: Cleridae).
Smithsonian Contributions to Zoology 227:1138.
Gunter, N. L., J. M. Leavengood, J. S. Bartlett, E. G. Chapman, and S. L. Cameron. 2013. A molecular
phylogeny of the checkered beetles and a description of Epiclininae a new subfamily (Coleoptera;
Cleridae). Systematic Entomology 38(3):626636.
Kolibac, J. 1992. Revision of Thanerocleridae n. stat. (Coleoptera: Cleroidea). Mitteilungen der Schweize-
rischen Entomologischen Gesellschaft 65:303340.
Mawdsley, J. R. 2002. Comparative ecology of the genus Lecontella Wolcott and Chapin (Coleoptera:
Cleridae: Tillinae), with notes on chemically defended species of the beetle family Cleridae.
Proceedings of the Entomological Society of Washington 104(1):164167.
Opitz, W. 1997. Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae).
Part I. The genera of Epiphloeinae. Insecta Mundi 11:5196.
Opitz, W. 1998. The classification and evolution of Apolopha Spinola (Coleoptera: Cleridae, Enopliinae).
The Coleopterists Bulletin 52:522.
Part II. The genera Chaetophloeus Opitz and Plocamocera Spinola. Bulletin of the American
Museum of Natural History 280:182.
Opitz, W. 2005. Classification, natural history, and evolution of the genus Aphelocerus Kirsch
(Coleoptera: Cleridae: Clerinae). Bulletin of the American Museum of Natural History 293:1128.
Part III. The genera Parvochaetus, n. gen., Amboakis, n. gen., and Ellipotoma Spinola. Insecta
Mundi 20:97164.
Opitz, W. 2007. Classification, natural history, and evolution of the Epiphloeinae (Coleoptera, Cleridae).
Part IV. The genera Pyticeroides Kuwert, 1894 and Diapromeces Opitz, 1997. Entomologica
Basiliensia et Collectionis Frey 29:77166.
Opitz, W. 2008a. Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae).
Part VI. The genera Epiphlaeus Spinola and Opitzius Barr. Annales Zoologici 38:134.
Opitz, W. 2008b. Classification, natural history, and evolution of Epiphloeinae (Coleoptera: Cleridae).
Part VII. The genera Hapsidopteris Opitz, Iontoclerus Opitz, Katamyurus Opitz, Megatrachys
Opitz, Opitzia Nemesio, Pennasolis Opitz, new genus, Pericales Opitz, new genus, Pteroferus Opitz,
new genus, and Turbophloeus Opitz, new genus. Zootaxa 1754:140.
Opitz, W. 2010a. Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae).
Part VIII. The genera Acanthocollum Opitz, Stegnoclava Opitz, and Ichnea Laporte. The
Coleopterists Society Monographs 9:165.
Opitz, W. 2010b. Classification, natural history, phylogeny, and subfamily composition of the Cleridae in
generic content of the subfamilies (Coleoptera: Cleridae). Entomologica Basiliensia et Collectionis
Frey 32:31128.
Opitz, W. 2011a. Classification, natural history, and evolution of the Epiphloeinae (Coleoptera, Cleridae).
Part IX. The genus Megaphloeus Opitz, 2010. Entomologica Basiliensia et Collectionis Frey
33:63132.
Opitz, W. 2011b. Classification, natural history, and evolution of the Epiphloeinae (Coleoptera, Cleridae).
Part X. The genus Madoniella Pic, 1935. Entomologica Basiliensia et Collectionis Frey 33:133248.
Peracchi, A. L. 1960. Contribucao ao estudo dos Cleridas neotropicais (Coleoptera, Cleridae). Revista
Brasileira Biologia 20:6368.
Pic, M. 1952. Coleopteres du Globe. LEchange, Revue Linneenne 68(527):14.
Pic, M. 1956. Coleopteres du Globe. LEchange, Revue Linneenne 72(543):14.
Rifkind, J. 2006. Stridulation in Cymatodera Gray, 1832 (Coleoptera: Cleridae). The Pan-Pacific
Entomologist 82(2):258261.
Solervicens, J. A. 2005. A prothoracic character that defines the subfamily Korynetinae (Coleoptera:
Cleridae). New Zealand Entomologist 28:4954.
Byrrhidae Latreille, 1804
Author(s): Paul J. Johnson

terms_of_use.
critical research.
88(2), 2015, pp. 267268

Byrrhidae Latreille, 1804
PAUL J. JOHNSON
Insect Biodiversity Lab, Box 2207A, 1010 Rotunda Drive, South Dakota State
University, Brookings, SD 57007
e-mail: paul.johnson@sdstate.edu
Diversity in Peru: 1 subfamily, 1 genus, 1 species.

Recognition: The adult beetles are immediately recognized by their very convex,
tight and compact body, strongly deflexed head, transverse front coxae, and
retractile antennae and legs. Tarsi are all pentamerous. Antennae are narrowly to
moderately clavate. Flight wings are reduced or lacking. They are commonly called
pill beetles, due to their compact appearance, and sometimes moss beetles because of
their associations with bryophytes.
Habitat: The known specimens of Peruvian byrrhid are from high elevations,
probably in mosses among grasses and broken rock. Lowland species of South
America remain unknown for Peru. Elsewhere, most species are bryophagous.
Notes: We report here the first record of the family Byrrhidae from Peru, based on
two specimens that were both collected at high elevations:
Specimen 1: PERU: JU. Carhuamayo, La Victoria 10u51959.50/75u569200 4384 m,
1516.iv.2012, I. Galindo y J. Grados [1 male, MUSM].
Specimen 2: PERU: Junin, 30.xi.78, R. Aloana/EEASC 7278/Peru [1, USNM].
There are no described byrrhids from Peru or elsewhere in the central or northern
Andes, or Amazonia. The most recent summary of six species representing five
genera of South American byrrhids is Blackwelder (1944), who repeated the taxa
listed by Dalla Torre (1911). Most reports of byrrhids in South America are from
forests of southern Chile and the Falkland Islands. The exception is Reichardt (1974,
1975) who reported Chaetophora striata (Pic) and C. monnei (Reichardt) from
southern Brazil and Uruguay. Based on above reports, it can be predicted that
presently unreported taxa of Chaetophora can be expected in lowland areas of Peru,
where they will probably be attracted to lights. As indicated above, montane species
remain undescribed. Johnson (2002) gave a general overview of the biology of the
family.
Checklist:
Byrrhinae: Simplocariini
Genus undescribed, species undescribed [may represent one or two species].
Syncalyptinae
Chaetophora Stephens, species remain unverified and unreported.
Acknowledgements
NSF-EPSCoR #66928 (PI: CS Chaboo) provided support for the Beetles of Peru
project and the University of Kansas Department of Ecology and Evolutionary
Accepted 22 April 2014; Revised 16 October 2014

Biology-General Research Fund (PI: CS Chaboo) for funding this publication. Luis
Figueroa and Mabel Alvarado are thanked for bringing the MUSM specimen to our
attention and to two anonymous reviewers.
Literature Cited
11492.
Dalla Torre, K. W. von 1911. Nosodendridae, Byrrhidae, Dermestidae. Coleopterorum Catalogus, Pars.
33. W. Junk, Berlin.
Johnson, P. J. 2002. Superfamily Byrrhoidea Latreille 1804. 42. Byrrhidae Latreille 1804. In Arnett, R. H.,
Jr., M. C. Thomas, P. E. Skelley, and J. H. Frank (eds.). American Beetles, vol. 2, Polyphaga:
Scarabaeoidea through Curculionoidea, pp. 113116. CRC Press, Boca Raton, Florida.
Reichardt, H. 1974. New and little known neotropical Coleoptera III. Syncalypta striata Pic, 1922, with
discussion on Syncalypta Stephens, 1830 (Byrrhidae). Papeis Avulsos de Zoologia 27(18):227234.
Reichardt, H. 1975. New and little known neotropical Coleoptera V. A new species of Syncalypta
(Byrrhidae). Papeis Avulsos de Zoologia 29(12):7980.
Elateridae Leach, 1815
Author(s): Paul J. Johnson and Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 269272

Elateridae Leach, 1815
PAUL J. JOHNSON1 AND CAROLINE S. CHABOO2,3

Recognition: The beetles of this family are immediately recognized by their
generally elongate and narrow body, a disproportionately large and loosely attached
prothorax that rapidly articulates with the mesothorax to snap the body for
predator escape and self-righting; serrate to pectinate antennae; large prosternum;
large subtriangular hypomeron; large subquadrate metaventrite; and slender legs
with long pentamerous tarsus, usually with ventral setal brushes or membranous
pads.
Habitat: Elaterids are found primarily in forests, savannas, thorn-forests, and
scrublands, with different taxa between wet and dry vegetation types. Some are
specialized by larval adaptions to arboreal and terrestrial bromeliads, epiphyte mats
on large tree limbs, sandy riparian soils, grassland soils, or various stages of internal
decay of dead and dying trees. Adult activity is short-lived and mostly crepuscular
and nocturnal, especially in warm to hot environments. Mid and upper elevation
species are often diurnal. They feed at high-carbohydrate sources such as plant
wounds and weeps, floral and extra-floral nectaries, decaying fruits, plant glandular
exudates, but also predate on small plant-feeding arthropods. Larvae are mostly
predators under loose bark, within decaying wood, among roots and rhizomes
of epiphytes, leaf axils of bromeliads, and in soil. Some species are rhizophages in
soil.
Notes: The checklist of Elateridae is based on Gemminger and Harold (1869),
Candeze (1891), Schwarz (19061908), Schenkling (1925, 1927), and Blackwelder
(1944, 1957), supplemented by corrections and additions from the original and more
recently published descriptions and records. Authors of non-original combinations
are given in ( ). Based on diversity recorded in neighboring countries the published
records for Peru indicate a large fauna awaiting discovery and reporting.
Checklist:
Agrypninae: Agrypnini
Dilobitarus bellus Casari, 2013 Dilobitarus quadrituberculatus Candeze, 1857
Dilobitarsus impressicollis Schwarz, 1902 Lacon palliatus (Latreille, 1811)
1
Insect Biodiversity Lab, Box 2207A, 1010 Rotunda Drive, South Dakota State University, Brookings,
South Dakota 57007, USA; e-mail: paul.johnson@sdstate.edu
2
Lawrence, Kansas, 66045, USA; e-mail: cschaboo@ku.edu
3
Agrypninae: Oophorini
Aeolus bivittatus Kirsch, 1873 Conoderus inquinatus (Candeze, 1859)
Aeolus gaudichaudi Guerin-Meneville, 1838 Conoderus intermissus (Kirsch, 1873)
Aeolus saulcyi Guerin-Meneville, 1838 Conoderus lacerosus (Schwarz, 1906)
Aeolus scriptus (Fabricius, 1801) Conoderus lituratus (Schwarz, 1900)
Aeolus terminatus Schwarz, 1906 Conoderus melanurus (Candeze, 1859)
Conoderus apiatus (Erichson, 1847) Conoderus peruanus (Schwarz, 1900)
Conoderus belti (Champion, 1895) Conoderus repandus (Erichson, 1847)
Conoderus bipartitus (Schwarz, 1900) Conoderus rufipennis (Schwarz, 1902)
Conoderus conspersus (Schwarz, 1903) Conoderus sticticus (Erichson, 1847)
Conoderus falli (Lane, 1956) Heteroderes amplicollis (Gyllenhal, 1817)
Conoderus ferrugineomarginatus (Brethes, 1920)
Agrypninae: Pyrophorini: Anaissiina

Alampoides alychnus (Kirsch, 1873) Anaissus girardi Riese, 2007
Alampoides badius Costa, 1975 Anaissus tarsalis Candeze, 1857
Alampoides fulvus Costa, 1975 Peralampes occiduus (Erichson, 1847)
Alampoides tessellatus (Candeze, 1863)
Agrypninae: Pyrophorini: Pyrophorina

Lygelater bifossulatus (Candeze, 1865) Pyrearinus scintillula (Candeze, 1881)
Lygelater indicus (Herbst, 1783) Pyrophorus angustus Blanchard, 1843
Opselater melanurus (Candeze, 1863) Pyrophorus divergens Eschscholtz, 1829
Pyrearinus amplicollis (Candeze, 1863) Pyrophorus dulcifer Costa, 1972
Pyrearinus basalis (Schwarz, 1902) Pyrophorus noctilucus Linnaeus, 1758
Pyrearinus lucidulus (Illiger, 1807)
Agrypnini: Hemirhipini
Alaus cinnamomoeus Casari, 2003 Chalcolepidius mocquerysi Candeze, 1857
Chalcolepidius aurulentus Candeze, 1874 Chalcolepidius peruanus Candeze, 1886
Chalcolepidius chalcantheus Candeze, 1857 Chalcolepidius porcatus (Linnaeus, 1787)
Chalcolepidius corpulentus Candeze, 1874 Chalcolepidius virens (Fabricius, 1787)
Chalcolepidius erythroloma Candeze, 1857 Chalcolepis similis Casari, 1999
Chalcolepidius fryi Candeze, 1874 Lacais glauca (Castelnau, 1836)
Chalcolepidius jansoni Candeze, 1874 Propalaus alicii (Pjatakowa, 1941)
Chalcolepidius limbatus Eschscholtz, 1829 Propalaus haroldi (Candeze, 1878)
Agrypninae: Platycrepidiini
Platycrepidius castus (Janson, 1882) Platycrepidius lamarrei Chassain, 2013
Negastriinae
Agrypnella squamifer (Candeze, 1865)
Semiotinae
Semiotus acutus Candeze, 1874 Semiotus cyrtomaris Wells, 2007
Semiotus affinis Guerin-Meneville, 1855 Semiotus fleutiauxi Szombathy, 1909
Semiotus angulatus (Drury, 1782) Semiotus fulvicollis Blanchard, 1843
Semiotus angustus Wells, 2007 Semiotus furcatus (Fabricius, 1792)
Semiotus antennalis Candeze, 1895 Semiotus gibbosus Wells, 2007
Semiotus bilineatus Candeze, 1857 Semiotus illigeri Guerin-Meneville, 1844
Semiotus bispinus Candeze, 1874 Semiotus imperialis (Guerin-Meneville, 1844)
Semiotus carus Janson, 1882 Semiotus intermedius (Herbst, 1806)
Semiotus convexicollis Blanchard, 1843 Semiotus kirschi Chassain, 1998
Semiotus cristatus Candeze, 1874 Semiotus ligneus (Linnaeus, 1767)
Semiotus cuspidatus splendidus, Candeze, 1881 Semiotus limatus Candeze, 1889
Semiotus nigriceps Candeze, 1857 Semiotus superbus Kirsch, 1866

Semiotus regalis Guerin-Meneville, 1844 Semiotus taeniatus Erichson, 1847
Semiotus schaumi Guerin-Meneville, 1844 Semiotus virgatus Erichson, 1847
Semiotus seladonius Guerin-Meneville, 1844
Lissominae
Lissomus argenteocaudatus Cobos, 1966 Lissomus mediotestaceus Cobos, 1966
Lissomus bifloccosus Castelnau, 1835
Cardiophorinae
Esthesopus biformis Schwarz, 1902 Horistonotus nigricollis Schwarz, 1902
Esthesopus hieroglyphicus Schwarz, 1902 Horistonotus peruvianus (Guerin-Meneville, 1839)
Esthesopus humeralis Steinheil, 1874 Horistonotus popularis (Erichson, 1847)
Esthesopus humilis Candeze, 1860 Horistonotus suturalis Schwarz, 1902
Horistonotus cleryi Guerin-Meneville, 1839 Horistonotus tetraspilotus (Guerin-Meneville, 1839)
Horistonotus exoletus (Erichson, 1840) Horistonotus unicolor Schwarz, 1902
Elaterinae: Elaterini
Diplostethus peruanus (Schwarz, 1903) Tomicephalus sanguinicollis Latreille, 1834
Probothrium furvum (Erichson, 1847)
Elaterini: Agriotini: Agriotina

Cardiorhinus baeri Fleutiaux, 1940
Elaterini: Agriotini: Pomachilina

Cosmesus adrastoides (Schwarz, 1900) Paracosmesus melanurus (Schwarz, 1900)
Cosmesus flavus (Schwarz, 1903) Paracosmesus terminatus Schwarz, 1902
Cosmesus gracilis Kirsch, 1873 Pomachilius andinus Brethes, 1937
Cosmesus maculipennis (Schwarz, 1900) Pomachilius apicalis Schwarz, 1900
Cosmesus pallidus Guerin-Meneville, 1838 Pomachilius maculifrons Schwarz, 1900
Cosmesus parvicollis (Schwarz, 1902) Pomachilius marginatis Schwarz, 1900
Cosmesus robustus (Schwarz, 1900) Pomachilius nigriceps Erichson, 1847
Paracosmesus basalis (Schwarz, 1900) Pomachilius spinosus Schwarz, 1900
Paracosmesus cruciatus (Schwarz, 1900) Pomachilius sulcifrons Schwarz, 1900
Paracosmesus dimidiatus (Schwarz, 1900) Pomachilius unifasciatus Schwarz, 1902
Elaterini: Ampedini: Ampedina

Agelasinus antennalis Schwarz, 1900 Agelasinus viridis Candeze, 1863
Elaterini: Ampedini: Dicrepidiina

Achrestus antennalis Schwarz, 1902 Crepidius flabellifer (Erichson, 1847)
Achrestus lamellicornis Schwarz, 1902 Crepidius ornatus Candeze, 1900
Achrestus suturalis Schwarz, 1902 Cyathodera spinipennis Schwarz, 1902
Achrestus trilineatus Schwarz, 1902 Dipropus ellipticus (Schwarz, 1904)
Achrestus venustus Champion, 1895 Dipropus ferrugatus (Schwarz, 1906)
Anoplischius anguinus (Erichson, 1847) Dipropus laterus (Schwarz, 1904)
Anoplischius conicipennis Schwarz, 1902 Dipropus nigrovittatus (Schwarz, 1902)
Anoplischius conicus Candeze, 1900 Dipropus puncticollis (Fabricius, 1801)
Anoplischius melanurus Schwarz, 1902 Heterocrepidius ventralis (Guerin-Meneville,
Anoplischius planicollis (Erichson, 1847) 1838)
Elaterinae: Ampedini: Physorhinini

Anchastus virgatus Candeze, 1897 Physorhinus lateralis Candeze, 1859
Physorhinus distigma Candeze, 1859 Physorhinus sexnotatus Steinheil, 1875
Physorhinus gudenzii Riese, 2009
Cebrioninae: Aplastini
Cylindroderus limbatus Schwarz, 1902 Octinodes aequatorius Candeze, 1889
Dodecacius nigricollis Schwarz, 1902 Octinodes dimidiatus (Schwarz, 1906)
Dodecacius testaceus Schwarz, 1902
Thylacosterninae
Balgus obconicus (Bonvouloir, 1875) Thylacosternus walckenaeri (Guerin-Meneville,
Balgus schnusei schnusei (Heller, 1914) 1843)
Acknowledgements
Evolutionary Biology-General Research Fund (PI: CS Chaboo). Jacques Chassain
and Matthew Gimmel are thanked for assisting in the compilation of species names.
We also thank two anonymous reviewers.
Literature Cited
11492.
Candeze, E. C. A. 1891. Catalogue Methodique des Elaterides connus en 1890. H. Vaillant-Carmanne,
Liege.
Gemminger, M., and E. de Harold. 1869. Catalogus Coleopterorum, hucusque descriptorum synonymicus
et systematicus. Part 5. Berlin.
Schenkling, S. 1925. Elateridae I. Coleopterorum Catalogus auspiciis et auxilio W. Junk. Pars 80. W. Junk,
Berlin.
Schenkling, S. 1927. Elateridae II Coleopterorum Catalogus auspiciis et auxilio W. Junk. Pars 80. W.
Junk, Berlin.
Schwarz, O. 19061907. Coleoptera, Fam. Elateridae, Fasc. 46A, Genera Insectorum. P. Wytsman,
Bruxelles. 370p+5pl.
Cerophytidae Latreille, 1834
Author(s): Caroline S. Chaboo and Paul J. Johnson

terms_of_use.
critical research.
88(2), 2015, pp. 273

Cerophytidae Latreille, 1834
CAROLINE S. CHABOO1,3 AND PAUL J. JOHNSON2

Recognition: This small family comprises only 22 extant species, with 17 species in
the New World (Costa et al., 2003, 2014)). The common name rare click beetles is
because of an adult clicking mechanism involving the prothorax and mesothorax like
the true click beetles (Elateridae). Adults are small, 49 mm long, strongly convex
dorsally, slightly flattened ventrally. The antennae are sexually dimorphic, being
pectinate in males and serrate in females (Johnson, 2002). The larvae are whitish,
have a soft, elongate body; the prognathous head has sucking mouthparts with
styliform mandibles and maxillae.
Habitat: Adults may be collected at light or Malaise traps, by beating dead
vegetation, in leaf litter, and in rotted and fungusy wood where larvae occur.
Notes: Costa et al. (2003) documented one species from Peru, a single male
specimen from Ucayali (Sinchona), collected in the 1940s. Phylogenies and keys to
the world genera are provided in Costa et al. (2003, 2014).
Checklist:
Brachycerophytum sinchona Costa et al., 2003
Acknowledgements
publication. I thank Cleide Costa, Museu de Zoologia da Universidade de Sao
Paulo, Brazil, and Matthew Gimmel, Santa Barbara Museum of Natural History,
for their help.
Literature Cited
Costa, C., S. A. Vanin, J. F. Lawrence, and S. Ide. 2003. Systematics and cladistic analysis of
Cerophytidae (Elateroidea: Coleoptera). Systematic Entomology, London, 28:375407. http://dx.
doi.org/10.1046/j.1365-3113.2003.00219.x
Costa, C., S. A. Vanin, and S. P. Rosa. 2014. Description of a new genus and species of Cerophytidae
(Coleoptera: Elateroidea) from Africa with a cladistic analysis of the family. Zootaxa 3878
(3):248260.
Johnson, P. J. 2002. 55. Cerophytidae Latreille 1834. In Arnett, R. H. Jr., M. C. Thomas, P. E. Skelley,
and J. H. Frank (eds.). American Beetles vol. 2, p. 150151.
1
Lawrence, Kansas 66045, USA; e-mail: cschaboo@ku.edu
2
Insect Biodiversity Lab, Box 2207A, 1010 Rotunda Drive, South Dakota State University Brookings,
South Dakota 57007, USA; e-mail: paul.johnson@sdstate.edu
3
Erotylidae Latreille, 1802
Author(s): Joseph V. McHugh and Caroline S. Chaboo

terms_of_use.
critical research.
88(2), 2015, pp. 274282
Beetles (Coleoptera) of Peru: A Survey of the Families. Erotylidae

Latreille, 1802
JOSEPH V. MCHUGH1 AND CAROLINE S. CHABOO2,3

Recognition: The common name for Erotylidae is the Pleasing fungus beetles
due to the attractive adult coloration and mycophagous habits of many species.
Erotylidae, including Languriidae, Lizard beetles, form a monophyletic group that
is supported by phylogenetic analyses using morphological (e.g., Wegrzynowicz,
2002; Leschen and Buckley, 2007) and molecular (e.g., Robertson et al., 2004) data.
Beetles in the erotylid familial clade may be recognized by the following combination
of derived features: subocular glandular ducts present, supraocular line present,
pronotal lateral carina simple, procoxal cavities with lateral extensions, trochantinal
notch present, mesocoxal cavities laterally closed by metaventrite, and aedeagus
laterally compressed with relatively long struts on penis (Leschen et al., 2010).
Adults of Peruvian erotylids are 1.223.0 mm. long and range in body form from
parallel-sided to oval. The base of the prothorax is not conspicuously narrower than
the elytral bases. They vary in convexity from being dorso-ventrally flattened to
having a hemispherical form. In some species (e.g., Gibbifer spp.) the elytra are
highly vaulted. The dorsal surfaces are typically smooth and either glabrous or have
fine setae. Erotylids bear fine pores for glandular ducts on many structures,
especially the prothoracic lateral carina, genae, mentum, thoracic ventrites, and
around the eyes.
The antennae are 11-segmented and capitate. The club is flattened and 36
segmented. The mandibles are short and broad with a bidentate or tridentate apex
and a well-developed mola and prostheca. The maxillae have a well-developed galea
and lacinia. The terminal segment on the maxillary and labial palpi varies from
fusiform to being strongly expanded and truncate apically.
The pronotum usually has a smooth lateral margin and a complete carina with
a raised margin or bead. The anterior lateral pronotal angles are either produced
forward or simple, the posterior angles are not produced. The posterior margin is
usually evenly rounded, but a mesal lobe occurs in some. The pronotal disc is
typically smooth, but at the base there may be a pair of weak basal impressions,
a pair of pits, or a transverse depression.
The scutellum is visible dorsally and usually broadly rounded or broadly
pentagonal. The elytra have striate to confused punctation. A scutellary striole
may be present or absent. The epipleura are usually complete to apex.
The procoxae do not project ventrally. The procoxal cavities are slightly transverse
or circular and externally open or closed. Protrochantins are not exposed. The
1
Department of Entomology, Biological Sciences Building, University of Georgia, Athens, Georgia,
30602, U.S.A.; e-mail: mchugh.jv@gmail.com
2
3
Corresponding author. e-mail: cschaboo@ku.edu
Accepted 11 May 2015; Revised 17 July 2015
mesoventrite and metaventrite are on same plane. The mesocoxal cavities are
subovate and moderately to widely separated. The metaventrite is slightly to strongly
convex and often bears a discrimen. The metacoxae are transverse and contiguous to
widely separated. The hind wings are relatively long, but can be reduced or lost in
some species. The venation is variable but usually includes a complete r4 vein and
a radial-medial loop that forms an acute angle. The medial fleck is present. An anal
lobe may be present or absent.
The legs are slender and long to very long. The tarsi are 555, but can appear 444.
Ventral lobes may be present on one or more tarsomeres. The abdomen has 5 ventrites.
Either all ventrites are free or ventrites 1 and 2 are connate. Ventrite 1 is only slightly
longer than ventrite 2. The intercoxal process is present and variable in form. The
aedeagus is usually symmetrical, uninverted, and laterally compressed with an
anteriorly rounded tegmen and parameres that are individually articulated to the
phallobase. In females, sternite VIII bears a well-developed spiculum ventrale. The
ovipositor is long and usually bears well developed styli. The spermatheca is sclerotized
and variable in form. For further details of adult morphology, please review: Vaurie,
1948; Boyle, 1956; McHugh et al., 1997; Lawrence et al., 1999a; Wegrzynowicz, 2002;
Leschen, 2003; Leschen et al., 2010; Skelley and Leschen, 2007; and Leschen et al., 2010.
It is difficult to characterize the larval form of Erotylidae because there is great
diversity in the gross morphology of immatures in this group, undoubtedly due in
part to the diversity of their feeding behaviors and habitats. Mature larvae range
from 225 mm. The body may be parallel-sided to fusiform, slightly flattened to
rounded in cross-section, and unpigmented to strongly pigmented. The dorsal
surfaces may be smooth or bear some form of integumental armature, including fine
asperities, setiferous tubercles, granules, spines, or complex scoli. The vestiture is
typically sparse to moderately dense, and usually simple.
The head is usually protracted. The epicranial stem may be present or absent. The
frontal arms are usually contiguous at the base and either lyriform or V-shaped.
There may be 6, 5, 2, or 0 stemmata. The antennae range from short to very long.
Ventral mouthparts are retracted. The labial palps are 2-segmented. The gular
sutures are separate, defining a gula that is wider than long. The thoracic and
abdominal terga may have plates. These plates may bear 2 or more transverse rows
of setiferous tubercles, senti, or scoli. The tergal surface often has patches of
asperities or granules. Lateral tergal processes may be present and can be complex.
The legs are well developed and 5-segmented, with a tarsungulus that is acute and
claw-like. The abdomen is more than twice as long as thorax. Tergum IX usually
bears a pair of urogomphi which vary in form from simple upturned hooks, to
complex spines, to very long threadlike structures. Pregomphi are often present.
Sternum IX is simple, not enclosed by sternum VIII. The anal region is
posteroventrally oriented. The spiracles are either annular or annular-biforous,
and sometimes occur at ends of spiracular tubes. For further review of juvenile
morphology, please review: Candeze, 1861; Boving and Craighead, 1931; Chujo,
1936; Roberts, 1939, 1958; Niimura, 1939; Nobuchi, 19541955; Hayashi et al., 1959;
Graves, 1965; Hayashi and Takanaka, 1965; Sen Gupta, 1969; Kompantsev, 1982;
Hayashi, 1986; Costa et al., 1988; Skelley, 1988; Skelley et al., 1997; Teixeira and
Casari, 1998; Lawrence, 1991; Lawrence et al., 1999b; and McHugh, 2001.
Habitat: Erotylidae typically occur in forested habitats and feed on fungi, plants, or
decaying plant matter. The mycophagous erotylids are usually specialists on
Basidiomycetes and are often found in close proximity to their host fungi. Some
erotylid species are associated with leaf litter, rotting logs, and other forms of decaying
plant matter that are permeated with fungi. Truly phytophagous species include stem
borers and pollen feeders. Only a few erotylids are of economic importance. Some
mycophagous species are considered pests of stored products and mushroom farms. A
few phytophagous species are pests of crops, especially legumes and composites
(Vaurie, 1948). Some pollenivorous erotylids are considered beneficial species due to
their role as pollinators of cycads (Franz and Skelley, 2008; Tang, 1987).
The family includes nocturnal and diurnal species. Many of the larger
mycophagous species are best sampled directly from their fungal hosts during the
day or at night. Phytophagous erotylids are best collected by sweeping host
vegetation. Active flying species are collected regularly at blacklights or with malaise
traps. Some of the smaller, cryptic species are best collected by Berlese funnel
extractions from decaying leaf litter or other rotting plant material.
The brilliant coloration of many Erotylinae and Languriinae is clearly aposematic.
Little is known about chemical defenses in Erotylidae; however, most species bear
pores on many parts of the body and some species are known to ooze glandular
secretions from these pores when disturbed (Arrow, 1925; McHugh et al., 1997;
Wegrzynowicz, 2002; Drilling et al., 2010; Drilling and Dettner, 2010). Strikingly
colorful erotylids appear to be components in mimicry complexes with other beetles in
many regions although their exact role in these systems remains unclear (Arrow, 1925).
Different types of gregariousness have been recorded for Erotylidae. Some species
only exhibit this behavior as adults. Other species are gregarious as larvae, and of
those, some form pupal aggregations as well (Graves, 1965; JVM, pers. obs.).
Gregariousness in erotylid species is triggered by daily cues in some species (Park
and Sejba, 1935; McHugh et al., 1997) and by seasonal cues in others (Goodrich and
Skelley, 1991; Navarrete-Heredia and Novelo-Gutierrez, 2000). In the genus
Pselaphacus, parental care has been documented for several species (see Chaboo
and McHugh, 2010). Pselaphacus larvae aggregate into conspicuous writhing balls
that move in unison across fruitings of their host fungus. The mother closely tends
these larval masses and shepherds them from one cluster of fungi to the next. Many
erotylids have stridulatory files on the dorsal surface of the head capsule which are
used in conjunction with a scraping device on the anterior portion of the pronotum.
Some erotylid species are known to use these organs to produce species-specific and
context-specific calls (Ohya, 2001).
The evolutionary relationships of Erotylidae have received attention recently in
formal phylogenetic studies that utilized different character suites and taxa (e.g.,
Wegrzynowicz, 2002; Robertson et al., 2004; Leschen and Buckley, 2007). While
a few general aspects of erotylid evolution were supported by all of these studies,
much uncertainty remains. Many difficult taxonomic issues in this family cannot be
addressed until the internal phylogenetic relationships are clarified.
Notes: The following list is derived from published works by Alvarenga (1994),
Blackwelder (1945), Bruce (1952), Campaner et al. (2008), Deelder (1942), ICZN
(1996), Lopes (2006), Mader (1942, 1944, 1951), Martins and Pereira (1965), Skelley
(1998), Skelley and Goodrich (1994), and Skelley et al. (1997). New biodiversity
occurrence data (indicated by *) were provided by an unpublished dissertation by
Peterson L. Lopes (2009) and from the following digitally published sources:
University of Georgia Collection of Arthropods (UGCA, accessed from holdings list
on museum website, http://ugca.uga.edu, 24th April 2015), Snow Entomological

Museum Collection (SEMC, accessed through iDigBio Specimen Data Portal,
https://www.idigbio.org/portal, 24th April 2015), Lund Museum of Zoology - Insect
collections (MZLU, accessed through GBIF.org, 24th April 2015, GBIF Occurrence
Download http://doi.org/10.15468/dl.fbgfgc.
In many museums, a large percentage of Neotropical erotylid specimens remain
undetermined. Revisionary work is greatly needed for most taxa that occur in this
region. The list provided below is only a preliminary step toward understanding the
rich Peruvian erotylid fauna.
Checklist:
Pharaxonothinae
Pharaxonotha kirschii (Reitter, 1875)
Loberinae
Loberus tropicus Kirsch, 1873
Languriinae: Hapalipini
Hapalips hispidus Bruce, 1952
Languriinae: Thallisellini
Acryptophagus sp. * [SEMC] Pseudhapalips lamellifer Champion, 1913 * [SEMC]
Platoberus sp. * [MZLU] Thallisella peruviana Crotch, 1876
Languriinae: Languriini
Brasilanguria flavipes (Fowler, 1886) Crotchia metallica Fowler, 1886
Camptocarpus longicollis cylindricollis (Kirsch, Crotchia nana Martins, 1965
1876) Goniolanguria colombiana Martins & Pereira, 1965
Compsolanguria reichei (Crotch, 1876) Goniolanguria excelsa Martins & Pereira, 1965
Crotchia coptengoides Fowler, 1886 Goniolanguria intermedia Martins & Pereira, 1965
Crotchia gibbosa Fowler, 1886 Langurites verticalis (Erichson, 1847)
Cryptophilinae: Toramini
Empocryptus sp.
Toramus bifasciatus Kirsch, 1873
Erotylinae: Megalodacnini
Megalodacne indica latifasciata Delkeskamp, 1952
Erotylinae: Tritomini
Pselaphacus contaminatus Erichson, 1847 Ischyrus duponti Lacordaire, 1842 * [UGCA]
Pselaphacus curvipes Guerin-Meneville, 1841 * Ischyrus fraternus (Lacordaire 1842) * [SEMC,
[UGCA] as Micrischyrus fraternus]
Pselaphacus gorhami Kuhnt, 1909 Ischyrus grammicus Gorham, 1883
Pselaphacus mysticus Gorham, 1883 Ischyrus kempferi (Alvarenga, 1977)
Pselaphacus nigropunctatus Percheron, 1835 Ischyrus mimus Skelley, 1998
Pselaphacus puncticollis Guerin-Meneville, 1841 Ischyrus mystacis Skelley, 1998
Pselaphacus rubricatus (Herbst, 1799) Ischyrus peruae Mader, 1942
Pselaphacus signatus Guerin-Meneville, 1841 Ischyrus peruvianus Gorham, 1883
Ischyrus auriculatus Lacordaire, 1842 Ischyrus quadripunctatus (Olivier, 1792)
Ischyrus boucardi Crotch, 1876 Ischyrus scriptus (Olivier, 1807)
Ischyrus chacojae Gorham, 1887 Ischyrus tetrasticus Gorham, 1887
Ischyrus disconigrum Mader, 1942 Ischyrus tripunctatus Crotch, 1873
Ischyrus velatus Lacordaire, 1842 Mycotretus lepidus Lacordaire, 1842 * [UGCA]

Megischyrus bartleti Gorham, 1883 Mycotretus maculatus Olivier, 1792 * [SEMC]
Megischyrus catenatus (Crotch, 1876) Mycotretus multimaculatus Taschenberg, 1870
Megischyrus circumcinctus Delkeskamp, 1957 Mycotretus opalescens Crotch, 1876 * [UGCA]
Megischyrus columbianus (Lacordaire, 1842) Mycotretus opalizans Mader, 1942
Megischyrus decempunctatus (Guerin-Meneville, Mycotretus ornatus (Duponchel, 1825)
1841) Mycotretus pebasensis Crotch, 1876
Megischyrus elongatus Gorham, 1883 Mycotretus pelliciens Kirsch, 1876
Megischyrus jurinei (Lacordaire, 1842) Mycotretus peruae Crotch, 1876
Megischyrus laetus (Schenkling, 1919) Mycotretus prioteloides Mader, 1942
Megischyrus planior (Kirsch, 1876) Mycotretus pygmaeus Lacordaire, 1842
Megischyrus semitinctus (Erichson, 1847) Mycotretus quadripunctatus Crotch, 1876
Megischyrus sicarius (Lacordaire, 1842) Mycotretus quadristriolatus Kuhnt, 1910
Megischyrus zonalis (Lacordaire, 1842) Mycotretus savignyi Lacordaire, 1842
Callischyrus cyanopterus (Erichson, 1847) Mycotretus scitulus Lacordaire, 1842 * [SEMC]
Callischyrus insignis (Laporte, 1840) Mycotretus sericeonitens Crotch, 1876
Cyrtomorphus pusillus Kirsch, 1876 Mycotretus suturalis Kirsch, 1876
Mycotretus adalioides Crotch, 1876 Mycotretus thoracicus Kuhnt, 1910
Mycotretus antesignatus Mader, 1942 Mycotretus tigrinoides Mader, 1942
Mycotretus apicalis Lacordaire, 1842 Mycotretus tigrinus (Olivier, 1792)
Mycotretus bicolor Taschenberg, 1876 Mycotretus tricolor Crotch, 1876
Mycotretus bicoloratus Kuhnt, 1911 Mycotretus tucuruiensis Alvarenga, 1983
Mycotretus cinctellus (Guerin-Meneville, 1841) Mycophtorus peruvianus Kirsch, 1876
Mycotretus cordiger Crotch, 1876 * [SEMC] Pseudolybas sp. * [SEMC]
Mycotretus cunctans Mader, 1942 Lybas atripennis Erichson, 1847
Mycotretus derasofasciatus Kuhnt, 1910 Lybas bicolor (Guerin-Meneville, 1841)
Mycotretus dichrous Kirsch, 1876 Lybas dorsalis Gorham, 1883
Mycotretus discoidalis Taschenberg, 1870 Lybas kleinei Mader, 1943
Mycotretus dispar Taschenberg, 1870 Lybas normalis Lacordaire, 1842 * [SEMC]
Mycotretus dorsonotatus Lacordaire, 1842 Lybas puncticollis Mader, 1942
Mycotretus duodecimguttatus (Duponchel, 1825) Lybanodes bicolor Skelley, 1997
Mycotretus fidelis Delkeskamp, 1939 Lybanodes stigmatus Skelley, 1997
Mycotretus floriger Lacordaire, 1842 * [UGCA] Mycolybas lucidus Lacordaire, 1842
Mycotretus gemmula Lacordaire, 1842
Erotylinae: Erotylini
Cyclomorphus octopunctatus Erichson, 1847 Aegithus punctatissimus (Fabricius, 1775)
Cyclomorphus sicardi Kuhnt, 1910 Aegithus sanguinans Dohrn, 1880
Strongylosomus alutaceus (Erichson, 1847) Aegithus satellitius Lacordaire, 1842
Strongylosomus dichrous (Lacordaire, 1842) Aegithus striatellus Crotch, 1876
Strongylosomus peruvianus (Crotch, 1876) Iphiclus (Megaprotus) circulus (Lacordaire, 1842)
Strongylosomus rugosus (Lacordaire, 1842) Iphiclus (Megaprotus) dilectus (Gorham, 1888)
Aegithus armitagei Gorham, 1889 Iphiclus (Megaprotus) duplicatus (Lacordaire, 1842)
Aegithus bartletti Gorham, 1889 Iphiclus (Megaprotus) laetus (Crotch, 1876)
Aegithus brunnipennis Lacordaire, 1842 Iphiclus (Megaprotus) octolinearis (Kuhnt, 1910)
Aegithus burmeisteri Lacordaire, 1842 Iphiclus (Megaprotus) octopunctatus (Kirsch, 1876)
Aegithus circumfusus Kuhnt, 1908 Iphiclus (Megaprotus) perlepidus (Lacordaire, 1842)
Aegithus clavicornis (Linnaeus, 1758) Iphiclus (Megaprotus) sedecimlinearis (Mader, 1942)
Aegithus consularis Guerin-Meneville, 1855 Iphiclus (Megaprotus) sedecimstrigatus (Mader, 1942)
Aegithus cribrosus Lacordaire, 1842 Iphiclus (Megaprotus) zonulus (Crotch, 1873)
Aegithus cyanipennis Guerin-Meneville, 1841 Iphiclus (Habrodactylus) fulvipennis (Erichson, 1847)
Aegithus dohrni Mader, 1942 Iphiclus (Habrodactylus) grammicus (Erichson, 1847)
Aegithus hemisphaericus Lacordaire, 1842 Iphiclus (Habrodactylus) obliquatus (Kuhnt, 1910)
Aegithus inflatus Crotch, 1876 Iphiclus (Habrodactylus) oblongonotatus (Lacor-
Aegithus luteus Erichson, 1847 daire, 1842)
Aegithus multistriatus Mader, 1942 Iphiclus (Habrodactylus) proximus (Guerin-Meneville,
Aegithus nigrocinctus Erichson, 1847 1844)
Aegithus politus Gorham, 1888 * [UGCA] Iphiclus (Habrodactylus) rufescens (Lacordaire, 1842)
Iphiclus (Habrodactylus) rufifrons (Lacordaire, 1842) Iphiclus (Brachymerus) quinquefasciatus Lacor-

Iphiclus (Habrodactylus) tabidus (Erichson, 1847) daire, 1842 * [UGCA]
Iphiclus (Habrodactylus) thoracicus (Kirsch, 1876) Iphiclus (Brachymerus) salamandra (Erichson, 1847)
Iphiclus (Sternolobus) luscus (Erichson, 1847) Iphiclus (Brachymerus) tricinctus (Duponchel, 1825)
Iphiclus (s. str.) guttiger (Kuhnt, 1910) Iphiclus (Brachymerus) ucayalensis (Gorham, 1889)
Iphiclus (s. str.) intersectus (Duponchel, 1825) * Iphiclus (Brachymerus) westwoodi (Guerin-Meneville,
[SEMC] 1841)
Iphiclus (s. str.) sedecimmaculatus Buquet, 1840 Erotylus crotchi Deelder, 1942
Iphiclus (s. str.) titschacki (Mader, 1942) Erotylus crucifer Kuhnt, 1908
Iphiclus (Saccomorphus) adamsi (Lacordaire, 1842) Erotylus decipiens Crotch, 1876
Iphiclus (Saccomorphus) erotyloides (Crotch, 1876) Erotylus dilaceratus Kirsch, 1876
Iphiclus (Saccomorphus) glyptoderus (Lacordaire, Erotylus elongatulus Crotch, 1876
1842) Erotylus flavopunctatus Kuhnt, 1908
Iphiclus (Saccomorphus) intercedens (Schenkling, Erotylus imitans Kirsch, 1876
1919) Erotylus incertus Lacordaire, 1842
Iphiclus (Saccomorphus) interruptus (Kuhnt, 1908) Erotylus incomparabilis Perty, 1832
Iphiclus (Saccomorphus) mutabilis (Gorham, 1889) Erotylus jucundus Mader, 1942
Iphiclus (Saccomorphus) nebulosus (Guerin-Meneville, Erotylus kuhnti Deelder, 1942
1841) Erotylus laetus Deelder, 1942
Iphiclus (Saccomorphus) placitus (Kirsch, 1876) Erotylus loratus Erichson, 1847
Iphiclus (Saccomorphus) procerus (Erichson, 1847) Erotylus margineguttatus Crotch, 1876
Iphiclus (Saccomorphus) quadrisignatus (Duponchel, Erotylus melanostictus Crotch, 1876
1825) Erotylus nautae Crotch, 1876
Iphiclus (Saccomorphus) ruficeps (Guerin-Meneville, Erotylus neglectus Mader, 1942
1841) Erotylus nigrocinctus Kuhnt, 1908
Iphiclus (Neomorphoides) atriventris (Mader, 1943) Erotylus olivieri Lacordaire, 1842
Iphiclus (Neomorphoides) disconigrum (Mader, 1942) Erotylus parcepunctatus Crotch, 1876
Iphiclus (Neomorphoides) lateripunctatus (Crotch, Erotylus parvus Kuhnt, 1908
1876) Erotylus peregrinus Mader, 1942
Iphiclus (Neomorphoides) nigritarsis (Mader, 1942) Erotylus peruvianus Crotch, 1876
Iphiclus (Neomorphoides) posticenigrum (Mader, Erotylus rudepunctatus Crotch, 1876
1942) Erotylus scenicus Erichson, 1847
Iphiclus (Neomorphoides) pyrrhocephalus (Erichson, Erotylus singularis Kirsch, 1876
1847) Erotylus spectrum Thomson, 1856
Iphiclus (Neomorphoides) rufipennis (Fabricius, Erotylus staudingeri Mader, 1935
1801) Erotylus subreticulatus Guerin-Meneville, 1841
Iphiclus (Neomorphoides) simplex (Lacordaire, 1842) Erotylus superbus Guerin, 1956
Iphiclus (Brachymerus) amictus (Erichson, 1847) Erotylus taeniatus Latreille, 1813
Iphiclus (Brachymerus) andicola (Kirsch, 1876) Erotylus trichromaticus Alvarenga, 1976
Iphiclus (Brachymerus) batesi (Gorham, 1889) * Erotylus ustulatus Erichson, 1847
[SEMC] Erotylus varians Crotch, 1876
Iphiclus (Brachymerus) bicinctus (Olivier, 1807) Erotylus variomaculatus Kuhnt, 1908
Iphiclus (Brachymerus) bistrifoliatus (Gorham, 1889) Erotylus voeti Lacordaire, 1842
Iphiclus (Brachymerus) dorsalis (Olivier, 1792) Erotylus ziczac Taschenberg, 1870
Iphiclus (Brachymerus) epipleuralis (Crotch, 1876) Erotylina atrotibialis Alvarenga, 1976
Iphiclus (Brachymerus) incas (Gorham, 1889) Erotylina bardia Curran, 1944
Iphiclus (Brachymerus) jacinthoi Alvarenga, 1977 Erotylina bassleri Curran, 1944
Iphiclus (Brachymerus) miles (Mader, 1942) Erotylina ecuadorica (Crotch, 1876)
Iphiclus (Brachymerus) mirus (Mader, 1942) Erotylina jaspidea (Erichson, 1847)
Iphiclus (Brachymerus) nigripennis (Demay, 1838) Erotylina maculiventris (Lacordaire, 1842)
* [SEMC] Erotylina multiguttata (Lacordaire, 1842)
Iphiclus (Brachymerus) nigrotrifasciatus (Mader, Erotylina scita Curran, 1944 * [UGCA]
1942) Erotylina toxophora (Lacordaire, 1842)
Iphiclus (Brachymerus) odyneroides Crotch, 1876 * Gibbifer anthracinus (Gorham, 1888)
[UGCA] Gibbifer armillatus (Erichson, 1847)
Iphiclus (Brachymerus) perplexus (Mader, 1942) Gibbifer ater (Kirsch, 1876)
Iphiclus (Brachymerus) peruvianus (Mader, 1942) Gibbifer badeni (Dohrn, 1883)
Iphiclus (Brachymerus) puncticollis (Kirsch, 1876) Gibbifer dromedarius (Lacordaire, 1842)
Gibbifer gibbosus (Linnaeus, 1763) Prepopharus herbsti (Lacordaire, 1842)

Gibbifer gracilis (Kuhnt, 1908) Prepopharus impluviatus (Lacordaire, 1842)
Gibbifer irroratus (Kuhnt, 1908) Prepopharus obliteratus (Erichson, 1847)
Gibbifer maximus (Crotch, 1876) Prepopharus partitus (Crotch, 1876)
Gibbifer miliaris (Lacordaire, 1842) Prepopharus tricolor (Mader, 1942)
Gibbifer patellatus (Gorham, 1888) Neopriotelus bimaculatus (Mader, 1942)
Gibbifer sticticus (Erichson, 1847) Neopriotelus elegans (Mader, 1942)
Gibbifer stillatus (Kirsch, 1865) Neopriotelus obsoletus Crotch, 1876 * [UGCA]
Gibbifer tigrinus Voet, 1778 Neopriotelus peruvianus (Mader, 1942)
Gibbifer zebu (Kirsch, 1876) Neopriotelus stellio (Erichson, 1847)
Barytopus dubitabilis (Crotch, 1876) Neopriotelus zebra (Kuhnt, 1910)
Barytopus hesitans (Crotch, 1876) Phricobacis arduus (Erichson, 1847)
Barytopus lunulatus (Olivier, 1792) Phricobacis flavomaculatus Mader, 1943
Oligocorynus convexus (Crotch, 1876) * Lopes Phricobacis marginatus (Guerin-Meneville, 1841)
(2009, unpub. dissertation) Ellipticus (s. str.) atripennis (Mader, 1942)
Oligocorynus hybridus (Erichson, 1847) Ellipticus (s. str.) dorbignyi (Guerin-Meneville, 1841)
Oligocorynus militarioides (Mader, 1942) Ellipticus (s. str.) hepaticus (Crotch, 1876)
Oligocorynus peruvianus (Mader, 1942) Ellipticus (s. str.) niger (Deelder, 1942)
Oligocorynus rugipunctatus Crotch, 1876 * Lopes Ellipticus (s. str.) octomaculatus (Crotch, 1876)
(2009, unpub. dissertation) Ellipticus (s. str.) orbignyanus (Lacordaire, 1842)
Oligocorynus zebra (Fabricius, 1787) Ellipticus (s. str.) peruvianus (Kirsch, 1876)
Scaphidomorphus bosci Guerin-Meneville, 1841 Ellipticus (s. str.) spinifer (Thomson, 1857)
Prepopharus americanus (Voet, 1778) Ellipticus (s. str.) terminalis (Erichson, 1847)
Prepopharus bitaeniatus (Lacordaire, 1842) Ellipticus (s. str.) testaceus (Fabricius, 1775)
Prepopharus diffinis Erichson, 1847 Ellipticus (Inflatotelus) inflatus (Mader, 1942)
Prepopharus eduardoi Alvarenga, 1976 Ellipticus (Inflatotelus) laevis (Delkeskamp, 1957)
Acknowledgements
publication. We thank Mathew Gimmel for help assembling some names and Paul
Skelley and one anonymous reviewer for comments that improved the manuscript.
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Staphylinidae Latreille, 1802
Author(s): Alfred F. Newton

terms_of_use.
critical research.
88(2), 2015, pp. 283304
Beetles (Coleoptera) of Peru: A Survey of the Families. Staphylinidae

Latreille, 1802
ALFRED F. NEWTON
Integrative Research Center, Field Museum of Natural History, 1400 South Lake
Shore Drive, Chicago, Illinois 60605, USA. E-mail: anewton@fieldmuseum.org
Diversity in Peru: 17 subfamilies, 221 genera, 877 species (confirmed).

Recognition: Staphylinidae or rove beetles range in size from 135 mm (most are
28 mm) and vary greatly in shape from very compact to extremely slender or even
antlike, but most may be recognized by having more or less truncate elytra (exposing
one to many abdominal tergites), six or seven visible abdominal sternites, and
contiguous procoxae which vary in shape but are often prominent and conical. The
antennae are usually 11segmented and filiform to weakly clavate (rarely with
a loose to compact antennal club or reduced segmentation), and tarsi are often five
segmented but may have only four, three or two segments or various heteromerous
combinations. Some members possess paired ocelli dorsally on the head.
Habitat: Rove beetles can be found almost everywhere beetles occur, from
seashore to alpine grasslands (e.g., Thayer, 2005). They are especially abundant in
forest habitats, including in leaf litter and rotting logs, on fungi and vegetation, and
attracted to dung and other decaying materials, and in non-forested moist habitats
such as marine beaches and near lakes, rivers and bogs. They tend to be scarce in
arid and curtivated or other highly disturbed habitats. A majority of species are
predators of other invertebrates as adults and larvae, but many are saprophages or
mycophages, and some feed on algae or pollen, but virtually none feed on living
green plants. Many species are inquilines of social insects, especially ants and
termites, and these are often well integrated with their host societies (e.g., Kistner,
1982), but some species are found in vertebrate nests or even, in one group, are
commensals living on the bodies of small mammals (Seevers, 1955). Some species of
the genus Paederus Fabricius in Peru (and elsewhere) have a toxic chemical, pederin,
in their haemolymph that can cause severe dermatitis or lesions in humans (known
locally as dermatitis purulenta, internationally as Paederus dermatitis or linear
dermatitis) and thus are of medical and even economic importance (e.g., Ojeda
Pena, 1969; Guillen, 1989; Vieira et al., 2014). Other Peruvian staphylinids are
potentially beneficial to humans as predators of pest arthropods (e.g., Schuller and
Sanchez, 2003), including species of the genus Holobus Solier that prey on red spider
mites (aranita roja) infesting cotton plants (Korytkowski, 1966, as Somatium
Wollaston sp.), and larvae and adults of Platystethus spiculus Erichson that, at least
under laboratory conditions, prey on eggs of the disease-carrying house fly Musca
domestica Linnaeus (Palomino and Dale, 1989).
Notes: Staphylinidae, with more than 61,300 described extant species, is now the
most diverse animal family in the world after addition of the former families
Pselaphidae, Scaphidiidae and Scydmaenidae as subfamilies, as noted by Grebenni-
kov and Newton (2009). The overall classification, phylogeny, world distribution,
Accepted 29 August 2014; Revised 9 March 2015

morphology and biology were reviewed most recently by Thayer (2005), from which
much of the information in this introduction was gleaned. The Neotropical region
has one of the most diverse regional faunas, but is relatively less well studied than the
Holarctic and Oriental regions. Guides including identification keys to genera, many
of which occur in Peru, have been published for Mexico (Navarrete-Heredia et al.,
2002) and the West Indies (Blackwelder, 1943) but not for any South American
countries, although a faunal overview for staphylinids of Argentina was presented by
Chani-Posse and Thayer (2008). In addition to the now-outdated catalog of
Neotropical Coleoptera prepared by Blackwelder (1944), recent species checklists for
Staphylinidae have been produced for Colombia (Newton et al., 2005), Chile and
southern Argentina (Newton and Thayer, 2005, ,http://archive.fieldmuseum.org/
peet_staph/db_1b.html.), and Brazil (Asenjo et al., 2013), and the monumental
worldwide catalog of Herman (2001) includes all except five of the 32 Recent
staphylinid subfamilies.
For Peru, the most recent published summary for Staphylinidae is the preliminary
checklist of Asenjo (2004), which listed 394 species in 124 genera, based largely on the
catalogs of Blackwelder (1944) and Herman (2001). The known staphylinid diversity
in Peru has now more than doubled in a decade, to 877 species in 221 genera, with
nearly three dozen species and a half dozen genera listed as probable additions below.
This large increase is due in part to the addition of Scydmaeninae (70 species) to the
family, but is also based on a combination of new data from the more recent regional
catalogs cited above, new or overlooked data accumulated in a world catalog in
preparation by A.F. Newton (to be made available online soon), and a recent increase
in taxonomic activity in certain groups of Neotropical staphylinids. A few subfamilies
have received relatively comprehensive recent treatments, so that a substantial
percentage of the available collections of these groups have been studied and many
new species described, e.g., in Megalopsidiinae (Puthz, 2012), Osoriinae (Irmler,
20032015), Piestinae (Caron et al., 2012) and Steninae (Puthz, 19882014). In
contrast, other subfamilies such as Oxytelinae, Pselaphinae, Scaphidiinae and
Tachyporinae have been virtually unstudied for more than a century, and even the
known collections of these moderately-to-very diverse groups consist of mostly
undescribed species. In addition, most of Peru has not yet been systematically
surveyed for staphylinids, so that large numbers of new species can be expected to be
added to collections when this is accomplished. Four hundred and twenty-four of the
currently known 877 species (48%) are endemic (precinctive) to Peru as far as known,
a high percentage for one country in a large continent, and indicative of its important
contribution to world biodiversity. Although these numbers do not yet reflect it
because of our poor state of knowledge of Neotropical Staphylinidae, this group is
almost certainly the most species-rich beetle family in Peru, and the actual faunal
diversity can be expected to approach that of the roughly 4000 or more species that
were suggested for Peru by Asenjo (2004), and the 5000 predicted for Colombia
(Newton et al., 2005) and Mexico (Navarrete-Heredia et al., 2002) and nearly realized
already for North America north of Mexico (Newton et al., 2000).
The checklist below includes the current name of each species known to occur in
Peru through June 2015, with authors of non-original combinations in (); a source
(usually a literature citation) for the occurrence of the species in Peru in []; and in
some cases synonyms that have been previously reported from Peru. Full
synonymies and references to original descriptions are generally not included but
may be found in the cited sources and/or the catalogs cited above. Species that are
endemic (precinctive) to Peru are preceded by an asterisk (*). The names of those
species considered unconfirmed because they were doubtfully or erroneously
reported from Peru, or are here considered probable for Peru due to their overall
distribution pattern, are enclosed in [] and an explanatory note added to each
(probable here means reported from Ecuador plus Bolivia and/or Chile). Species
that are newly recorded from Peru here are listed in bold, with an indication of who
made the identification and where the specimens are deposited using the following
collection codens: BMNH (The Natural History Museum, London), FMNH (Field
Museum of Natural History, Chicago) and SEMC (Snow Entomological Museum,
University of Kansas, Lawrence). Species names are listed alphabetically within
subfamilies, which are also in alphabetical sequence.
Checklist:
Subfamily ALEOCHARINAE
Abroteles beaumonti Casey, 1890 [Jacobson et al. *Aleochara (Xenochara) divergens Pace, 1990
(1986)] [Asenjo (2004)]
*Acrocyusa inca Pace, 1986 [Asenjo (2004)] Aleochara (Xenochara) lacertina Sharp, 1883 [Pace
*Acrotona etontii (Pace, 2011) [Pace (2011b), as (2009)]
Atheta (Acrotona)] *Aleochara (Xenochara) lacustris Bernhauer, 1908
*Acrotona loreticola (Pace, 2008) [Pace (2008), as [Asenjo (2004)]
Atheta (Acrotona)] [Aleochara (Xenochara) puberula Klug, 1832] [un-
[Acrotona parcior (Bernhauer, 1927)] [unrecorded recorded but probably in Peru]
but probably in Peru] Aleochara (Xenochara) taeniata Erichson, 1839
Adinopsis angusta (Sharp, 1883) [Maruyama and [Asenjo (2004)]
Kamezawa (2013)] Aleochara (Xenochara) verberans Erichson, 1839
Adinopsis myllaenoides (Kraatz, 1857) [Pace (2008)] [Asenjo et al. (2013)]
*Adinopsis peruviana Pace, 2008 [Pace (2008)] Aleochara (Xenochara) weiseri Bernhauer, 1921
Adinopsis pubescens Klimaszewski, 1982 [Klimas- [Asenjo et al. (2013)]
zewski and Frank (1992)] *Aleodorus peruvianus Pace, 1990 [Asenjo (2004)]
Aleochara (Aleochara) boliviana Pace, 1990 [New- Aleodorus scissus (Erichson, 1839) [Asenjo (2004)]
ton et al. (2005)] Amazoncharis aspera Pace, 1990 [Asenjo (2004)]
Aleochara (Aleochara) centralis Sharp, 1883 [Pla- Amazoncharis laminata Pace, 1990 [Hlavac et al.
vilstschikov (1929)] (2011)]
Aleochara (Aleochara) cribricollis Fairmaire & Amazonopora brooksi Ahn & Ashe, 1999 [Asenjo
Germain, 1862 [Solsky (1872)] (2004)]
Aleochara (Aleochara) curtula (Goeze, 1777) *Amazonopora lescheni Ahn & Ashe, 1999 [Asenjo
[Asenjo et al. (2013)] (2004)]
Aleochara (Aleochara) lata Gravenhorst, 1802 *Apalonia incaica (Pace, 1986) [Asenjo (2004)]
[Newton et al. (2005)] *Apalonia machupicchuensis (Pace, 1986) [Asenjo
[Aleochara (Aleochara) lustrica Say, 1832] [un- (2004)]
recorded but probably in Peru] *Apalonia quillabambana (Pace, 1986) [Asenjo
Aleochara (Aleochara) parvicollis Bernhauer, 1904 (2004)]
[Asenjo (2004)] *Apalonia tahuantinsuyiensis (Pace, 1986) [Asenjo
[Aleochara (Coprochara) bimaculata Gravenhorst, (2004)]
1802] [unrecorded but probably in Peru] *Arecaceopora couturieri Pace, 2011 [Pace (2011a)]
Aleochara (Coprochara) notula Erichson, 1839 *Arecaceopora delgadoi Pace, 2011 [Pace (2011a)]
[Asenjo et al. (2013)] *Arecaceopora perlifera Pace, 2011 [Pace (2011a)]
*Aleochara (Coprochara) pycnostichia Maus, 2000 *Atheta (Bessobia) ruinarum Pace, 1986 [Asenjo
[Maus (2000)] (2004)]
Aleochara (Coprochara) signaticollis Fairmaire & Atheta (Chaetida) peruviana Pace, 1986 [Asenjo
Germain, 1862 [Asenjo et al. (2013)] (2004)]
Aleochara (Coprochara) solieri Bernhauer & Atheta (Datomicra) brevis (Sharp, 1876) [Asenjo et
Scheerpeltz, 1926 [Moussallem et al. (2014)] al. (2013)]
[Atheta (Datomicra) conformis (Erichson, 1839)] *Gyrophaena peruminima Pace, 2008 [Pace (2008)]
[unrecorded but probably in Peru] *Gyrophaena peruviana Pace, 2008 [Pace (2008)]
*Atheta (Evaniodera) apicilutea Pace, 1986 [Asenjo *Gyrophaena westerduijni Pace, 2008 [Pace (2008)]
(2004)] *Habramicrada incaica Pace, 2007 [Pace (2007)]
*Atheta (Kladobrachydotina) couturieri Pace, 2011 *Habramicrada peruviana Pace, 2007 [Pace (2007)]
[Pace (2011a)] *Haplochara franziana Pace, 1986 [Asenjo (2004)]
*Atheta (Kladobrachydotina) sacsahuamanensis Pace, *Heliconandria gynochela Hanley, 2002 [Hanley
1986 [Asenjo (2004)] (2002)]
[Atheta (Pseudobessobia) atacamensis Pace, 1999] [Heterostiba castanea Pace, 1986] [Asenjo (2004)];
[unrecorded but probably in Peru] delete, type locality in Bolivia, not Peru
Atheta (Pseudobessobia) cuzcoensis Pace, 1986 [Heterostiba quechua Pace, 1986] [Asenjo (2004)];
[Asenjo (2004)] delete, type locality in Bolivia, not Peru
Atheta (Pseudobessobia) machupicchuensis Pace, *Heterostiba tahuantinsuyiensis Pace, 1986 [Asenjo
1986 [Asenjo (2004)] (2004)]
*Atheta (Pseudobessobia) mochicorum Pace, 1986 Holobus minutus (Cameron, 1931) [Asenjo et al.
[Asenjo (2004)] (2013)]
Atheta (Pseudobessobia) quillabambana Pace, 1986 [Holobus oviformis (Casey, 1894)] [Casanova and
[Asenjo (2004)] Korytkowski (1968)]; Peru record requires con-
*Bessoglossa peruviana Pace, 1986 [Asenjo (2004)] firmation
*Brachida improvisa Pace, 2008 [Pace (2008)] Holobus pigmaeus Solier, 1849 [Newton et al.
*Brundinia semisericea (Bernhauer, 1941) [Bern- (2005)]
hauer (1951)] *Hoplandria alboapicalis Pace, 2009 [Pace (2009)]
*Cousya peruviana Pace, 1986 [Asenjo (2004)] *Hoplandria angulosa Solsky, 1875 [Asenjo (2004)]
*Ctenopeuca heynei Bernhauer, 1915 [Asenjo Hoplandria globula Bernhauer & Scheerpeltz, 1926
(2004)] [Asenjo (2004)]; 5H. convexa Bernhauer, 1921
[Dactyloglossa inca Pace, 1986] [Asenjo (2004)]; *Hoplandria inca Pace, 1990 [Asenjo (2004)]
[delete, type locality in Bolivia, not Peru] Hoplandria lividula Solsky, 1877 [Asenjo (2004)]
[Diestota angulicollis Bernhauer, 1929] [Asenjo *Hoplandria mirabilis Bernhauer, 1904 [Asenjo
(2004)]; delete, nomen nudum in Plavilstschikov (2004)]
(1929) Hoplandria mochicorum Pace, 1990 [Asenjo (2004)]
*Diestota calida Bernhauer, 1920 [Asenjo (2004)] [Hydrosmecta pseudoimbellis (Pace, 1987)] [unre-
*Diestota chavin Pace, 1986 [Asenjo (2004)] corded but probably in Peru]
Diestota diffusa (Fauvel, 1901) [Asenjo et al. (2013)] [Ischnopoda fissicollis (Fairmaire & Germain,
Diestota ecuadorensis Pace, 1996 [Pace (2008)] 1862)] [unrecorded but probably in Peru]
Diestota laesicollis (Erichson, 1839) [Asenjo et al. *Lamprostiba etontii Pace, 2011 [Pace (2011b)]
(2013)] *Lamprostiba machupicchuensis Pace, 2001 [Asenjo
Diestota loretensis Pace, 2008 [Pace (2008)] (2004)]
Eburniola leucogaster Mann, 1923 [Kistner [Lamprostiba nazcaorum Pace, 1986] [Asenjo
(2006)] (2004)]; delete, type locality in Bolivia, not Peru
Ecitomorpha arachnoides Wasmann, 1889 [Asenjo Lamprostiba schwabei (Bernhauer, 1939) [Asenjo
et al. (2013)]; 5E. melanotica Mann, 1926 (2004)]
Ecitophya gracillima Mann, 1925 [Asenjo et al. *Leptandria ashei Hanley, 2003 [Hanley (2003)]
(2013)] *Leptanillophilus similis Holmgren, 1908 [Asenjo
Ecitophya rapaxae Mann, 1926 [Asenjo (2004)] (2004)]
Ecitophya simulans (Wasmann, 1889) [Asenjo et al. *Leptonia athaualpai Pace, 1986 [Asenjo (2004)]
(2013)]; 5E. bicolor Reichensperger, 1933 Leptonia comarapensis Pace, 1986 [Asenjo (2004),
[Ecitopora brevicornis Mann, 1925] [unrecorded but misspelled as comaparensis]
probably in Peru] *Leptonia decolorata Pace, 1990 [Pace (1990)]
[Eurydiotyphla franzi Pace, 1986] [Asenjo (2004)]; [Leptonia franzi Pace, 1986] [Asenjo (2004)]; delete,
delete, type locality in Bolivia, not Peru type locality in Bolivia, not Peru
Feluva pichinchaensis Pace, 2008 [Pace (2011a)] *Leptonia garavitensis Pace, 1986 [Asenjo (2004)];
Gansiella peruviana Pace, 2008 [Pace (2008)] 5L. inkarrii Pace, 1986
*Gnypeta etontii Pace, 2011 [Pace (2011b)] *Leptonia incaria Pace, 1986 [Asenjo (2004)]
*Gyrophaena bigranulata Pace, 2008 [Pace (2008)] *Leptonia moenium Pace, 1986 [Asenjo (2004)]
*Gyrophaena cornelli Pace, 2008 [Pace (2008)] *Leptonia nazcaensis Pace, 1986 [Asenjo (2004)]
*Gyrophaena perpunctum Pace, 2008 [Pace (2008)] *Leptonia nobilis Pace, 1986 [Asenjo (2004)]
*Gyrophaena perpusillina Pace, 2008 [Pace (2008)] *Leptonia peruviana Pace, 2011 [Pace (2011b)]
[Leptonia tahuantinsuyiensis Pace, 1986] [Asenjo *Peruusa antegilva Pace, 2008 [Pace (2008)]
(2004)]; delete, type locality in Bolivia, not Peru *Peruusa peruviana Pace, 2008 [Pace (2008)]
*Leptonia tiahuanacorum Pace, 1986 [Asenjo Phanerota baloghi (Pace, 1990) [Pace (2008), as
(2004)] Gyrophaena (Phanerota)]
*Macrogerodonia cuzcoensis Pace, 1986 [Asenjo Phanerota loretensis (Pace, 2008) [Pace (2008), as
(2004)] Gyrophaena (Phanerota)]
*Macrogerodonia magnicollis (Bernhauer, 1941) Phanerota perinsidiosa (Pace, 2008) [Pace (2008), as
[Pace (1990)] Gyrophaena (Phanerota)]
*Macrogerodonia nigrogibba Pace, 2008 [Pace *Phanerota perucastanea (Pace, 2008) [Pace (2008),
(2008)] as Gyrophaena (Phanerota)]
*Macrogerodonia peruviana (Bernhauer, 1908) *Phanerota peruflexa (Pace, 2008) [Pace (2008), as
[Asenjo (2004)]; M. peruviana longicollis Pace, Gyrophaena (Phanerota)]
1986 is a subspecies described from Peru. Phanerota vecta (Pace, 1990) [Pace (2008), as
Meronera cinctella (Motschulsky, 1858) [Asenjo et Gyrophaena (Phanerota)]
al. (2013)] Phoinixiusa ecuadorensis Pace, 2011 [Pace (2011a)]
*Mesoplandria loreticola Pace, 2008 [Pace (2008)] *Placusa loretensis Pace, 2008 [Pace (2008)]
Metadeinopsis brunnea Klimaszewski, 1979 [Asenjo Placusa peruviana Pace, 2008 [Pace (2008)]
et al. (2013)] *Placusa ruficollis Pace, 2008 [Pace (2008)]
Metadeinopsis longicornis (Sharp, 1876) [Asenjo et *Placusa westerduijni Pace, 2008 [Pace (2008)]
al. (2013)] *Platandria loretensis Pace, 2008 [Pace (2008)]
*Myllaena pallida Bernhauer, 1941 [Caron and Platandria peruviana (Bernhauer, 1941) [Asenjo
Klimazewski (2008)] (2004)]
Myrmedonota inca (Pace, 2001) [Asenjo (2004), as [Plesiomalota fasciatipennis (Fairmaire & Germain,
Apalonia]; to Myrmedonota, Eldredge (2012) 1862)] [unrecorded but probably in Peru]
*Myrmedonota myrmecophila (Pace, 2008) [Pace [Plesiomalota hispidula (Fauvel, 1866)] [unrecorded
(2008), as Apalonia]; to Myrmedonota, Eldredge but probably in Peru]
(2012) [Plesiomalota huascari Pace, 1986] [Asenjo (2004)];
delete, type locality in Bolivia, not Peru
*Myrmedonota westerduijni (Pace, 2008) [Pace
*Plesiomalota incaica Pace, 1986 [Asenjo (2004)]
(2008), as Apalonia]; to Myrmedonota, Eldredge
*Plesiomalota palmarum Pace, 2011 [Pace (2011a)]
(2012)
*Plesiomalota peruviana Pace, 1986 [Asenjo (2004)]
Nehemitropia lividipennis (Mannerheim, 1830)
Plesiomalota roedingeri (Bernhauer, 1941) [Asenjo
[Asenjo et al. (2013)]
(2004)]
[Neolara alboguttata (Erichson, 1839)] [unrecorded
*Plesiomalota tarapotensis Pace, 2007 [Pace (2007)]
but probably in Peru]
*Plesiomalota yucaicola Pace, 2008 [Pace (2008)]
[Oligota parva Kraatz, 1862] [unrecorded but
Polylobus bicolor (Solier, 1849) [Asenjo (2004)]
probably in Peru]
Pseudomniophila unidentata Pace, 1985 [Asenjo
*Orphnebius iquitosensis Pace, 2008 [Pace (2008)]
(2004)]
*Orphnebius loretensis Pace, 2008 [Pace (2008)]
*Pseudomyllaena franzi Pace, 1986 [Asenjo (2004)]
Orphnebius marginalis (Reichensperger, 1935) [Ja- Pseudophilotermes mcmahanae (Kistner, 1971)
cobson and Kistner (1998), Pace (2008)] [Kistner (2006)]
*Orphnebius peruvianus Pace, 1986 [Asenjo (2004)] Rothium evansi Ahn & Ashe, 1996 [Ahn and Ashe
*Orphnebius pungitius Solsky, 1875 [Asenjo (2004)] (1996)]
*Orphnebius satan Pace, 2009 [Pace (2009)] *Schistoglossa aymaraorum Pace, 1986 [Asenjo
*Orphnebius tarapotensis Pace, 2009 [Pace (2009)] (2004)]
*Orphnebius transversus Pace, 2009 [Pace (2009)] *Serikendusa peruviana Pace, 2008 [Pace (2008)]
*Ousipaliaglossa peruviana Pace, 2011 [Pace *Sharpidium peruensis Santiago-Jimenez, 2010
(2011a)] [Santiago-Jimenez (2010)]
[Parabainusa inca Pace, 2008] [Pace (2008)]; delete, *Sharpidium seeversi Santiago-Jimenez, 2010 [San-
nomen nudum tiago-Jimenez (2010)]
*Parasilusa inca Pace, 1986 [Asenjo (2004)] *Somasterochara peruviana Pace, 1986 [Asenjo
[Peliusa villosa Motschulsky, 1858] [unrecorded but (2004)]
probably in Peru] Spanioda pectoralis (Solier, 1849) [Asenjo (2004), as
Perinthus dudleyanus Casey, 1889 [Kistner (2004)] Calodera]
Perinthus major Seevers, 1957 [Kistner (2004)] *Termitogaster ashei Kistner, 2006 [Kistner (2006)]
*Perinthus praeterhirsutus Kistner, 2004 [Kistner *Termitogaster fontesi Kistner, 2006 [Kistner
(2004)] (2006)]
[Termitogaster impressicollis Seevers, 1937] [un- [Termitophya inornata Seevers, 1937] [unrecorded
recorded but probably in Peru] but probably in Peru]
*Termitogaster neoemersoni Kistner, 2006 [Kistner *Termitophya peruanus Kistner, 2006 [Kistner
(2006)] (2006)]
Termitogaster wasmanni (Holmgren, 1911) [Asenjo *Tetradonella amazoniensis Jacobson & Kistner,
(2004), as Xenogaster] 1998 [Jacobson and Kistner (1998)]
*Termitogaster wurlitzeri Kistner, 2006 [Kistner *Tetradonella breviapex Jacobson & Kistner, 1998
(2006)] [Jacobson and Kistner (1998)]
Termitohospes guianae Seevers, 1941 [Kistner Tetradonella latiarea Jacobson & Kistner, 1998
Termitomorpha hauchii (Seevers, 1946) [Kistner *Tetradonella peruensis Jacobson & Kistner, 1998
Termitomorpha meinerti Wasmann, 1894 [Asenjo et Thyreoxenus pulchellus Mann, 1923 [Asenjo et al.
al. (2013)] (2013)]
*Termitonannus calvitium Kistner, 2006 [Kistner *Tricholpochila armata Bernhauer, 1908 [Asenjo
(2006)] (2004)]; 5T. peruviana Bernhauer, 1908
*Termitonannus schmitzorum Kistner, 2006 [Kist- *Tricholpochila flavipennis Bernhauer, 1908
ner (2006)] [Asenjo (2004)]
Termitophya holmgreni Wasmann, 1911 [Kistner *Xenogaster lineis Kistner, 2006 [Kistner (2006)]
(2006)]; 5T. wasmanni Holmgren, 1911
Subfamily EUAESTHETINAE
*Edaphus angulosicollis Puthz, 2014 [Puthz (2014a)] Edaphus villac Puthz, 2007 [Puthz (2007b)]
Edaphus apo Puthz, 2007 [Puthz (2007b)] Octavius batesi (Sharp, 1876) [Herman (2001)]
*Edaphus atahualpa Puthz, 2014 [Puthz (2014a)] *Octavius peruanus Puthz, 2001 [Asenjo (2004)]
*Edaphus bufo Puthz, 1985 [Herman (2001)] Stenaesthetus amedegnatoae Orousset, 1990 [Her-
*Edaphus cinche Puthz, 2007 [Puthz (2007b)] man (2001)]
*Edaphus foraminosus Puthz, 2014 [Puthz (2014a)] Stenaesthetus castaneus Orousset, 1990 [Herman
*Edaphus foveolicollis Puthz, 2014 [Puthz (2014a)] (2001)]
*Edaphus lescheni Puthz, 2006 [Puthz (2006a)] Stenaesthetus illatus Sharp, 1876 [Herman (2001)]
*Edaphus longepilosus Puthz, 2006 [Puthz (2006a)] [Stenaesthetus immarginatus (Erichson, 1840)]
*Edaphus peruanus Puthz, 1973 [Herman (2001)] [Asenjo et al. (2013), but Peru with ?]
*Edaphus poulaini Orousset, 1986 [Herman (2001)] *Stenaesthetus peruanus Orousset, 1990 [Herman
Edaphus rishwani Makhan, 1995 [Asenjo et al. (2001)]
(2013)] Stenaesthetus puthzi Orousset, 1990 [Herman
Edaphus siphonifer Puthz, 2007 [Puthz (2007b)] (2001)]
*Edaphus variipunctus Puthz, 2014 [Puthz (2014a)] Tamotus hirsutus Puthz, 2002 [Asenjo (2004)]
Edaphus ventricula Puthz, 2006 [Puthz (2006c)] Tamotus rugicollis Puthz, 2002 [Asenjo (2004)]
Subfamily MEGALOPSIDIINAE
[Megalopinus adjecticius Puthz, 2012] [unrecorded Megalopinus duplicatus Puthz, 1994 [Herman
but probably in Peru] (2001)]
Megalopinus aimara Puthz, 1994 [Herman (2001)] Megalopinus elegantior Puthz, 1994 [Herman
*Megalopinus amauta Puthz, 2012 [Puthz (2012)] (2001)]
Megalopinus bicuspidiventris Puthz, 2012 [Puthz Megalopinus endofalcatus Puthz, 2012 [Puthz
(2012)] (2012)]
Megalopinus bolivianus (Bernhauer, 1909) [Puthz [Megalopinus erichsoni (Benick, 1916)] [unrecorded
(2012)] but probably in Peru]
Megalopinus brunneiventris Puthz, 2012 [Puthz (2012)] Megalopinus fascifer Puthz, 2012 [Puthz (2012)]
*Megalopinus brunneus Puthz, 2012 [Puthz (2012)] Megalopinus franzi Puthz, 1994 [Herman (2001)]
*Megalopinus brzoskai Puthz, 2012 [Puthz (2012)] Megalopinus gestroi (Bernhauer, 1909) [Herman
*Megalopinus cacique Puthz, 2012 [Puthz (2012)] (2001)]
Megalopinus chinan Puthz, 1994 [Herman (2001)] Megalopinus glabripennis (Benick, 1952) [Puthz
Megalopinus coya Puthz, 2012 [Puthz (2012)] (2012)]
Megalopinus cruciger (Sharp, 1886) [Asenjo et al. *Megalopinus harpaginiventris Puthz, 2012 [Puthz
(2013)] (2012)]
[Megalopinus humerigemmatus Puthz, 2012] [un- *Megalopinus quadrigutta Puthz, 2012 [Puthz
recorded but probably in Peru] (2012)]
Megalopinus humerinaevulus Puthz, 2012 [Puthz Megalopinus quadrimaculatus (Bernhauer, 1909)
(2012)] [Puthz (2012)]
Megalopinus inka Puthz, 1994 [Herman (2001)] Megalopinus quadrimaculosus Puthz, 2012 [Puthz
Megalopinus laevipennis (Bernhauer, 1909) [Puthz (2012)]
(2012)] Megalopinus robustus (Motschulsky, 1857) [Puthz
*Megalopinus lescheni Puthz, 1994 [Herman (2001)] (2012)]
Megalopinus longicornis Puthz, 2012 [Puthz (2012)] Megalopinus rossi Puthz, 2012 [Puthz (2012)]
Megalopinus minutus Puthz, 2012 [Puthz (2012)] [Megalopinus rufipectus Puthz, 2012] [unrecorded
Megalopinus moderatus Puthz, 1994 [Herman but probably in Peru]
(2001)] Megalopinus schlingeri Puthz, 2012 [Puthz (2012)]
Megalopinus nigellus Puthz, 2012 [Puthz (2012)] Megalopinus schuhi Puthz, 2012 [Puthz (2012)]
*Megalopinus nigrificatus Puthz, 1989 [Herman *Megalopinus serratipenis Puthz, 2012 [Puthz
(2001)] (2012)]
Megalopinus nobilis Puthz, 2012 [Puthz (2012)] Megalopinus speculifrons Puthz, 2012 [Puthz
Megalopinus obliquestriatus (Bernhauer, 1920) (2012)]
[Puthz (2012)] Megalopinus spiniventris (Benick, 1926) [Puthz
Megalopinus ogloblini (Bernhauer, 1933) [Herman (2012)]
(2001)] Megalopinus spinosus (Sharp, 1876) [Puthz (2012)];
*Megalopinus ornatus (Waterhouse, 1883) [Herman 5M. impressus (Sharp, 1876)
(2001)] [Megalopinus suavis Puthz, 2012] [unrecorded but
*Megalopinus pakarina Puthz, 2012 [Puthz (2012)] probably in Peru]
*Megalopinus peruanus Puthz, 2012 [Puthz (2012)] Megalopinus suavivestis Puthz, 2012 [Puthz (2012)]
Megalopinus pulcher Puthz, 1989 [Herman (2001)] Megalopinus tubericollis Puthz, 1994 [Herman
[Megalopinus punctatus (Erichson, 1840)] [Asenjo et (2001)]
al. (2013)]; Peru unconfirmed, Puthz (2012) *Megalopinus vicuna Puthz, 2012 [Puthz (2012)]
Subfamily MICROPEPLINAE
Peplomicrus uyttenboorgarti (Bernhauer, 1928) [Herman (2001)]
*Peplomicrus watrousi Campbell, 1986 [Herman (2001)]
Subfamily OMALIINAE
*Phyllodrepa peruviana Bernhauer, 1941 [Herman (2001)]
Subfamily OSORIINAE
*Allotrochus minor Irmler, 2000 [Asenjo (2004)] Espeson subtilis Bernhauer, 1910 [Bernhauer
Aneucamptus crassus (Sharp, 1876) [Herman (1951)]
(2001)] Espeson titschacki Bernhauer, 1941 [Herman
Aneucamptus excisicollis (Motschulsky, 1860) (2001)]
[Asenjo (2004)] Holotrochopsis danoffburgi (Irmler, 2001) [Irmler
*Aneucamptus geroi Irmler, 2001 [Asenjo (2004)] (2015)]
*Arborilispinus mirabilis Irmler, 2010 [Irmler Holotrochopsis pubescens (Sharp, 1876) [Irmler
(2010b)] (2015)]
Clavilispinus exiguus (Erichson, 1840) [Irmler Holotrochopsis setifera (Irmler, 2010 [Irmler (2015)]
(2003a)] *Holotrochus ashei Irmler, 2005 [Irmler (2005a)]
Clavilispinus junkii Irmler, 2003 [Irmler (2003a)] Holotrochus centralis Sharp, 1887 [Asenjo et al.
Clavilispinus megacephalus (Fauvel, 1864) [Herman (2013)]
(2001)] *Holotrochus hanagarthi Irmler, 1982 [Herman
(2001)]
Clavilispinus minutus (Sharp, 1887) [Irmler (2003a)]
Holotrochus lescheni Irmler, 2005 [Irmler (2005a)]
Clavilispinus politus (Sharp, 1887) [Herman (2001)]
*Holotrochus lineatus Irmler, 1982 [Herman (2001)]
Dirocephalus andinus Irmler, 2005 [Irmler (2005d)]
*Holotrochus loretoi Irmler, 2005 [Irmler (2005a)]
*Dirocephalus politus Irmler, 2005 [Irmler (2005d)]
*Holotrochus neotropicus Irmler, 1982 [Herman
*Dirocephalus rugosus Irmler, 2005 [Irmler (2005d)] (2001)]
Eleusis interrupta (Erichson, 1840) [Newton et al. *Holotrochus peruvianus Bernhauer, 1941 [Herman
(2005)] (2001)]
Holotrochus poundi Blackwelder, 1943 [Herman Lispinus lunaris Irmler, 1994 [Irmler (2009)]
(2001)] *Lispinus minimus Irmler, 2009 [Irmler (2009)]
*Holotrochus pseudoleticiae Irmler, 2013 [Irmler Lispinus newtoni Irmler, 1994 [Irmler (2009)]
(2013)] *Lispinus peruanus Irmler, 2009 [Irmler (2009)]
*Holotrochus pumilus Irmler, 1982 [Herman (2001)] Lispinus quadripunctulus Fauvel, 1865 [Irmler
Holotrochus syntheticus Sharp, 1876 [Herman (2001)] (2009)]
Leptochirus arcifer Fauvel, 1902 [Herman (2001)] Lispinus sinuatocollis Bernhauer, 1942 [Irmler
Leptochirus brunneoniger Perty, 1830 [Asenjo et al. (2009)]; 5L. costaricensis Irmler, 1994
(2013)] [Lispinus sobrinus Fauvel, 1864] [unrecorded but
*Leptochirus consentaneus Wendeler, 1957 [Herman probably in Peru]
(2001)] Lispinus socialis Irmler, 1994 [Irmler (2009)]
Leptochirus eximius Wendeler, 1957 [Herman *Lispinus speciosus Irmler, 2009 [Irmler (2009)]
(2001)] Lispinus striola Erichson, 1840 [Herman (2001)]
*Leptochirus giganteus Wendeler, 1927 [Herman Lispinus tardus Sharp, 1887 [Irmler (2009)]
(2001)] *Mimogonia andina Irmler, 2005 [Irmler (2005b)]
Leptochirus helleri Bernhauer, 1903 [Herman (2001)] *Mimogonia antennata Irmler, 1981 [Herman
Leptochirus iconnicoffi Bernhauer, 1917 [Herman (2001)]
(2001)] *Mimogonia brunnea Irmler, 1981 [Herman (2001)]
Leptochirus incertus Bernhauer, 1903 [Herman *Mimogonia hanagarthi Irmler, 2007 [Irmler
(2001)] (2007a)]
Leptochirus kolbei Bernhauer, 1903 [Asenjo (2004)] *Mimogonia huggerti Irmler, 2007 [Irmler (2007a)]
[Leptochirus laeviventris Fauvel, 1902] [unrecorded *Mimogonia longoelytrata Irmler, 2005 [Irmler
but probably in Peru] (2005b)]
[Leptochirus latro Sharp, 1876] [unrecorded but *Mimogonia pumilia Irmler, 1981 [Herman (2001)]
probably in Peru] Mimogonia similis Irmler, 1981 [Irmler (2007a)]
Leptochirus maxillosus (Fabricius, 1801) [Herman *Mimogonia subopaca Irmler, 1981 [Herman (2001)]
(2001)]; 5L. anthracinus Fauvel, 1902, 5L. Mimogonia tricolor Irmler, 1981 [FMNH collection
collaris Fauvel, 1902 (det. A. Newton)]
[Leptochirus mery Asenjo & Ribeiro-Costa, 2013] Nacaeus bicolor (Sharp, 1887) [Irmler (2003b)]
[unrecorded but probably in Peru] Nacaeus brasiliensis Irmler, 2003 [Irmler (2003b)]
Leptochirus peruvianus Bernhauer, 1917 [Herman Nacaeus claviger (Cameron, 1913) [Irmler (2003b)]
(2001)] Nacaeus cordiger Irmler, 2003 [Irmler (2003b)]
Leptochirus proteus Fauvel, 1864 [Herman (2001)] Nacaeus flavipennis (Fauvel, 1864) [Irmler (2003b)]
Leptochirus scoriaceus Germar, 1824 [Asenjo et al. *Nacaeus flavoelytratus Irmler, 2006 [Irmler (2006)]
(2013)] [Nacaeus funebris (Bernhauer, 1921)] [unrecorded
Leptochirus tenuis Bernhauer, 1903 [Bernhauer but probably in Peru]
(1903)]; type locality Chanchomayo in Peru, *Nacaeus inkae Irmler, 2003 [Irmler (2003b)]
not Bolivia Nacaeus nigrifrons (Fauvel, 1863) [Asenjo et al. (2013)]
Lispinus attenuatus Erichson, 1840 [Herman (2001)] Nacaeus opacus (Fauvel, 1864) [Irmler (2003b)]
*Lispinus blandus Irmler, 2009 [Irmler (2009)] Nacaeus peruvianus Irmler, 2003 [Irmler (2003b)]
Lispinus bolivianus Bernhauer, 1929 [Irmler (2009)]; Nacaeus planellus (Sharp, 1887) [Irmler (2003b)]
5L. opacipennis Bernhauer, 1921 Nacaeus simplex (Sharp, 1876) [Irmler (2003b)]
Lispinus canalis Irmler, 1994 [FMNH collection Nacaeus spegazzinii (Bernhauer, 1933) [Irmler
(Irmler labeled paratype)] (2003b)]
Lispinus catena Sharp, 1876 [Irmler (2009)] *Neolosus obscurus Irmler, 1999 [Herman (2001)]
Lispinus cordilliensis Irmler, 1994 [Newton et al. Oryssoma schwarzi Notman, 1925 [Irmler (2010c)]
(2005)] *Osoriellus acutus Irmler, 2014 [Irmler (2014)]
Lispinus fungicola Irmler, 2001 [Asenjo (2004)] *Osoriellus adustus Irmler, 2014 [Irmler (2014)]
Lispinus granadensis Fauvel, 1864 [Solsky (1872)] *Osoriellus anceps Irmler, 2014 [Irmler (2014)]
[Lispinus insularis Fauvel, 1863] [Herman (2001)]; *Osoriellus bicolor Irmler, 2014 [Irmler (2014)]
delete, Peru in error, teste Irmler (2000) *Osoriellus bicornis Irmler, 2014 [Irmler (2014)]
Lispinus laeviusculus Bernhauer, 1910 [Herman Osoriellus boliviensis Irmler, 2014 [Irmler (2014)]
(2001)]; 5L. laevigatus Bernhauer, 1906 Osoriellus carinicollis (Notman, 1925) [Irmler
Lispinus laticollis Erichson, 1840 [Irmler (2009)] (2014)]
*Lispinus lescheni Irmler, 2001 [Asenjo (2004)] *Osoriellus digitatus Irmler, 2014 [Irmler (2014)]
Lispinus linearis Erichson, 1840 [Irmler (2009)] *Osoriellus diversus Irmler, 2014 [Irmler (2014)]
Lispinus listenbarthi Irmler, 1994 [Asenjo (2004)] *Osoriellus fumarius Irmler, 2014 [Irmler (2014)]
Osoriellus hanagarthi Irmler, 2014 [Irmler (2014)] *Osorius pusillus Irmler, 2010 [Irmler (2010c)]
*Osoriellus infuscatus Irmler, 2014 [Irmler (2014)] Osorius stipes Sharp, 1876 [Asenjo et al. (2013)]
*Osoriellus lescheni Irmler, 2014 [Irmler (2014)] *Osorius verhaaghi Irmler, 2010 [Irmler (2010c)]
*Osoriellus loreti Irmler, 2014 [Irmler (2014)] Ouloglene barberi Notman, 1925 [Irmler (2010c)]
Osoriellus luteus Irmler, 2014 [Irmler (2014)] Pseudespeson nitens (Fauvel, 1902) [Asenjo et al.
*Osoriellus majusculus Irmler, 2014 [Irmler (2014)] (2013)]
Osoriellus melancholicus Irmler, 2014 [Irmler (2014)] *Quadricephalus obstrusus Irmler, 2005 [Irmler
Osoriellus obtusicollis Irmler, 2014 [Irmler (2014)] (2005d)]
Osoriellus opticus Irmler, 2014 [Irmler (2014)] Tannea amazonica Irmler, 2003 [Irmler (2003b)]
*Osoriellus rubripennis Irmler, 2014 [Irmler (2014)] Tannea brevicollis (Fauvel, 1865) [Asenjo et al.
*Osoriellus rufescens Irmler, 2014 [Irmler (2014)] (2013)]
*Osoriellus rugipennis (Bernhauer, 1941) [Irmler *Tannea fulgens (Bernhauer, 1941) [Herman
(2014)] (2001), Irmler (2003b)]
*Osoriellus seriatus Irmler, 2014 [Irmler (2014)] Tannea laticeps Irmler, 2005 [Irmler (2005c)]
*Osoriellus setifer Irmler, 2014 [Irmler (2014)] Tannea latinota Irmler, 2003 [Irmler (2003b)]
*Osoriellus sinuatoangularis Irmler, 2014 [Irmler *Tannea longula Irmler, 2005 [Irmler (2005c)]
(2014)] Tannea parallelonota Irmler, 2003] [Irmler (2007b)]
*Osoriellus spinosus Irmler, 2014 [Irmler (2014)] Tannea picata Irmler, 2003 [Irmler (2003b)]
Osoriellus sticticus Irmler, 2014 [Irmler (2014)] Tannea salasi Irmler, 2003 [Irmler (2003b)]
*Osoriellus subsinuatus Irmler, 2014 [Irmler (2014)] Tannea tenella (Erichson, 1840) [Irmler (2003b)]
Osoriellus trinitatus (Blackwelder, 1943) [Irmler Tannea varablancae Irmler, 2003] [Irmler (2007b)]
(2014)] Thoracophorus denticollis (Erichson, 1840) [Herman
Osoriellus verhaaghi Irmler, 2014 [Irmler (2014)] (2001)]
Osoriellus weberi (Blackwelder, 1943) [Irmler Thoracophorus guadalupensis Cameron, 1913 [Her-
(2014)] man (2001)]
*Osoriosetus hanagarthi Irmler, 2015 [Irmler (2015)] *Thoracophorus peruvianus Bernhauer, 1941 [Her-
Osorius alternans Bernhauer, 1920 [Irmler (2010c)] man (2001)]
[Osorius brasiliensis Guerin-Meneville, 1829] [un- Thoracophorus sallaei Sharp, 1887 [Herman (2001)]
recorded but probably in Peru] Thoracophorus silvaticus Irmler, 2010 [Irmler
Osorius columbinus Bernhauer, 1920 [Irmler (2010a)]
(2010c)] Thoracophorus susannae Irmler, 2001 [Asenjo
Osorius laevicollis Bernhauer, 1908 [Irmler (2010c)] (2004)]
Osorius nitens Sharp, 1876 [Irmler (2010c)]; 5O. *Tumboecus verhaaghi Irmler, 2015 [Irmler (2015)]
latimargo Bernhauer, 1908 *Verhaaghiella becki (Irmler, 2001) [Irmler (2005d)]
Osorius peruvianus Bernhauer, 1908 [Irmler (2010c)]
Subfamily OXYTELINAE
Anotylus dentifrons (Erichson, 1840) [Asenjo et al. Parosus longipennis Makranczy, 2014 [Makranczy
(2013)] (2014)]
Anotylus insignitus Gravenhorst, 1806 [Asenjo (2004)] Parosus major Makranczy, 2014 [Makranczy (2014)]
Anotylus peruvianus (Bernhauer, 1941) [Herman *Parosus simplex Makranczy, 2014 [Makranczy
(2001)] (2014)]
Bledius pumilio Erichson, 1840 [Herman (2001)] *Parosus thayerae Makranczy, 2014 [Makranczy
[Carpelimus fulvipes (Erichson, 1840)] [unrecorded (2014)]
but probably in Peru] Platystethus spiculus Erichson, 1840 [Herman (2001)]
*Carpelimus opacellus (Bernhauer, 1941) [Herman [Thinodromus binotatus (Bernhauer, 1922)] [unre-
(2001)] corded but probably in Peru]
*Oxytelus jelskii Fauvel, 1905 [Herman (2001)] Thinodromus signatus (Erichson, 1834) [Herman
Oxytelus speculum Fauvel, 1904 [Herman (2001)] (2001)]
*Parosus longicornis Makranczy, 2014 [Makranczy
(2014)]
Subfamily PAEDERINAE
*Baryopsis glabra Assing, 2012 [Assing *Baryopsis montivagans (Bernhauer, 1906) [Assing
(2012a)] (2012a)]
*Baryopsis ingens Assing, 2012 [Assing (2012a)] *Cylindroxystus furvus Herman, 1991 [Herman
*Baryopsis minor Assing, 2012 [Assing (2012a)] (1991)]
*Cylindroxystus messus Herman, 1991 [Herman Paederus bolivianus Bernhauer, 1905 [Asenjo
(1991)] (2004)]
Dacnochilus bilobatus Jimenez-Sanchez & Galian, Paederus brasiliensis Erichson, 1840 [Asenjo et al.
2013 [Jimenez-Sanchez and Galian (2013)] (2013)]
Dacnochilus grandioculus Jimenez-Sanchez & Galian, Paederus columbinus Laporte, 1835 [Asenjo et al.
2013 [Jimenez-Sanchez and Galian (2013)] (2013)]
Dacnochilus paganus (Sharp, 1876) [Jimenez-Sanchez Paederus curticollis Bernhauer, 1905 [Asenjo
and Galian (2013)] (2004)]
Dibelonetes biplagiatus Sahlberg, 1847 [Asenjo [Paederus ferus Erichson, 1840] [unrecorded but
(2004)] probably in Peru]
*Echiaster roedingeri Bernhauer, 1941 [Bernhauer *Paederus gigas Bernhauer, 1905 [Asenjo (2004)]
(1951)] Paederus magniceps Bernhauer, 1905 [Asenjo (2004)]
*Ecitonides volans Assing, 2012 [Assing (2012b)] Paederus mutans Sharp, 1876 [Asenjo (2004)]
Eupaederus cyanipennis (Guerin-Meneville, 1830) Paederus ornaticornis Sharp, 1891 [Ojeda Pena
[Asenjo (2004), as Paederus]; to Eupaederus, (1969), Guillen (1989)]; 5P. irritans Chapin,
Scheerpeltz (1957) 1926
Lathropinus haenschi Bernhauer, 1918 [Bernhauer Paederus ovaliceps Bernhauer, 1905 [Asenjo (2004)]
(1951)]; 5L. peruvianus Bernhauer, 1941 *Paederus peruanus Wendeler, 1931 [Asenjo (2004)]
Lithocharis infuscata Erichson, 1840 [Asenjo Paederus subnitidus Bernhauer, 1905 [Asenjo
(2004)] (2004)]
Lithocharis mendax Sharp, 1886 [Asenjo (2004)] [Paederus tempestivus Erichson, 1840] [unrecorded
*Lithocharis nitida Erichson, 1840 [Asenjo but probably in Peru]
(2004)] Paederus tridens Sharp, 1876 [Asenjo (2004)]
Lithocharis vilis Kraatz, 1859 [Asenjo et al. (2013)] Palaminus brevicollis Sahlberg, 1847 [Asenjo et al.
*Medon semiopacus Bernhauer, 1941 [Bernhauer (2013)]
(1951)] Palaminus gracilis Sharp, 1876 [Asenjo et al.
*Monista maculipennis Bernhauer, 1941 [Bernhauer (2013)]
(1951)] Palaminus pallidus Sahlberg, 1847 [Asenjo et al.
*Neolindus amazonicus Irmler, 1981 [Herman (2013)]
(1991)] *Palaminus sharpi Bernhauer, 1905 [Asenjo (2004)]
*Neolindus cephalochymus Herman, 1991 [Herman Palaminus variabilis Erichson, 1840 [Asenjo et al.
(1991)] (2013)]
*Neolindus hanagarthi Irmler, 1981 [Herman Palaminus vittiger Bernhauer, 1917 [Asenjo (2004)]
(1991)] Rugilus chilensis Solier, 1849 [Scheerpeltz (1972)]
*Neolindus luciamans Assing, 2012 [Assing (2012c)] *Rugilus peruvianus (Bernhauer, 1921) [Asenjo
*Neolindus peruvianus Irmler, 1981 [Herman (2004)]
(1991)] Rugilus rufipennis (Bernhauer, 1921) [Asenjo
*Neolindus punctiventris Irmler, 1981 [Herman (2004)]
(1991)] Sciocharis oculata (Sharp, 1876) [Asenjo et al. (2013)]
*Neolindus verhaaghi Irmler, 2011 [Irmler *Scioporus peruvianus (Bernhauer, 1941) [Bern-
(2011)] hauer (1951)]
Ochthephilum bugnioni (Fauvel, 1901) [Asenjo et al. *Scopaeus grandipennis Bernhauer, 1941 [Bern-
(2013)] hauer (1951)]
*Ochthephilum diehli (Bernhauer, 1941) [Bernhauer Suniotrichus sordidus Sharp, 1886 [Asenjo et al.
(1951)] (2013)]
Ochthephilum disjunctum (Lynch, 1884) [Scheer- Sunius debilicornis (Wollaston, 1857) [Asenjo (2004)]
peltz (1972)] *Taenodema banghaasi Bernhauer, 1908 [Asenjo
*Ochthephilum flavipes (Erichson, 1847) [Asenjo (2004)]
(2004)] *Taenodema elegans Solsky, 1871 [Asenjo (2004)]
Ochthephilum longicorne (Sharp, 1876) [Asenjo *Taenodema hemichlora Solsky, 1875 [Asenjo
(2004)] (2004)]
*Oedichirus hamatus Herman, 2013 [Herman *Taenodema peruviana Bernhauer, 1908 [Asenjo
(2013)] (2004)]
Paederus amazonicus Sharp, 1876 [Vieira et al. *Thinocharis palcoensis Bernhauer, 1941 [Bernhauer
(2014)] (1951)]
*Paederus andinus Wendeler, 1931 [Asenjo (2004)] *Thinocharis peruviana Bernhauer, 1941 [Bernhauer
*Paederus baeri Bernhauer, 1921 [Asenjo (2004)] (1951)]
Subfamily PIESTINAE
Hypotelus laevis Solsky, 1872 [Caron et al. (2012)] Piestus pygmaeus Laporte, 1835 [Caron et al.
Piestus angularis Fauvel, 1864 [Caron et al. (2012)] (2012)]
Piestus aper Sharp, 1876 [Caron et al. (2012)] Piestus rugosus Sharp, 1876 [Caron et al. (2012)]
Piestus bicornis (Olivier, 1811) [Caron et al. (2012)]; Piestus spinosus (Fabricius, 1801) [Caron et al.
5P. oxytelinus of authors (misidentification) (2012)]
Piestus buquetii Fauvel, 1864 [Caron et al. (2012)] Piestus sulcatus Gravenhorst, 1806 [Caron et al.
Piestus capricornis Laporte, 1835 [Caron et al. (2012)] (2012)]
Piestus chiriquensis Sharp, 1887 [Caron et al. (2012)] Piestus surrufus Caron et al., 2012 [Caron et al.
*Piestus elegans Caron et al., 2012 [Caron et al. (2012)]
(2012)] Piestus termitis Caron et al., 2012 [Caron et al.
Piestus lacordairei Laporte, 1835 [Caron et al. (2012)] (2012)]
Piestus minutus Erichson, 1840 [Caron et al. Piestus validus Sharp, 1876 [Caron et al. (2012)]
(2012)]; 5P. nigrator Fauvel, 1902 Piestus zischkai Scheerpeltz, 1951 [Caron et al.
Piestus pennicornis Fauvel, 1864 [Caron et al. (2012)]
(2012)]
Subfamily PROTEININAE
*Megarthrus machu Cuccodoro, 2011 [Cuccodoro (2011)]
Subfamily PSELAPHINAE
*Arthmius elegantulus Fletcher, 1928 [Asenjo (2004)] *Hamotus brevimarginatus Schaufuss, 1887 [Asenjo
*Arthmius laevipennis Fletcher, 1928 [Asenjo (2004)] (2004)]
*Arthmius magnus Fletcher, 1928 [Asenjo (2004)] *Jubogaster towai Parker & Maruyama, 2013
*Arthmius peruvianus (Raffray, 1882) [Asenjo (2004)] [Parker and Maruyama (2013)]
*Bryaxis proportionalis Schaufuss, 1887 [Schau- *Mayetia peruana Orousset, 1995 [Orousset (1995)]
fuss (1887), type locality in Peru (not Chile as *Metopioxys hamatus Raffray, 1897 [Asenjo et al.
subsequently reported)]; not Bryaxis but genus (2013)]
uncertain, teste D. S. Chandler (pers. comm. to Metopioxys reichei (Schaufuss, 1872) [Asenjo et al.
A. Newton) (2013)]
*Bythinoplectus vaucheri Comellini, 1985 [Asenjo Metopioxys tavakiliani Comellini, 1983 [Asenjo et
(2004)] al. (2013)]
*Decarthron lamellipes Fletcher, 1928 [Asenjo Metopioxys tricuspidatus Raffray, 1897 [Asenjo et
(2004)] al. (2013)]
*Eutrichites skelleyi Carlton, 1996 [Asenjo (2004)] Oxarthrius simplex (Raffray, 1897) [Asenjo (2004)]
*Fustiger testudineus Schaufuss, 1882 [Asenjo *Reichenbachia femoralis Fletcher, 1928 [Asenjo
(2004)] (2004)]
*Goniacerus franzi Comellini, 1990 [Comellini *Reichenbachia limpida Fletcher, 1928 [Asenjo
(1990)] (2004)]
Goniacerus setifer (Schaufuss, 1877) [Asenjo et al. *Rhytus oberthueri (Raffray, 1891) [Asenjo et al.
(2013)] (2013)]
Subfamily PSEUDOPSINAE
[Pseudopsis columbica Fauvel, 1865] [Herman (2001)]; Peru probably in error, teste Asenjo and Ribeiro-
Costa (2010)
*Pseudopsis monica Asenjo & Ribeiro-Costa, 2010 [Asenjo and Ribeiro-Costa (2010)]
Subfamily SCAPHIDIINAE
[Cyparium championi Matthews, 1888] [unrecorded Scaphidium fasciatomaculatum Oberthur, 1884
but probably in Peru] [Fierros-Lopez (2006)]
*Cyparium inclinans Kirsch, 1873 [Asenjo (2004)] *Scaphidium vittipenne Oberthur, 1884 [Lobl
*Cyparium mathani Oberthur, 1884 [Asenjo (2004)] (1997)]
*Cyparium nigronotatum Pic, 1931 [Asenjo (2004)] Scaphisoma tropicum Kirsch, 1873 [Asenjo
*Cyparium peruvianum Pic, 1947 [Lobl (1997)] (2004)]
Subfamily SCYDMAENINAE
*Alloraphes dentatus Jaoszynski, 2013 [Jaoszynski *Euconnus (Napoconnus) fulgurans Franz, 1980
(2013b)] [Franz (1980)]
*Alloraphes peruanus Franz, 1980 [Franz (1980)] *Euconnus (Napoconnus) santanae Franz, 1980
*Archiconnus huallaganus Franz, 1980 [Franz [Franz (1980)]
(1980)] *Euconnus (Napoconnus) yapacarii Franz, 1980
*Euconnomorphus pilosissimus Franz, 1980 [Franz [Franz (1980)]
(1980)] *Euconnus (Pycnophus) paludicola Franz, 1980
*Euconnus angustus Franz, 1980 [Franz (1980)] [Franz (1980)]
*Euconnus compactus Franz, 1980 [Franz (1980)] *Euconnus (Tetramelus) alloscydmaenoides Franz,
*Euconnus differens Franz, 1980 [Franz (1980)] 1967 [Franz (1980)]
*Euconnus garavitoi Franz, 1967 [Franz (1980)] *Euconnus (Tetramelus) cuzcoanus Franz, 1967
*Euconnus gigas Franz, 1980 [Franz (1980)] [Franz (1980)]
*Euconnus loretoi Franz, 1989 [Franz (1989)] *Euconnus (Tetramelus) cuzcoi Franz, 1967 [Franz
(1980)]
*Euconnus minutus Franz, 1980 [Franz (1980)]
*Euconnus (Tetramelus) garavitanus Franz, 1967
*Euconnus nubigena Franz, 1980 [Franz (1980)] [Franz (1980)]
*Euconnus pachecoi Franz, 1967 [Franz (1980)]
*Euconnus (Tetramelus) peruanus Franz, 1967
*Euconnus pilosipennis Franz, 1980 [Franz (1980)] [Franz (1980)]
*Euconnus pilosissimus Franz, 1980 [Franz (1980)] *Euconnus (Tetramelus) scydmaenoides Franz,
*Euconnus princeps Franz, 1967 [Franz (1980)] 1967 [Franz (1980)]
*Euconnus quillabambae Franz, 1967 [Franz (1980)] *Euconnus (Tetramelus) ventrecarinatus Franz,
*Euconnus samiriae Franz, 1989 [Franz (1989)] 1980 [Franz (1980)]
*Euconnus samirianus Franz, 1989 [Franz (1989)] *Homoconnus novus Franz, 1980 [Franz (1980)]
*Euconnus tarapotanus Franz, 1980 [Franz (1980)] *Microscydmus huallagae Franz, 1980 [Franz
*Euconnus vaucheri Franz, 1989 [Franz (1989)] (1980)]
*Euconnus (Napochus) alloquillabambanus Franz, *Neuraphanax mirificus Franz, 1967 [Franz (1980)]
1967 [Franz (1980)] *Parapseudoconnus fraudulentus (Franz, 1980) [Ja-
[Euconnus (Napochus) campestris (Schaufuss, oszynski (2013a)]
1866)] [unrecorded but probably in Peru] *Parapseudoconnus monticola (Franz, 1980) [Jaos-
Euconnus (Napochus) cirnelimai Franz, 1967 zynski (2013a)]
[Asenjo et al. (2013)] *Parapseudoconnus simulator (Franz, 1980) [Jaos-
*Euconnus (Napochus) consimilis Franz, 1980 zynski (2013a)]
[Franz (1980)] *Protandroconnus peruensis Franz, 1989 [Franz
*Euconnus (Napochus) huallagae Franz, 1980 (1989)]
[Franz (1980)] *Protoconnus andicola Franz, 1967 [Franz (1980)]
*Euconnus (Napochus) juanguerrae Franz, 1980 *Protoconnus minutissimus Franz, 1980 [Franz
[Franz (1980)] (1980)]
*Euconnus (Napochus) koepkei Franz, 1967 [Franz *Protoconnus minutus Franz, 1980 [Franz (1980)]
(1980)] *Protoconnus peruensis Franz, 1967 [Franz (1980)]
*Euconnus (Napochus) parashanusii Franz, 1980 *Pseudocephennium peruvianum Jaoszynski, 2012
[Franz (1980)] [Jaoszynski (2012)]
*Euconnus (Napochus) potreroi Franz, 1967 [Franz
*Pseudoraphes huallagae Franz, 1980 [Franz (1980)]
(1980)] *Pseudoraphes minor Franz, 1980 [Franz (1980)]
*Euconnus (Napochus) punctipennis Franz, 1980
*Sciacharis garavitoensis Franz, 1997 [Franz (1997)]
[Franz (1980)]
*Scydmaenus garavitoensis Franz, 1980 [Franz
*Euconnus (Napochus) quillabambanus Franz, 1967
(1980)]; S. quillabambanus Franz, 1980 (in key,
[Franz (1980)]
p. 71) is an apparent lapsus calami for S.
*Euconnus (Napochus) rufus Franz, 1980 [Franz
garavitoensis (pp. 58, 59, 224)
(1980)]
*Scydmaenus heissianus Franz, 1980 [Franz (1980)]
*Euconnus (Napochus) shanusii Franz, 1980 [Franz
(1980)] *Scydmaenus huallagae Franz, 1980 [Franz (1980)]
*Euconnus (Napochus) tarapotae Franz, 1980 *Scydmaenus iquitosanus Franz, 1980 [Franz (1980)]
[Franz (1980)] *Scydmaenus iquitosi Franz, 1980 [Franz (1980)]
*Euconnus (Napoconnus) angustithorax Franz, *Scydmaenus terricola Franz, 1980 [Franz (1980)]
1980 [Franz (1980)] *Scydmaenus tingomariae Franz, 1988 [Franz (1988)]
*Euconnus (Napoconnus) curticornis Franz, 1980 *Scydmoraphes peruanus Franz, 1980 [Franz
[Franz (1980)] (1980)]
Subfamily STAPHYLININAE
Agerodes coeruleus Motschulsky, 1858 [Herman Belonuchus haemorrhoidalis (Fabricius, 1801)
(2001)] [Asenjo et al. (2013)]
Agerodes kraatzii (Kirsch, 1865) [Herman (2001)] [Belonuchus iconnicovi Plavilstschikov, 1929]
*Agerodes nobilis Bernhauer, 1916 [Herman (2001)] [Asenjo (2004)]; delete, nomen nudum
*Agerodes puncticollis Bernhauer, 1906 [Herman *Belonuchus ielskii (Solsky, 1872) [Herman (2001)]
(2001)] Belonuchus longiceps Bernhauer, 1906 [Herman
Agerodes punctipennis Bernhauer, 1906 [Herman (2001)]
(2001)] Belonuchus mordens Erichson, 1840 [Asenjo et al.
*Agerodes semicyaneus Schubert, 1911 [Herman (2013)]
(2001)] *Belonuchus paradoxus Bernhauer, 1906 [Herman
Agerodes semiviolaceus Bernhauer, 1904 [Pla- (2001)]
vilstschikov (1929)] [Belonuchus peruvianus Plavilstschikov, 1929]
Agrodes conicicollis Sharp, 1876 [Asiain et al. [Asenjo (2004)]; delete, nomen nudum
(2007)]; 5A. longiceps Sharp, 1876 *Belonuchus pulchripennis Bernhauer, 1908 [Her-
Agrodes elegans Nordmann 1837 [Asiain et al. man (2001)]
(2007)]; 5A. melancholicus (Schubert, 1911) Belonuchus rufipennis (Fabricius, 1801) [Asenjo
*Amblyopinodes amazonicus Guerrero, 1996 [Her- et al. (2013)]
man (2001)] *Belonuchus rugicollis Bernhauer, 1916 [Herman
Amblyopinodes gahani (Fauvel, 1901) [Herman (2001)]
(2001)]; 5A. similis Seevers, 1865 Belonuchus sharpi Bernhauer, 1907 [Herman
*Amblyopinodes peruvianus Seevers, 1955 [Herman (2001)]; 5B. grandiceps Sharp, 1876
(2001)] *Belonuchus skalitzkyi Bernhauer, 1906 [Herman
*Amblyopinus akodoni Seevers, 1955 [Herman (2001)] (2001)]
*Amblyopinus ancashi Seevers, 1955 [Herman (2001)] Belonuchus taczanovskii Solsky, 1872 [Herman
*Amblyopinus brandesi Kraatz, 1900 [Herman (2001)]
(2001)] *Belonuchus vulneratus Bernhauer, 1906 [Herman
*Amblyopinus ielskii Solsky, 1875 [Herman (2001)] (2001)]
*Amblyopinus kalinowskii Seevers, 1955 [Herman Belonuchus xanthopterus (Nordmann, 1837)
(2001)] [Asenjo et al. (2013)]
*Amblyopinus kofordi Seevers, 1955 [Herman (2001)] Chroaptomus flagrans (Erichson, 1840) [Chani-
*Amblyopinus machadoi Barrera 1972 [Herman Posse (2006)]
(2001)] Creophilus maxillosus (Linnaeus, 1758) [Clarke (2011)]
*Amblyopinus monticolus Seevers, 1955 [Herman Creophilus variegatus Mannerheim, 1830 [Clarke
(2001)] (2011)]
*Amblyopinus pacae Seevers, 1955 [Herman (2001)] Delgadobius amazonensis Chani-Posse & Couturier,
*Amblyopinus piurae Seevers, 1955 [Herman 2012 [Chani-Posse and Couturier (2012)]
(2001)] Edrabius alticolus Seevers, 1955 [Herman (2001)]
*Amblyopinus punae Seevers, 1955 [Herman (2001)] *Edrabius pearsoni Seevers, 1955 [Herman (2001)]
*Amblyopinus sanborni Seevers, 1944 [Herman Edrabius peruanus Seevers, 1955 [Herman (2001)]
(2001)] *Elecatopselaphus peruvianus Scheerpeltz, 1972
*Amblyopinus spinipennis Seevers, 1955 [Herman [Herman (2001)]
(2001)] Elmas gigas Ashe & Chatzimanolis, 2003 [Ashe and
*Atopocentrum mirabile Bernhauer, 1906 [Herman Chatzimanolis (2003)]
(2001)] Elmas lescheni Ashe & Chatzimanolis, 2003 [Ashe
*Belonuchus angusticollis Bernhauer, 1906 [Herman and Chatzimanolis (2003)]
(2001)] Eulissus amicus Bernhauer, 1916 [Herman
Belonuchus asperatus Bernhauer, 1906 [Herman (2001)]
(2001)] Eulissus chalybaeus Mannerheim, 1830 [Herman
Belonuchus candens (Erichson, 1840) [Asenjo et al. (2001)]
(2013)] Eulissus rutilus (Perty, 1830) [Herman (2001)]
Belonuchus clypeatus Sharp, 1876 [FMNH collec- Gastrisus mimetes Sharp, 1884 [FMNH collection
tion (det. M. Bernhauer)] (det. M. Bernhauer)]
Belonuchus convexus (Bernhauer, 1908) [Herman Gastrisus opaculus Sharp, 1884 [Asenjo (2004)]
(2001)] *Gastrisus putidiusculus (Tottenham, 1939) [Her-
*Belonuchus frater Bernhauer, 1906 [Herman (2001)] man (2001)]
Glenus ielskii Solsky, 1872 [Herman (2001)] Nordus nigroscutellatus (Bernhauer, 1905) [Chatzi-
Glenus vestitus Sharp, 1876 [FMNH collection (det. manolis (2004)]
A. Newton)] *Nordus periergos Chatzimanolis, 2004 [Chatzima-
Heterolinus xanthogaster (Solsky, 1875) [Asenjo nolis (2004)]
(2004)]; 5H. distinctus Bernhauer, 1905 *Nordus picticornis (Solsky, 1872) [Chatzimanolis
Heterothops paralleliceps Bernhauer, 1917 [Scheer- (2004)]
peltz (1960)] *Nordus portokalis Chatzimanolis, 2004 [Chatzi-
*Heterothops titschacki Bernhauer, 1941 [Herman manolis (2004)]
(2001)] *Nordus seferisi Chatzimanolis, 2004 [Chatzima-
Holisus analis Erichson, 1839 [Herman (2001)] nolis (2004)]
Homalolinus canaliculatus (Erichson, 1839) [Her- Nordus simplex (Sharp, 1876) [Chatzimanolis
man (2001)] (2004)]
*Isanopus eptaskouros Chatzimanolis, 2008 [Chat- Nordus solitarius (Sharp, 1884) [Herman (2001)];
zimanolis (2008)] 5N. occipitalis (Bernhauer, 1905)
Isanopus tenuicornis Sharp, 1876 [Chatzimanolis Nordus speciosus (Bernhauer, 1905) [Chatzimanolis
(2008)] (2004)]
Leistotrophus versicolor (Gravenhorst, 1806) [Her- Nordus styloceros (Sharp, 1876) [Chatzimanolis
man (2001)] (2004)]
Leptopeltus flavipennis (Erichson, 1840) [Chani- [Nordus testaceus (Fabricius, 1801)] [Plavilstschi-
Posse and Asenjo (2013)] kov (1929)]; Peru probably in error, teste
Leptopeltus montivagans Bernhauer, 1906 [Chani- Chatzimanolis (2004)
Posse and Asenjo (2013)] [Nordus xanthocerus (Nordmann, 1837)] [Herman
*Leptopeltus weyrauchi Chani-Posse & Asenjo, (2001)]; Peru in error, teste Chatzimanolis (2004)
2013 [Chani-Posse and Asenjo (2013)] Oligotergus fasciatus (Nordmann, 1837) [Herman
*Megamblyopinus germaini (Fauvel, 1900) [Her- (2001)]
man (2001)] Ontholestes brasilianus Bernhauer, 1906 [SEMC
*Megamblyopinus mniszechi (Solsky, 1875) [Her- collection (det. A. Newton)]
man (2001)] *Ophionthus serpentinus Bernhauer, 1908 [Herman
Mimosticus pseudosharpi Brunke & Solodovnikov, (2001)]
2014 [Brunke & Solodovnikov (2014)] Paederomimus aberrans (Sharp, 1876) [FMNH
Mimosticus sharpi Brunke & Solodovnikov, 2014 collection (det. M. Bernhauer)]
[Brunke & Solodovnikov (2014)] Paederomimus aeneiceps (Sharp, 1876) [Asenjo
Mimosticus tenuiformis Brunke & Solodovnikov, (2004)]
2014 [Brunke & Solodovnikov (2014)] Paederomimus cribricollis (Erichson, 1840) [Her-
Mimosticus viridipennis (Sharp, 1884) [Brunke & man (2001)]
Solodovnikov (2014)] *Paederomimus sulciceps Bernhauer, 1917 [Herman
Nordus aeneipennis (Bernhauer, 1917) [Chatzima- (2001)]
nolis (2004)] *Paraxenopygus peruvianus Bernhauer, 1915 [Her-
Nordus amazonicus (Sharp, 1876) [Chatzimanolis man (2001)]
(2004)] Phanolinus coelestus (Solsky, 1875) [Herman (2001)]
Nordus antennatus (Sharp, 1876) [Chatzimanolis Phanolinus cyanescens (Guerin-Meneville, 1844)
(2004)] [Asenjo et al. (2013)]
Nordus batesi (Sharp, 1876) [Chatzimanolis (2004)] *Phanolinus dives (Erichson, 1847) [Herman (2001)]
Nordus cribricollis (Sharp, 1876) [Chatzimanolis *Phanolinus divinus Bernhauer, 1911 [Herman
(2004)] (2001)]
Nordus diversiventris (Bernhauer, 1917) [Chatzima- Phanolinus episcopus Bernhauer, 1906 [Asenjo (2004)]
nolis (2004)] Phanolinus exornatus (Solsky, 1875) [Herman
*Nordus elegans Chatzimanolis, 2004 [Chatzima- (2001)]
nolis (2004)] *Phanolinus peruvianus Bernhauer, 1917 [Herman
Nordus fungicola (Sharp, 1884) [Newton et al. (2001)]
(2005)] *Phanolinus princeps Bernhauer, 1906 [Herman
Nordus gebieni (Bernhauer, 1917) [Chatzimanolis (2001)]
(2004)] *Philonthus abdominalis Erichson, 1847 [Herman
Nordus maculiceps (Sharp, 1876) [Chatzimanolis (2001)]
(2004)]; 5N. schuberti (Wendeler, 1926) *Philonthus fassli Bernhauer, 1916 [Herman (2001)]
Nordus major (Bernhauer, 1905) [Chatzimanolis Philonthus feralis Erichson, 1840 [Asenjo et al.
(2004)] (2013)]
Philonthus figulus Erichson, 1840 [Asenjo (2004)] Platydracus fassli (Bernhauer, 1908) [BMNH,
Philonthus flavolimbatus Erichson, 1840 [Herman SEMC collections (det. A. Newton)]
(2001)] Platydracus femoratus (Fabricius, 1801) [Asenjo et
Philonthus iopterus Erichson, 1840 [Plavilstschikov al. (2013)]
(1929)] [Platydracus fervidus (Sharp, 1884)] [Asenjo
*Philonthus limaensis Solsky, 1872 [Herman (2001)] (2004)]; delete, Peru in error (Newton in prep.)
Philonthus longicornis Stephens, 1832 [Asenjo Platydracus gratiosus (Sharp, 1876) [SEMC collec-
(2004)] tion (det. A. Newton)]
Philonthus lynchi Bernhauer & Schubert, 1914 Platydracus gratus (Sharp, 1876) [FMNH, SEMC
[Chani-Posse (2010)]; 5P. affinis Lynch, 1884 collections (det. A. Newton)]
Philonthus metallifer Bernhauer, 1916 [Herman Platydracus mimeticus (Bernhauer, 1917) [FMNH
(2001)] collection (det. A. Newton)]
*Philonthus ophthalmicus Bernhauer, 1941 [Her- *Platydracus notatus (Solsky, 1872) [Herman (2001)]
man (2001)] Platydracus ochropygus (Nordmann, 1837) [Her-
*Philonthus palcoensis Bernhauer, 1941 [Herman man (2001)]
(2001)] Platydracus osculatii (Guerin-Meneville, 1855)
[Philonthus pallipes Blanchard, 1842] [unrecorded [Asenjo et al. (2013)]
but probably in Peru] Platydracus parviceps (Sharp, 1876) [Asenjo et al.
*Philonthus pavidus Erichson, 1847 [Herman (2013)]
(2001)] Platydracus scabrosus (Curtis, 1839) [BMNH col-
Philonthus peruvianus Bernhauer, 1916 [Herman lection (det. A. Newton)]
(2001)] Platydracus semicyaneus (Bernhauer, 1915) [Her-
*Philonthus pulcher Bernhauer, 1908 [Herman man (2001)]
(2001)] Platydracus vetustus (Sharp, 1876) [Asenjo et al.
Philonthus punctipennis (Solier, 1849) [Chani-Posse (2013)]
(2008)] Platyprosopus major Sharp, 1876 [Herman (2001)]
Philonthus succinctus Guerin-Meneville, 1844 [Her- Plochionocerus fulgens (Fabricius, 1793) [Asiain et
man (2001)] al. (2007)]; 5P. carinatus (Bernhauer, 1916)
*Philonthus supernus Herman, 2001 [Herman Plochionocerus humeralis (Sharp, 1885) [Asiain et
(2001)]; 5P. excelsus Bernhauer, 1941 al. (2007)]
*Philonthus titschacki Bernhauer, 1941 [Herman Plochionocerus igneus (Fauvel, 1901) [Asiain et al.
(2001)] (2007)]
*Philonthus tristipennis Bernhauer, 1941 [Herman Plochionocerus impressipennis (Sharp, 1891) [Asiain
(2001)] et al. (2007)]
*Philothalpus juanecoi Asenjo & Ribeiro-Costa, Plochionocerus janthinus (Erichson, 1847) [Asiain et
2009 [Asenjo and Ribeiro-Costa (2009)] al. (2007)]; 5P. peruvianus (Bernhauer, 1907)
Philothalpus loksos Chatzimanolis & Ashe, 2005 Plochionocerus splendens (Blanchard, 1842) [Asiain
[Chatzimanolis and Ashe (2005)] et al. (2007)]; 5P. fulgens (Brulle, 1836)
*Philothalpus lucieae Asenjo & Ribeiro-Costa, 2009 Plociopterus chlorocephalus Bernhauer, 1917 [Her-
[Asenjo and Ribeiro-Costa (2009)] man (2001)]
Philothalpus mundus (Sharp, 1876) [Chatzimanolis Plociopterus ielskii Solsky, 1872 [Herman (2001)]
and Ashe (2005)] Plociopterus laetus Sharp, 1876 [Asenjo (2004)]
Philothalpus pecki Chatzimanolis & Ashe, 2005 Plociopterus mirandus Sharp, 1876 [Asenjo (2004)]
[Chatzimanolis and Ashe (2005)] Plociopterus miriventris Bernhauer, 1905 [Herman
Philothalpus stravos Chatzimanolis & Ashe, 2005 (2001)]
[Chatzimanolis and Ashe (2005)] Plociopterus traili Sharp, 1876] [Soukup (1942), as
Platydracus amazonicus (Sharp, 1876) [FMNH, Brachydirus]
SEMC collections (det. A. Newton)] Plociopterus tricolor Sharp, 1876 [Asenjo (2004)]
[Platydracus antiquus (Nordmann, 1837)] [Asenjo Plociopterus zischkai Scheerpeltz, 1960 [Asenjo
(2004)]; delete, Peru in error (Newton in prep.) (2004)]
Platydracus bolivianus (Bernhauer, 1908) [FMNH Quedius clypealis Sharp, 1876 [Soukup (1942)]
collection (det. A. Newton)] Quedius fulgidus (Fabricius, 1793) [Asenjo (2004)]
Platydracus buquetii (Laporte, 1835) [Herman (2001)] Quedius mesomelinus (Marsham, 1802) [Herman
Platydracus chrysotrichopterus (Scheerpeltz, 1933) (2001)]
[BMNH, FMNH collections (det. A. Newton)] Renda brendelli Marquez, 2010 [Marquez (2010)]
Platydracus emeritus (Herman, 2001) [Asenjo et al. *Renda brevipennis Marquez, 2010 [Marquez
(2013)]; P. priscus (Sharp 1876) is a synonym (2010)]
Renda clavicornis (Sharp, 1876) [Marquez (2010)] Xanthopygus calidus (Erichson, 1839) [Herman
Renda fasciata Marquez, 2010 [Marquez (2010)] (2001)]
Renda fimetaria (Sharp, 1876) [Marquez (2010)] Xanthopygus cognatus Sharp, 1876 [Herman (2001)]
Renda flagellicornis (Nordmann, 1837) [Marquez Xanthopygus faustus (Erichson, 1839) [Asenjo et al.
(2010)] (2013)]
Renda formicaria (Laporte, 1835) [Marquez *Xanthopygus max Blackwelder, 1944 [Herman
(2010)] (2001)]; 5X. peruvianus (Bernhauer, 1916)
Renda lescheni Marquez, 2010 [Marquez (2010)] Xanthopygus punctatus Bernhauer, 1905 [Herman
Renda minor (Sharp, 1876) [Marquez (2010)] (2001)]
Renda nitida Marquez, 2010 [Marquez (2010)] Xanthopygus sapphirinus (Erichson, 1839) [Herman
Renda ophthalmica Marquez, 2010 [Marquez (2010)] (2001)]
*Xanthopygus skalitzkyi (Bernhauer, 1906) [Her-
*Renda palpalis Marquez, 2010 [Marquez (2010)]
man (2001)]
*Scaponopselaphus mutator (Sharp, 1876) [Asenjo
Xanthopygus xanthopygus (Nordmann, 1837) [Her-
et al. (2013)]
man (2001)]
*Scytalinus spectabilis Bernhauer, 1915 [Herman
Xenopygus analis (Erichson, 1840) [Herman (2001)]
(2001)]
Xenopygus bicolor (Laporte, 1835) [Asenjo et al.
Styngetus egregius (Erichson, 1840) [Newton et al.
(2013)]
(2005)] *Xenopygus peruvianus (Solsky, 1872) [Herman
*Styngetus heynei Bernhauer, 1917 [Herman (2001)] (2001)]
Styngetus langei Bernhauer, 1905 [Herman (2001)] *Zackfalinus antennatus Chatzimanolis, 2012
Tesba gigas Sharp, 1876 [Herman (2001)] [Chatzimanolis (2012)]
*Thyreocephalus fassli Bernhauer, 1916 [Herman *Zackfalinus brzoskai Chatzimanolis, 2012 [Chat-
(2001)] zimanolis (2012)]
Thyreocephalus lynceus (Erichson, 1839) [Herman Zackfalinus leptophallos Chatzimanolis, 2012
(2001)] [Chatzimanolis (2012)]
Thyreocephalus subtilis Sharp, 1885 [Herman Zackfalinus napus Chatzimanolis, 2012 [Chatzima-
(2001)] nolis (2012)]
Subfamily STENINAE
*Stenus amata Puthz, 1984 [Herman (2001)] Stenus koerneri Puthz, 2007 [Puthz (2007a)]
Stenus asserculifer Puthz, 2006 [Puthz (2006b)] *Stenus latilabris Puthz, 2006 [Puthz (2006b)]
Stenus atromicans Puthz, 1972 [Puthz (2006d)] Stenus latinus Puthz, 1984 [Puthz (2006e)]
*Stenus aurantiacoornatus Puthz, 2005 [Puthz Stenus nigricans Sharp, 1876 [Herman (2001)]
(2005)] *Stenus nodicollis Puthz, 2000 [Herman (2001)]
Stenus cerritulus Puthz, 1988 [Herman (2001)] *Stenus nusta Puthz, 2005 [Puthz (2005)]
*Stenus cincinnifer Puthz, 2000 [Herman (2001)] *Stenus obscurefactus Puthz, 1987 [Herman
Stenus colonus Erichson, 1840 [Newton et al. (2001)]
(2005)] Stenus ornativentris Sharp, 1886 [FMNH collection
*Stenus colossipennis Puthz, 2000 [Herman (det. Puthz)]
(2001)] Stenus palpalis Erichson, 1840 [Soukup (1942)]
*Stenus curaca Puthz, 1984 [Herman (2001)] Stenus perpulcher Solsky, 1875 [Herman (2001)]
*Stenus peruvianus Bernhauer, 1908 [Herman
Stenus cursitor Sharp, 1876 [Puthz (2006b)]
(2001)]
Stenus cuzcoensis Puthz, 1971 [Herman (2001)]
*Stenus praecellens Bernhauer, 1916 [Herman
Stenus distinctus Benick, 1917 [Puthz (1988)]
(2001)]
*Stenus flavocinctus Puthz, 2005 [Puthz (2005)]
*Stenus pulchripennis Puthz, 2005 [Puthz (2005)]
*Stenus flavofasciatus Puthz, 2005 [Puthz (2005)] *Stenus pygmalion Puthz, 1984 [Herman (2001)]
Stenus flavotaeniatus Puthz, 2005 [Puthz (2005)] *Stenus rectifrons Benick, 1917 [Herman (2001)]
Stenus fronto Puthz, 2007 [Puthz (2007a)] *Stenus sagittifer Puthz, 1988 [Herman (2001)]
*Stenus garavitoensis Puthz, 1979 [Herman (2001)] Stenus subcoeruleus Bernhauer, 1916 [Puthz (2007a)]
Stenus gayi Solier, 1849 [Herman (2001)] Stenus titicacanus Bernhauer, 1908 [Herman
Stenus hromadkaianus Puthz, 1993 [Herman (2001)]
(2001)]; 5S. comatus Hromadka, 1986 Stenus tricolor Sharp, 1876 [Puthz (2007a)]
*Stenus inka Puthz, 1969 [Herman (2001)] *Stenus tridentifer Puthz, 1984 [Herman (2001)]
*Stenus inti Puthz, 1988 [Herman (2001)] *Stenus zabolicus Puthz, 2014 [Puthz (2014b]
Subfamily TACHYPORINAE
Coproporus pulchellus (Erichson, 1839) [Asenjo et *Tachinomorphus andinus Bernhauer, 1918 [Her-
al. (2013)] man (2001)]
Coproporus segnis (Sharp, 1883) [Asenjo et al. (2013)] Vatesus goeldii (Wasmann, 1900) [Herman (2001)]
*Sepedophilus peruvianus (Bernhauer, 1917) [Her-
man (2001)]
Note
The important work cited below appeared in March 2015, but was not available in
time to include the many new Peruvian taxa or country records in this checklist. It
includes 5 new genera and 60 new species of Aleocharinae found in Peru (all but one
species endemic to Peru) as well as a few genera and species that are reported for the
first time from Peru. This brings the confirmed species count for Peruvian
Staphylinidae to 950 (through June 2015).
Pace, R. 2015. Biodiversity of Aleocharinae from Andean and Brazilian sub-
regions: new data, new genera and new species (Coleoptera: Staphylinidae). In
Biodiversity of South America. II. Coleoptera Staphylinidae Aleocharinae, pp. 39
173. Memoirs on Biodiversity 3, WBA Project, Verona. 173 pp.
Acknowledgements
I thank Angelico Asenjo for providing an updated draft of his checklist of
Peruvian Staphylinidae for comparison to this list, and for comments on the
inclusion of certain species; Matthew Gimmel for producing the initial draft of this
contribution (he earned but declined coauthorship); and Caroline Chaboo for useful
comments on it. I also thank many contributors of electronic checklists that served
as starting points for parts of my world database catalog of Staphyliniformia against
which this contribution was checked, notably J. Milton Campbell and Anthony
Davies (Tachyporinae), Donald S. Chandler (Pselaphinae), Lee H. Herman
(Paederinae), Ivan Lobl (Scaphidiinae) and Sean T. OKeefe (Scydmaeninae). I
acknowledge NSF-EPSCoR # 66928 (to Caroline S. Chaboo) for supporting the
Peru beetle project and Matthew Gimmels research in the Chaboo lab, and
a University of Kansas Ecology and Evoltuonary Biology-General Research Fund
(to Chaboo) for funding this publication. Finally, I thank Edilson Caron and an
anonymous reviewer for comments that improved the manuscript.
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