Accessibility / Report Error

Extending the geographic and altitudinal range of Popilius gibbosus (Burmeister, 1847) (Coleoptera: Passalidae) with taxonomical comments

Abstract

Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.) was known only from localities in the Cordillera Oriental of the northern Andes in Colombia and Venezuela and the southern section in Colombia between 1,350-3,000 m. Herein, we provide the first record for Bolivia at low elevations (< 1,000 m), thus extending the geographic and altitudinal range of this species. Information about characters not yet used and the first description of the aedeagus are provided, accompanied with figures of those characters that facilitate the identification of the species. An identification key for Popilius species in Bolivia and a P. gibbosus distribution map are also provided.

Keywords
Amazon Andean; Andes; Saproxilophagus beetles; Range distribution; South America; Taxonomy

INTRODUCTION

PopiliusKaup, 1871Kaup, J.J. 1871. Monographie der Passaliden. Berliner Entomologische Zeitschrift, 15: 1-125. is a Neotropical genus of Passalidae belonging to the Proculini tribe which, according to the most recent concept promoted by Boucher (2006Boucher, S. 2006. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Annales de la Société Entomologique de France, N.S., 41(3/4): 239-604.), comprises 15 species, distributed from the intertropical region of South America, including Trinidad and the Isthmus of Panama. Based on a phylogenetic analysis for Proculini, Boucher (2006Boucher, S. 2006. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Annales de la Société Entomologique de France, N.S., 41(3/4): 239-604.) considered the genus a natural group supported by one autapomorphy: the lower apical spines of the meso- and metatibiae with a beveled apex. However, Beza-Beza et al. (2020Beza-Beza, C.F.; Jiménez-Ferbans, L. & McKenna, D.D. 2020. Phylogeny and Systematics of Wood-Degrading Neotro­pical Bess Beetles (Coleoptera: Passalidae: Passalinae). Arthropod Systematics & Phylogeny, 78(2): 287-308. https://doi.org/10.26049/ASP78-2-2020-05.
https://doi.org/10.26049/ASP78-2-2020-05...
) do not corroborate this hypothesis, claiming that Popilius species are mixed with others of Petrejoides Kuwert, 1896.

Boucher (2006Boucher, S. 2006. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Annales de la Société Entomologique de France, N.S., 41(3/4): 239-604.) still considered that the genus is subdivided into two groups of species with morphological and biogeographic differences: the “marginatus” group, extra-Andean, dependent on low to medium altitudinal forests (< 1,500 m); and the “hebes” group, intra-Andean north (from 1,000 to more than 3,000 m above sea level). Gillogly (2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.) also reported that almost all species of Popilius have an extremely limited range in distance or altitude.

The second group includes Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.) with distribution on the eastern north Andean face between 1,350 and 3,000 m (Gillogly, 2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.; Boucher, 2006Boucher, S. 2006. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Annales de la Société Entomologique de France, N.S., 41(3/4): 239-604.; Jiménez-Ferbans et al., 2018Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2018. Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia, with the Description of Two New Species. Neotropical Entomology, 47(5): 642-667. https://doi.org/10.1007/s13744-017-0584-1.
https://doi.org/10.1007/s13744-017-0584-...
). Gillogly (2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.) was the only author to carry out a taxonomic review for the genus, which unfortunately was not published in indexed and peer-reviewed journals. However, it has added significant information to the taxonomic knowledge of the genus. In his work, Gillogly (2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.) commented that P. gibbosus forms a complex of seven species, all being synonyms with a single species (P. gibbosus) with morphological variations.

As this species is still the subject of discussions regarding morphological characters used for its identification and with restricted geographical distribution, the present work aims to formally carry out the first record of the occurrence of Popilius gibbosus for Bolivia. Thus, expanding the geographical and altitudinal distribution of the species, now in a lower altitude region (< 1,000 m) and provide a detailed diagnosis, containing images that help to visualize the main taxonomic characters.

MATERIAL AND METHODS

The analyzed material comes from the Laboratório de Sistemática e Ecologia de Coleoptera (LASEC) of the Instituto Nacional de Pesquisas da Amazônia (INPA). The specimens of P. gibbosus were identified using the the original description by Burmeister (1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.), taxonomic revision by Gillogly (2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.), and the diagnosis present in Jiménez-Ferbans et al. (2018Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2018. Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia, with the Description of Two New Species. Neotropical Entomology, 47(5): 642-667. https://doi.org/10.1007/s13744-017-0584-1.
https://doi.org/10.1007/s13744-017-0584-...
) for Colombian specimens. The terminology for the cephalic capsule and mandibles follows Boucher (2006Boucher, S. 2006. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Annales de la Société Entomologique de France, N.S., 41(3/4): 239-604.), and to other parts of the body, it follows Reyes-Castillo (1970Reyes-Castillo, P. 1970. Coleoptera, Passalidae: morfología e división em grandes grupos; géneros Americanos. Folia Entomológica Mexicana, 20-22: 1-240.).

The images were obtained in a Leica M165C stereomicroscope with a coupled DFC295 camera and processed in the LAS Version 4.2 software to perform the morphometric measurements. Then, they were stacked in layers by the software Helicon Focus Version 7.6.1 to generate a single image of combined focus. For uniform and more effective illumination, we used the geodesic dome of lighting according to Kawada & Buffington (2016Kawada, R. & Buffington, M.L. 2016. A scalable and modular dome illumination system for scientific microphotography on a budget. PloS One, 11(5): e0153426. https://doi.org/10.1371/journal.pone.0153426.
https://doi.org/10.1371/journal.pone.015...
). The images were edited for light and contrast correction, and the figures were made using Adobe Photoshop, following the guide proposed by Bevilaqua (2020Bevilaqua, M. 2020. Guide to image editing and production of figures for scientific publications with an emphasis on taxonomy Image editing for scientific publications. Zoosystematics and Evolution, 96(1): 139-158. https://doi.org/10.3897/zse.96.49225.
https://doi.org/10.3897/zse.96.49225...
). The distribution map was produced with aid of the website SimpleMappr (Shorthouse, 2010Shorthouse, D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps. [Retrieved from https://www.simplemappr.net]
https://www.simplemappr.net...
).

RESULTS

Popilius gibbosus ( Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p. )

Figs. 1 - 2

Figure 1
Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.), habitus. (A) dorsal; (B) ventral; (C) lateral.

Figure 2
Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.). (A) Head and prothorax, dorsal; (B) Head, ventral (C) Head and prothorax, dorsolateral; (D) Prosternal process and mesosternum; (E) Metasternum (left side); (F) Aedeagus (ventral, dorsal and lateral view).

Passalus gibbosusBurmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.: 507.

Pertinax gibbosus: Kaup, 1869Kaup, J.J. 1869. Prodromus zu einer Monographie der Passaliden. Coleopterologische Hefte, 4: 1-40.: 21.

Popilius gibbosus: Kaup, 1871Kaup, J.J. 1871. Monographie der Passaliden. Berliner Entomologische Zeitschrift, 15: 1-125.: 76; Kuwert, 1897Kuwert, A. 1897. Die Passaliden Dichotomisch Bearbeitet. Novitates Zoologicae, 4: 274-306.: 299; Hincks & Dibb, 1935Hincks, W.D. & Dibb, J.R. 1935. Coleopterorum catalogus. Pars 142: Passalidae. s’Gravenhage, W. JUNK. 118p.: 17; Reyes-Castillo, 1970Reyes-Castillo, P. 1970. Coleoptera, Passalidae: morfología e división em grandes grupos; géneros Americanos. Folia Entomológica Mexicana, 20-22: 1-240.: 107; Jiménez-Ferbans et al., 2018Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2018. Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia, with the Description of Two New Species. Neotropical Entomology, 47(5): 642-667. https://doi.org/10.1007/s13744-017-0584-1.
https://doi.org/10.1007/s13744-017-0584-...
: 17.

Oileoides gibbosus: Pereira, 1941Pereira, F.S. 1941. Contribuição para o conhecimento da subfamília dos Pseudacanthinae (Col. Passalidae). Arquivos de Zoologia do Estado de São Paulo, 3(5): 93-113.: 99; Hincks & Dibb, 1958Hincks, W.D. & Dibb, J.R. 1958. Coleopterorum catalogus. Suplementa Pars 142: Passalidae. ‘sGravenhage, Uitgevery Dr. W. JUNK. 32p.: 7.

Material examined: BOLIVIA: Region Chapare - 400 m, 06-ix-58, leg. Zischka // Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.) M. Bevilaqua det. 2020Bevilaqua, M. 2020. Guide to image editing and production of figures for scientific publications with an emphasis on taxonomy Image editing for scientific publications. Zoosystematics and Evolution, 96(1): 139-158. https://doi.org/10.3897/zse.96.49225.
https://doi.org/10.3897/zse.96.49225...
. 1 ♂ (INPA). Bolivia oriental, Santa Cruz - 500 m, 24-xi-55, Zischka // Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.) M. Bevilaqua det. 2020Bevilaqua, M. 2020. Guide to image editing and production of figures for scientific publications with an emphasis on taxonomy Image editing for scientific publications. Zoosystematics and Evolution, 96(1): 139-158. https://doi.org/10.3897/zse.96.49225.
https://doi.org/10.3897/zse.96.49225...
. 1 ♀ (INPA).

Diagnosis: medium-sized specimens (24.5-26 mm), body flat (Fig. 1). Labrum with a concave anterior edge; clypeus large and wide with slightly concave anterior edge, with anterior angles large and acute; posterior angles large, almost fused with mediofrontal tubercles; clypeofrontal groove strong (Fig. 2A). Frontal area straight or slightly convex, finely punctate. Posterior mediofrontal area with longitudinal groove and that extends until the central tubercle. Posterofrontal ridges weak, flat, straight, reaching the inner tubercles, which are inconspicuous. Central tubercle small, with apex not free, and deep groove on posterior region, fused and placed behind the lateroposterior tubercles, which are large, well-marked, and transverse. Lateropostfrontal areas shallow, with pubescent punctation. Canthus ocular with apex rounded, not reaching middle of eye. Antennal club with three large and narrow lamellae (Fig. 2A). Central area of mentum not dilated, glabrous, impunctate; sides with large and shallow scars. Hypostomal process glabrous and convex laterally, without longitudinal groove (Fig. 2B). Prothorax pubescent ventrally (Fig. 1B); anterior angles of prothorax rounded, not protruding (Figs. 1A, 2C); pronotal punctation present only on lateral fossae, with presence of short setae (Fig. 1C). Marginal pronotal groove wide, deep and impunctate, practically reaching the longitudinal groove (Fig. 2C). Prosternal process rhomboidal, with long and sparse setae (Fig. 2D). Mesosternal scars narrow, shallow, glabrous, with opaque surface (Fig. 2D). Metasternal disc not delimited by punctuations; metasternum without punctures on posterior region and densely pubescent laterally (Fig. 2E). Humeri glabrous. Aedeagus slightly longer than wide; median lobe globose, slightly larger than basal piece and parameres together, and strongly sclerotized ventrally. Parameres smaller than basal piece, separated by a conspicuous suture (except centrally) ventrally. In dorsal view, projections of the parameres are not joined. The sclerotization of basal piece is truncate laterally, and membranous toward dorsal surface (Fig. 2F).

Distribution: Colombia (Boyacá, Cundinamarca, Magdalena, Meta, Valle del Cauca), Venezuela (Aragua, Trujillo) (Gillogly, 2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.; Jiménez-Ferbans et al., 2018Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2018. Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia, with the Description of Two New Species. Neotropical Entomology, 47(5): 642-667. https://doi.org/10.1007/s13744-017-0584-1.
https://doi.org/10.1007/s13744-017-0584-...
); Bolivia (Cochabamba, Santa Cruz) (Fig. 3).

Figure 3
Geographical distribution of Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.) in South America, with previous and new records.

Remarks: According to Gillogly (2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.) and Jiménez-Ferbans et al. (2018Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2018. Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia, with the Description of Two New Species. Neotropical Entomology, 47(5): 642-667. https://doi.org/10.1007/s13744-017-0584-1.
https://doi.org/10.1007/s13744-017-0584-...
), Popilius gibbosus is restricted to high-altitudes (1,350-3,000 m) in eastern portion of the Andes of Colombia and Venezuela. The specimens from Bolivia were collected at the slope of the eastern portion of the Cordillera at low altitudes (400 m and 500 m). This makes it possible to expand both the information on the distribution and the altitude at which this species can be found, being able to assist in the collection of this species, which is relatively rare in several collections.

Also, according to Gillogly (l.c.), the absence of specimens from the western Cordillera and the Central Cordillera may be an effect of the lack of collection, as mentioned by Jiménez-Ferbans et al. (2019Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2019. The Passalidae (Coleoptera, Scarabaeoidea) from Bolivia, with the descriptions of three new species. ZooKeys, 882: 51-85. https://doi.org/10.3897/zookeys.882.35532.
https://doi.org/10.3897/zookeys.882.3553...
) referring to Bolivia’s bess beetle fauna. Jiménez-Ferbans et al. (2019Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2019. The Passalidae (Coleoptera, Scarabaeoidea) from Bolivia, with the descriptions of three new species. ZooKeys, 882: 51-85. https://doi.org/10.3897/zookeys.882.35532.
https://doi.org/10.3897/zookeys.882.3553...
) carried out a checklist of Passalidae species from Bolivia, finding two Popilius species: P. marginatus (Percheron, 1835) and P. tetraphyllus (Eschscholtz, 1829). The authors commented that the number of species of Bolivia is low and reflects the lack of a systematic exploration of this country; more surveys are needed, especially in ecosystems such as montane forest and tropical rain forest, pointing out the need for sampling in the mid-range montane ecosystems of Bolivia (Jiménez-Ferbans et al., 2019Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2019. The Passalidae (Coleoptera, Scarabaeoidea) from Bolivia, with the descriptions of three new species. ZooKeys, 882: 51-85. https://doi.org/10.3897/zookeys.882.35532.
https://doi.org/10.3897/zookeys.882.3553...
).

Key to Popilius species from Bolivia (Adapted from Jiménez-Ferbans et al., 2019 Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2019. The Passalidae (Coleoptera, Scarabaeoidea) from Bolivia, with the descriptions of three new species. ZooKeys, 882: 51-85. https://doi.org/10.3897/zookeys.882.35532.
https://doi.org/10.3897/zookeys.882.3553...
)

1. Antennal club with three lamellae ......................................................... 2

1’. Antennal club with four or five lamellae ................................................................................................. Popilius tetraphyllus (Eschscholtz, 1829)

2. Metasternum glabrous anterolaterally ....................................................................................................... Popilius marginatus (Percheron, 1835)

2’. Metasternum pubescent anterolaterally ...................................................................................................... Popilius gibbosus (Burmeister, 1847Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.)

DISCUSSION

Gillogly (2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.) commented that P. gibbosus forms a complex of seven species that must be treated as synonyms. He also commented that there are subtle morphological variations between the species studied (including type specimens), but that such variations are not sufficient to distinguish the species to the point of being treated as different. The characters used by Gillogly in his review agree with those of our specimens, except for the presence of setae on the hypostomal process, elytral humeri, and on the last abdominal segment, which are absent in the specimens we have. In the diagnosis provided by Jiménez-Ferbans et al. (2018Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2018. Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia, with the Description of Two New Species. Neotropical Entomology, 47(5): 642-667. https://doi.org/10.1007/s13744-017-0584-1.
https://doi.org/10.1007/s13744-017-0584-...
) the only character that does not match our specimens are the antennal clubs, which according to the authors are short, although in the image available in the work it is possible to observe that they are long and thin, similar to our specimens.

Gillogly (2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.) did not explore characters from the aedeagus, although male genitalia has already proved useful in the identification of species, groups of species, or genera (Reyes-Castillo, 1970Reyes-Castillo, P. 1970. Coleoptera, Passalidae: morfología e división em grandes grupos; géneros Americanos. Folia Entomológica Mexicana, 20-22: 1-240.; Bührnheim, 1978Bührnheim, P.F. 1978. O edeago na sistemática de passalídeos americanos (Coleoptera: Passalidae). Acta Amazonica, 8(1): 5-61.; Cano, 1994Cano, E.B. 1994. Estado actual sobre el conocimiento del edeago en Passalidae (Coleoptera), con la descripción del edeago en seis especies del Viejo Mundo. Acta Zoológica Mexicana (nueva serie), 61: 21-34.; Boucher, 2006Boucher, S. 2006. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Annales de la Société Entomologique de France, N.S., 41(3/4): 239-604.). The aedeagus of P. gibbosus, which has not been described so far, does not resemble any other known in the literature, making this an excellent structure to differentiate the species from the others in the genus.

In a phylogenetic analysis proposed by Beza-Beza et al. (2020Beza-Beza, C.F.; Jiménez-Ferbans, L. & McKenna, D.D. 2020. Phylogeny and Systematics of Wood-Degrading Neotro­pical Bess Beetles (Coleoptera: Passalidae: Passalinae). Arthropod Systematics & Phylogeny, 78(2): 287-308. https://doi.org/10.26049/ASP78-2-2020-05.
https://doi.org/10.26049/ASP78-2-2020-05...
) that aimed to understand the relationships between the genera of New World Passalidae, Popilius was not rescued as a monophyletic group, becoming paraphyletic concerning Petrejoides. Popilius gibbosus, one of the species used in the analysis, was recovered as a sister group of Petrejoides tenius Kuwert, 1896. Therefore, as other authors (Gillogly, 2005Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.; Beza-Beza et al., 2020Beza-Beza, C.F.; Jiménez-Ferbans, L. & McKenna, D.D. 2020. Phylogeny and Systematics of Wood-Degrading Neotro­pical Bess Beetles (Coleoptera: Passalidae: Passalinae). Arthropod Systematics & Phylogeny, 78(2): 287-308. https://doi.org/10.26049/ASP78-2-2020-05.
https://doi.org/10.26049/ASP78-2-2020-05...
) have already suggested, there is a need for an extensive taxonomic review accompanied by phylogenetic analysis of Popilius to clarify the limits of the genus and the species that are included in it. We hope that the information available here may help to resolve this issue in the future.

ACKNOWLEDGMENTS:

We would like to thank the reviewers for providing important considerations to the article.

REFERENCES

  • Bevilaqua, M. 2020. Guide to image editing and production of figures for scientific publications with an emphasis on taxonomy Image editing for scientific publications. Zoosystematics and Evolution, 96(1): 139-158. https://doi.org/10.3897/zse.96.49225
    » https://doi.org/10.3897/zse.96.49225
  • Beza-Beza, C.F.; Jiménez-Ferbans, L. & McKenna, D.D. 2020. Phylogeny and Systematics of Wood-Degrading Neotro­pical Bess Beetles (Coleoptera: Passalidae: Passalinae). Arthropod Systematics & Phylogeny, 78(2): 287-308. https://doi.org/10.26049/ASP78-2-2020-05
    » https://doi.org/10.26049/ASP78-2-2020-05
  • Boucher, S. 2006. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Annales de la Société Entomologique de France, N.S., 41(3/4): 239-604.
  • Bührnheim, P.F. 1978. O edeago na sistemática de passalídeos americanos (Coleoptera: Passalidae). Acta Amazonica, 8(1): 5-61.
  • Burmeister, H. 1847. Handbuch der Entomologie: Besondere Entomologie, Fortsetzung: Coleoptera Lamellicornia Xylophila et Pectinicornia. Berlin, G. Reimer. v. 5, viii + 584p.
  • Cano, E.B. 1994. Estado actual sobre el conocimiento del edeago en Passalidae (Coleoptera), con la descripción del edeago en seis especies del Viejo Mundo. Acta Zoológica Mexicana (nueva serie), 61: 21-34.
  • Gillogly, A.R. 2005. Review of the Genus Popillius and preliminary phylogeny of Passalidae (Coleoptera). - Texas A&M University, Texas, USA. PhD Dissertation.
  • Hincks, W.D. & Dibb, J.R. 1935. Coleopterorum catalogus. Pars 142: Passalidae. s’Gravenhage, W. JUNK. 118p.
  • Hincks, W.D. & Dibb, J.R. 1958. Coleopterorum catalogus. Suplementa Pars 142: Passalidae. ‘sGravenhage, Uitgevery Dr. W. JUNK. 32p.
  • Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2018. Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia, with the Description of Two New Species. Neotropical Entomology, 47(5): 642-667. https://doi.org/10.1007/s13744-017-0584-1
    » https://doi.org/10.1007/s13744-017-0584-1
  • Jiménez-Ferbans, L.; Reyes-Castillo, P.; Schuster, J.C. 2019. The Passalidae (Coleoptera, Scarabaeoidea) from Bolivia, with the descriptions of three new species. ZooKeys, 882: 51-85. https://doi.org/10.3897/zookeys.882.35532
    » https://doi.org/10.3897/zookeys.882.35532
  • Kaup, J.J. 1869. Prodromus zu einer Monographie der Passaliden. Coleopterologische Hefte, 4: 1-40.
  • Kaup, J.J. 1871. Monographie der Passaliden. Berliner Entomologische Zeitschrift, 15: 1-125.
  • Kawada, R. & Buffington, M.L. 2016. A scalable and modular dome illumination system for scientific microphotography on a budget. PloS One, 11(5): e0153426. https://doi.org/10.1371/journal.pone.0153426
    » https://doi.org/10.1371/journal.pone.0153426
  • Kuwert, A. 1897. Die Passaliden Dichotomisch Bearbeitet. Novitates Zoologicae, 4: 274-306.
  • Pereira, F.S. 1941. Contribuição para o conhecimento da subfamília dos Pseudacanthinae (Col. Passalidae). Arquivos de Zoologia do Estado de São Paulo, 3(5): 93-113.
  • Reyes-Castillo, P. 1970. Coleoptera, Passalidae: morfología e división em grandes grupos; géneros Americanos. Folia Entomológica Mexicana, 20-22: 1-240.
  • Shorthouse, D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps. [Retrieved from https://www.simplemappr.net]
    » https://www.simplemappr.net
  • FUNDING INFORMATION:

    The first author is grateful to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the doctorate scholarship to support this research project (number process: 001/2015).

Edited by

Edited by: Simone Policena Rosa

Publication Dates

  • Publication in this collection
    21 Feb 2022
  • Date of issue
    2022

History

  • Received
    04 Feb 2021
  • Accepted
    05 Nov 2021
Museu de Zoologia da Universidade de São Paulo Av. Nazaré, 481, Ipiranga, 04263-000 São Paulo SP Brasil, Tel.: (55 11) 2065-8133 - São Paulo - SP - Brazil
E-mail: einicker@usp.br