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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Polyphaga
Infraordo: Scarabaeiformia
Superfamilia: Scarabaeoidea

Familia: Passalidae
Subfamiliae (4): Aulacocyclinae - Leptaulacinae - Macrolininae - Passalinae
Overview of genera (71)

Aceraius – Ameripassalus – Analaches – Aponelides – Aulacocyclus – Austropassalus – Basilianus – Cacoius – Ceracupes – Cetejus – Cicernonius – Comacupes – Cylindrocaulus – Didimus – Epishenus – Episphenoides – Eumelosomus – Flaminius – GonatasHeliscus – Hincksius – Labienus – Leptaulax – Macrolinus – Malagasulus – Mastochilus – Neleides – Neleuops – Nelues – NinusOdontotaeniusOgyges – Oileus – Ophrygonius – Orgyes – Paratiberioides – Passalus – Passipassalus – Paxillus – Pelopides – Pelops – Pentalobus – Petrejoides – Petrejus – Pharochilus – Phoroneus – Plesthenus – Pleurarius – PopiliusProculejusProculus – Protomocoelus – Pseudepisphenus – Pseudoarrox – Ptichopus – Publius – Rhodocanthopus – Semicyclus – Solenocyclus – SpasalusSpurius – Taeniocerus – Tarquinius – Tiberioides – Trichopleurus – UnduliferVerresVeturiusVindex – Vitellinus – Xylopassaloides

Passalidae Leach, 1815

selected references

Smith, A.B.T. 2006: A review of the family-group names for the superfamily Scarabaeoidea (Coleoptera) with corrections to nomenclature and a current classification. Coleopterists Society monograph, 5: 144–204. DOI: 10.1649/0010-065X(2006)60[144:AROTFN]2.0.CO;2 PDF
Vasconcelos da Fonseca, C.R.; Barbosa, M.L.L.; Farnandez, M.F.S. 2011: A hypothetical evolutionary history of passalid beetles narrated by the comparative anatomy of the hindgut (Coleoptera: Passalidae). Zootaxa, 3012: 1–20. Preview

Additional references

Boucher, S. 2004: The species of Passalidae (Insecta : Coleoptera) described by Johann Jakob Kaup: Historical overview and critical catalogue, with the description of four new species. Kaupia 13: 99-121. Reference page.
Boucher, S. 2006: Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Les taxons du groupe famille. La tribu des Proculini et son complexe Veturius. Annales de la Société entomologique de France (N.S.) 41(3-4): 239-604. Full article (PDF). Reference page.
Fonseca, C.R.V.D.; Reyes-Castillo, P. 2004: Synopsis on Passalidae family (Coleoptera: Scarabaeoidea) of Brazil with description of a new species of Veturius Kaup, 1871. Zootaxa, 789: 1–26. Abstract & excerpt
Gutiérrez-Velázquez, A. et al. 2013: The classic theory of Mexican Transition Zone revisited: the distributional congruence patterns of Passalidae (Coleoptera). Invertebrate systematics 27(3): 282–293. DOI: 10.1071/IS12056 Reference page.
Jiménez-Ferbans, L. et al. 2013: A checklist and key for the identification of bess beetles (Coleoptera: Passalidae) of Argentina. Zootaxa 3701(2): 192–206. DOI: 10.11646/zootaxa.3701.2.4 Reference page.
Jiménez-Ferbans, L., Reyes-Castillo, P. & Schuster, J.C. 2015: Passalidae (Coleoptera: Scarabaeoidea) of the Greater and Lesser Antilles. Zootaxa 3956(4): 491–512. DOI: 10.11646/zootaxa.3956.4.3. Preview (PDF). Reference page.
Catalogue of Palearctic Coleoptera. Vol. 3 ed. I. Lobl, & A. Smetana, Apollo Books, Stenstrup, Denmark, 2006
ISBN 87-88757-59-5, p.77


Australian Faunal Directory
Tree of Life Web Project. 2007. Passalidae. Bess beetles. Version 02 May 2007 (temporary) in The Tree of Life Web Project

Vernacular names
Deutsch: Zuckerkäfer
English: bess beetles
français: Passale
русский: Сахарные жуки

Passalidae is a family of beetles known variously as "bessbugs",[1] "bess beetles",[1] "betsy beetles"[1] or "horned passalus beetles". Nearly all of the 500-odd species are tropical; species found in North America are notable for their size, ranging from 20–43 mm, for having a single "horn" on the head, and for a form of social behavior unusual among beetles.

Bodies elongate-cylindrical and black overall; ventral surfaces may be covered with yellow setae. The head is narrower than the thorax, with antennae consisting of 10 antennomeres with a three-segment club. The elytra are elongate with parallel sides, and heavily striated.

They are subsocial (brood caring) beetles that live in groups within rotting logs or stumps.[2] The beetles will excavate tunnel systems within rotting wood where the females then lay their eggs.[3] They care for their young by preparing food for them and helping the larvae construct the pupal case. Both adults and larvae must consume adult feces which have been further digested by microflora for a time; an arrangement that might be described as a sort of external rumen.

In addition, they are also able to produce fourteen acoustical signals, more than many vertebrates. Adults produce the sounds by rubbing the upper surface of the abdomen against the hind wings. The larvae produce the sounds by rubbing the third leg against a striated area on the coxa of the second leg.

While the taxonomy of Nearctic species is well-known (four species in the US, and 90 in Mexico), bess beetles in other parts of the world need further study.

Of North American species, Odontotaenius disjunctus (synonym: Popilius disjunctus) is the familiar bessbug found throughout the eastern US and Canada, while O. floridanus has only been found in Florida on sand hills that used to be islands when Florida was flooded thousands of years ago. Ptichopus angulatus was recently discovered near the border of Mexico in Arizona. Its habitat is south to Colombia and it is commonly associated with the detritus chambers of leafcutter ant nests (Atta spp.). Two other species were reported from Arizona at the beginning of the 20th century, but have not been seen there since that time; they may have been brought from Mexico by a train hauling firewood.

The oldest records of the family go back to the Cretaceous, with the genus Ceracyclus known from two species found in the Cenomanian aged Burmese amber, the genus appears to be closely related to the living genus Cylindrocaulus.[4] The family has been suggested to have a close relationship with the extinct family Passalopalpidae.[5]

Selected species

Genus Aceraius

Aceraius laevicollis from under a rotten log in Mindanao, Philippines

Aceraius grandis

Genus Aulacocyclus

Aulacocyclus edentulus

Genus Ceracupes

Ceracupes arrowi

Genus Chondrocephalus

Chondrocephalus debilis
Chondrocephalus granulifrons

Genus Cylindrocaulus

Cylindrocaulus patalis

Genus Didimus

Didimus laevis
Didimus parastictus

Genus Heliscus

Heliscus tropicus

Genus Leptaulax

Leptaulax bicolor

Genus Odontotaenius

Odontotaenius disjunctus (patent-leather beetle)
Odontotaenius floridanus
Odontotaenius striatopunctatus

Genus Ogyges

Ogyges laevior

Genus Oileus

Oileus rimator

Genus Passalus

Passalus affinis
Passalus caelatus
Passalus elfriedae
Passalus inops
Passalus interruptus
Passalus interstitialis
Passalus jansoni
Passalus latifrons
Passalus pugionifer
Passalus punctatostriatus
Passalus punctiger
Passalus spiniger
Passalus unicornis

Genus Paxillus

Paxillus leachi
Paxillus pentataphylloides

Genus Pentalobus

Pentalobus barbatus

Genus Petrejoides

Petrejoides orizabae

Genus Popilius

Popilius eclipticus

Genus Proculus

Proculus burmeisteri
Proculus mniszechi

Genus Ptichopus

Ptichopus angulatus

Genus Publius

Publius agassizi

Genus Spasalus

Spasalus crenatus

Genus Spurius

Spurius bicornis

Genus Verres

Verres corticicola
Verres hageni

Genus Veturius

Veturius transversus

See also

Austroplatypus incompertus


John L. Foltz (2001-07-08). "Family Identification. Coleoptera: Passalidae". University of Florida. Archived from the original on 2008-12-18.
"Generic Guide to New World Scarab Beetles-Scarabaeoidea-Passalidae Overview". Retrieved 2017-01-22.
"What Are Bess Beetles?". Education. Retrieved 2017-01-22.
Boucher, Stéphane Verfasser (2017). Ceracyclini, tribe nov. of Passalidae Aulacocyclinae for Cylindrocaulus Fairmaire and Ceracyclus, gen. nov., with two new species from the Cenomanian Burmese amber (Coleoptera, Scarabaeoidea). Les Cahiers Magellanes. ISBN 978-2-35387-135-3. OCLC 1098209160. {{cite book}}: |last= has generic name (help)

Boucher, Stéphane; Bai, Ming; Wang, Bo; Zhang, Weiwei; Yang, Xingke (September 2016). "†Passalopalpidae, a new family from the Cretaceous Burmese amber, as the possible sister group of Passalidae Leach (Coleoptera: Scarabaeoidea)". Cretaceous Research. 64: 67–78. doi:10.1016/j.cretres.2016.03.017.


Jack C. Schuster, "Passalidae", in Ross H. Arnett, Jr. and Michael C. Thomas, American Beetles (CRC Press, 2002), vol. 2
BugGuide Passalidae

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