# Passalidae

Passalidae is a family of beetles known variously as "bessbugs", "bess beetles", "betsy beetles", "betsy bugs", 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 to 43 mm, for having a single "horn" on the head, and for a form of social behavior unusual among beetles.
Bodies are 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 quasisocial (common nest + brood caring) beetles that live in groups within rotting logs or stumps. The beetles will excavate tunnel systems within rotting wood where the females then lay their eggs. 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.
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. The family has been suggested to have a close relationship with the extinct family Passalopalpidae.

## Genera

* Aceraius [es] – Analaches [es] – Aponelides – Aulacocyclus [es] – Austropassalus [es] – Basilianus [es] – Cacoius – Ceracupes – Cetejus [es] – Cicernonius – Comacupes [es] – Cylindrocaulus – Didimus – Epishenus – Episphenoides – Eumelosomus – Flaminius – Gonatas [es] – Heliscus – Hincksius – Labienus [es] – Leptaulax [es] – Macrolinus [es] – Malagasulus – Mastochilus – Neleides – Neleuops – Nelues – Ninus – Odontotaenius – Ogyges [es] – Oileus – Ophrygonius – Orgyes – Paratiberioides – Passalus – Passipassalus – Paxillus [es] – Pelopides [es] – Pelops – Pentalobus – Petrejoides [es] – Petrejus – Pharochilus – Phoroneus – Plesthenus – Pleurarius – Popilius – Proculejus – Proculus [es] – Protomocoelus – Pseudepisphenus [es] – Pseudoarrox [es] – Ptichopus – Publius [es] – Rhodocanthopus – Semicyclus [es] – Solenocyclus [es] – Spasalus [es] – Spurius – Taeniocerus [es] – Tarquinius [es] – Tiberioides – Trichopleurus – Undulifer – Verres – Veturius [es] – Vindex – Vitellinus [es] – Xylopassaloides [es]

## Selected species

* Genus Aceraius
    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