127
The Bolboce a idae, Geo upidae (Coleop e a, Sca abaeoidea),
and Sca abaeinae (Coleop e a, Sca abaeoidea, Sca abaeidae)
auna o Hondu as: Taxonomy and dis ibu ion
José D. Ri e a-Dua e1, Rena o P. Salomão2, Ángel Solís3, Mau icio Michel4,
Ma cela I. Ma amo os1, Be Kohlmann3
1 Labo a o io de Hid obiología, Depa amen o de Ecología y Recu sos Na u ales, Escuela de Biología, Facul ad de Ciencias, Uni e sidad Nacional Au ónoma de
Hondu as, F ancisco Mo azán, Hondu as
2 Facul ad de Es udios Supe io es Iz acala, Uni e sidad Nacional Au ónoma de México, Tlalnepan la de Baz, Mexico
3 BioAl a Ba coding P ojec , San o Domingo de He edia, Cos a Rica
4 Escuela Ag ícola Paname icana, Depa amen o de Ciencia Ag ícola y P oducción, Colección de Insec os, Zamo ano, Hondu as
Co esponding au ho : Rena o P. Salomão ([email p o ec ed])
Copy igh : © José D. Ri e a-Dua e e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
A biodi e si y and dis ibu ion analysis o he bee le amilies Bolboce a idae and Geo up-
idae, and he sub amily Sca abaeinae (Sca abaeidae) o Hondu as is p esen ed o he
i s ime. The checklis con ains 98 species: wo Bolboce a idae, one Geo upidae, and 95
Sca abaeinae. The auna o Hondu as includes wo gene a o Bolboce a idae, one genus
o Geo upidae, and 22 o Sca abaeinae. On hophagus is he mos speciose genus, wi h
21 species, ollowed by Can hon (10 species) and Cop is (9 species). Valid names and
synonyms a e p o ided. Fi s coun y eco ds, no es on p esen ly ecognized species, dis-
ibu ion maps, and biodi e si y compa isons a e included, and nomencla u al p oblems
associa ed wi h species in ques ion a e discussed. Phanaeus lempi a Kohlmann, Solís
and Ri e a, sp. no . (Coleop e a: Sca abaeidae) is desc ibed as a new species. Can hon
sallei Ha old, 1863 and Can hon spinosus Ha old, 1863 a e eins a ed as alid species.
Can hon cyanellus LeCon e, 1859 does no con ain any subspecies. Two new synonymies
a e es ablished: Can hon cyanellus iole ae Hal e , 1961, syn. no . o Can hon sallei Ha -
old, 1863; Can hon speciosus Ha old, 1868, syn. no . o Can hon spinosus Ha old, 1863.
Key wo ds: Cen al Ame ica, dis ibu ion a las, endemics, iden i ica ion key, Neo opics,
new species, species e alida ion, synonymiza ion
In oduc ion
The dung bee les o he sub amily Sca abaeinae comp ise 6,939 desc ibed spe-
cies and belong o he Sca abaeidae amily, which con ains a o al o 36,455
species (Schoolmees e s 2025). Dung bee les a e among he mos s udied and
well-known insec g oups (Hanski and Cambe o 1991; Nichols e al. 2007;
Schol z e al. 2009; Cupello e al. 2023). Al hough hey a e dis ibu ed wo ldwide,
hey a e mo e abundan in he opics han in o he geog aphical egions (Gill
1991). Mos Sca abaeinae use dung as hei main ood esou ce; he e a e o he
common die s, such as nec ophagy, sap ophagy, and myce ophagy (Simmons
Academic edi o : And ey F olo
Recei ed:
18 Feb ua y 2025
Accep ed:
25 June 2025
Published:
10 Oc obe 2025
ZooBank: h ps://zoobank.
o g/15F7AF4E-9EC3-40EF-9C6D-
E6B131FA5A0C
Ci a ion: Ri e a-Dua e JD,
Salomão RP, Solís Ángel, Michel M,
Ma amo os MI, Kohlmann B (2025)
The Bolboce a idae, Geo upidae
(Coleop e a, Sca abaeoidea),
and Sca abaeinae (Coleop e a,
Sca abaeoidea, Sca abaeidae)
auna o Hondu as: Taxonomy
and dis ibu ion. ZooKeys 1255:
127–206. h ps://doi.o g/10.3897/
zookeys.1255.150586
ZooKeys 1255: 127–206 (2025)
DOI: 10.3897/zookeys.1255.150586
128
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
and Ridsdill-Smi h 2011). As wi h hei die , Sca abaeinae nes ing beha io has
been well s udied, and h ee main s a egies ha e been desc ibed: olle s, un-
nelle s, and dwelle species. Pe haps a ou h ca ego y should be es ablished,
he pushe s, he ones ha push dung wi h hei heads and o wa d pa s o
hei body. Ne e heless, hese ood eloca ion beha io s ha e e ol ed mul i-
ple imes in mul iple phyle ic lineages (Hal e and Edmonds 1982; Emlen and
Philips 2006; Hue a e al. 2023). Because o hei eeding and nes ing beha io ,
dung bee les play i al ecological oles, including nu ien ecycling, soil u no e ,
seconda y seed dispe sal, and li es ock pa asi e con ol (Nichols e al. 2008;
And esen and U ea-Galeano 2022). Al hough he la ges and dominan dung
bee le gene a a e all s ill incomple ely o ully un e ised (Cupello e al. 2023),
hei quick esponsi eness o habi a al e a ions make dung bee les highly e -
ec i e biological indica o s (Fa ila and Hal e 1997; Kohlmann e al. 2007), an
applica ion ha has been ising since he 1990s (Ri e a and Fa ila 2022).
Bolboce a idae and Geo upidae a e pa o he Sca abaeoidea supe ami-
ly and comp ise app oxima ely 641 and 499 species, espec i ely (see Zunino
1984; Howden 1985a, b; Schol z and B owne 1996; B owne and Schol z 2002;
Schoolmees e s 2025). Al hough Geo upidae a e p edominan ly cop oph-
agous, some species eed on o ganic ma e (e.g., aco ns, ungi, g ape ine
lea es) and ca ion (Pé ez-Ramos e al. 2007; Pé ez-Villama es e al. 2016).
Geo upidae exhibi s i s highes di e si y in sub opical and empe a e ecosys-
ems, bu hey can also be ound in opical egions (T o a-Mo eu e al. 2007;
A iaga-Jiménez e al. 2023). Fu he mo e, Geo upidae species show a ious
pa e ns o spa ial dis ibu ion, anging om eu y opic species o hose special-
ized in conse ed (e.g., pine-oak o es s) o dis u bed habi a s (e.g., pas u e-
lands, plan a ions) (Kohlmann 1991; T o a-Mo eu e al. 2007; A iaga-Jiménez
e al. 2023). Rega ding Bolboce a idae, some species a e myce ophagids, eed-
ing on he ungus mycelium, while o he s may be aphagous (Nikolaje 1987).
Adul bee les dig e ical holes and use he soil humus o la al eeding (How-
den 1982). Bolboce a idae is a cosmopoli an amily, wi h i s highes axonomic
di e si y in he sou he n hemisphe e.
Despi e he popula i y o dung bee les and ela ed g oups as a subjec o
s udy in La in Ame ica (see Fa ila e al. 2023), esea ch in Hondu as is consid-
e ably limi ed. To da e, he coun y has only wo scien i ic a icles (Ri e a and
Can a e o 2011; Ri e a and Schlein 2014), one iden i ica ion manual (C eedy
and Mann 2012), and ou unde g adua e heses (Ju Soló zano 2012; Ce i os
Ga cía and Ochoa Cadena 2017; Al a ado López and Oso io Ka an 2020; Engle
2020) ha a e ocused on dung bee le biological and ecological aspec s. The
knowledge o dung bee le di e si y and dis ibu ion in Hondu as is hus p ima i-
ly con ined o hese s udies, supplemen ed by a ew axonomic e isions o a -
ious gene a ha occasionally include species collec ed wi hin Hondu an e i o-
y (e.g., Solís and Kohlmann 2004; Edmonds and Zídek 2010, 2012; Da ling and
Génie 2018; Moc ezuma and Hal e 2021a). To da e, no speci ic esea ch has
been conduc ed on he Bolboce a idae and Geo upidae amilies in Hondu as,
and simila o Sca abaeinae species, p e ious eco ds o he coun y o igina e
om axonomic e isions ha include some species collec ed om he egion.
This wo k p esen s an upda ed checklis o he Bolboce a idae, Geo upidae,
and Sca abaeidae (Sca abaeinae) bee le auna o Hondu as. Addi ionally, we
include he desc ip ion o a new species o Phanaeus and a axonomic e iew
129
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
o Can hon sallei, C. spinosus, and Sisyphus mexicanus. This s udy aims o lay
he g oundwo k o u u e esea ch on dung bee les in Hondu as, encou aging
a g ea e unde s anding o hei di e si y and biogeog aphy in he Ame icas.
Ma e ials and me hods
The checklis o dung bee les o Hondu as was compiled om h ee main
sou ces: (1) ieldwo k in ol ing sys ema ic collec ions using pi all aps bai -
ed wi h human eces, conduc ed ac oss a ious si es in he cen al, wes e n,
no heas e n, and eas e n egions o he coun y du ing 2023 and 2024 (see
below, S udy si es and Dung bee le sampling); (2) a e iew o he en omological
collec ion a he Escuela Ag ícola Paname icana El Zamo ano; and (3) a ho -
ough e iew o eliable bibliog aphic sou ces ocused on eco ds o specimens
collec ed wi hin Hondu as and hei sampling locali ies.
S udy si es
Hondu as is loca ed in he cen e o he Cen al Ame ican is hmus, be ween
13°33'16" and 15°01'08"N and be ween 83°08'89" and 89°21'49"W. I is bo -
de ed o he no hwes by Gua emala, o he sou hwes by El Sal ado , o he
no h by he Ca ibbean Sea, o he sou h by he Gol o de Fonseca o he Paci ic
Ocean, and o he eas by Nica agua. I is he second-la ges coun y in Cen al
Ame ica, wi h a e i o ial ex ension o 112,492 km2 and a o es ed a ea ha
ep esen s 56% o he e i o y (6,315 million hec a es), o which 36% is p o ec -
ed wi hin he 91 decla ed p o ec ed a eas. These include e es ial, aqua ic,
and coas al ma ine ecosys ems (V eugdenhil e al. 2002).
Due o olcanic ac i i y and o ogenic p ocesses in pas imes, i is he mos
moun ainous e i o y in he egion, wi h wo- hi ds o i s e i o y co e ed by
moun ains (63%) and a smalle p opo ion o plains along i s coas s (37%) (Me-
jía O dóñez and House 2002). Such o og aphic condi ions c ea e egions wi h
dis inc and con as ing p ecipi a ion and empe a u e pa e ns. Mejía O dóñez
and House (2002) classi y he e i o y in o h ee la ge geomo phic egions: he
Paci ic Coas al Plain, he No he n Coas al Plain, and he Moun ainous Region
made up o he No he n, Cen al, and Sou he n idges.
Hondu as has a e age annual empe a u es in ypical anges o he opical
zone be ween 18 and 30 °C. The clima e is cha ac e ized by a p ecipi a ion e-
gime wi h wo dis inc seasons: he d y season, om No embe o May, and he
ainy season, om June o Oc obe , esul ing in an a e age annual p ecipi a ion
o 800–3,200 mm (ICF 2022). Such clima ic condi ions de e mine he di e en
li e zones o he coun y (Fig. 1), which ange om opical pine sa annas o
low mon ane ain o es s (cloud o es ) and sub opical ain o es s (pine o es )
(ICF 2022). Sampling was pe o med in se en egions o Hondu as ep esen -
ing con as ing ecosys ems o he coun y: opical d y o es s, opical cloud
o es s, opical ain o es s, and sub opical pine o es s (Fig. 1).
Celaque Moun ain Na ional Pa k (PNMC, 14°34'N, 88°40'W) is loca ed in he
sou hwes e n egion o Hondu as and co e s an a ea o 26,393 hec a es. The
pa k spans pa s o he depa men s o Lempi a (19,630.53 ha), Oco epeque
(5,943.75 ha), and Copán (804.14 ha). The al i ude in he egion a ies om 1,400
o 2,849 m a.s.l., wi h he la e ep esen ing he highes poin in he coun y.
130
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
The a ea ecei es annual ain all be ween 1,600 mm and 1,800 mm, wi h el-
a i e humidi y anging om 74% o 78%. Fu he mo e, empe a u es luc ua e
be ween a maximum o 35 °C and a minimum o 18 °C. The d y season ex ends
om No embe o Ap il, wi h Janua y and Feb ua y being he d ies mon hs (ICF
and MAPANCE 2016). Acco ding o he Hold idge classi ica ion, wo li e zones
a e iden i ied wi hin he PNMC: he sub opical humid o es , which co e s 40%
o he a ea in he bu e zones and ea u es species such as Liquidamba s y aci-
lua L. swee gum, Cle h a spp., and Nec and a spp.; and he lowe mon ane hu-
mid o es , which co e s ~60% o he PNMC’s a ea, including he pa k’s highes
ele a ions, whe e species such as Pinus maximinoi H.E. Moo e and Pinus ooca -
pa Schiede ex Schl dl a e ound (ICF and MAPANCE 2016).
La Tig a Na ional Pa k (PNLT; 14°09'N, 87°08'W) is loca ed in he cen al egion
o Hondu as, in he depa men o F ancisco Mo azán, no heas o he capi al,
Tegucigalpa, M.D.C. I co e s a o al a ea o 24,040 hec a es, spanning he mu-
nicipali ies o he Cen al Dis ic , Valle de Ángeles, San a Lucía, and Can a anas
(ICF 2013a). La Tig a is pa o he San Juanci o moun ain ange, wi h ele a ions
anging om 1,200 o 2,290 m a.s.l. The annual a e age empe a u e a ies be-
ween 13.6 °C and 22.8 °C. Two p ima y li e zones a e iden i ied in he a ea. The
i s is he lowe mon ane we sub opical o es , which ac s as a ansi ional
zone be ween d y o es and pine o es . This zone is loca ed a ele a ions ang-
ing om 1,300 o 1,800 m a.s.l. and ecei es annual p ecipi a ion o 1,500–2,000
mm. The ege a ion is p edominan ly domina ed by Pinus maximinoi, o en in
associa ion wi h L. s y aci lua. The second li e zone is he lowe mon ane, which
comp ises cloud o es s ound abo e 1,800 m a.s.l. This zone expe iences an
a e age annual p ecipi a ion o ~ 2,000 mm and is cha ac e ized by b oadlea
Figu e 1. Na u al li e zones o Hondu as.
131
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
o es co e , including species such as gian oak (Que cus sp.) and a ious spe-
cies o “aguaca illo” (Nec and a sp., Oco ea sp., Pe sea sp.) (ICF 2013a).
Ce o Azul Meámba Na ional Pa k (PANACAM; 14°50'N, 87°47'W) is loca ed
in he cen al egion o Hondu as, eas o Lake Yojoa. The pa k encompasses
an a ea o 31,339 hec a es, co e ing he municipali ies o Meámba , Sigua e-
peque, and Taulabé in he depa men o Comayagua, as well as he municipal-
i y o San a C uz de Yojoa in he depa men o Co és. The al i ude wi hin he
pa k anges om 400 o 2,090 m a.s.l., wi h mean annual p ecipi a ion anging
om 1,600–2,000 mm and a e age empe a u es be ween 8 °C and 25 °C, de-
pending on he ele a ion (ICF 2012). The a ea is p edominan ly composed o
mon ane and submon ane e e g een o es s (Mejía O dóñez and House 2002),
ea u ing ee species such as P. ooca pa, A bu us xalapensis Kun h, Magnolia
hondu ensis A.M.Molina, and Acacia a nesiana (L.).
Habi a /Species Managemen Re uge o he Hondu an Colib í Esme alda
(Olanchi o, 15°25'N, 86°52'W) is a ese e loca ed in Olanchi o municipali y, Yo o
depa men . This ese e consis s p ima ily o lowland opical d y o es s, wi h an
a e age ele a ion o 400 m a.s.l., mean empe a u es anging om 20°C o 33°C,
and an a e age annual ain all o 500 mm (ASESORA 2009). The ege a ion is
composed mainly o cac i (e.g., Opun ia hondu ensis S andl., S enoce eus yuncke i
(S andl.) M. B a o and Sánchez-Mej), sh ubs (e.g., Apheland a deppeana Schl dl. &
Cham., Lonchoca pus i olius S andl. & L.O. Williams.), and ees (e.g., Guaiacum
sanc um L., Leucaena lempi ana C.E.Hughes), o ming a mosaic o open and closed
canopy ege a ion (House 2004). The egion is highly seasonal, wi h a ainy pe iod
(mean mon hly ain all: 545 mm) om May o Oc obe and a d y pe iod (mean
mon hly ain all: 187 mm) om No embe o Ap il (ASESORA 2009). Al hough he
ese e is conse ed, wi hin i s su oundings, he e a e pa ches o an h opogenic
ac i i ies dedica ed o li es ock p ac ices and Hondu an A my ield aining.
Na ional Pa k Sie a de Agal a (PNSA, 14°54'N, 85°52'W, wi h ele a ions ang-
ing om 600 o 1,000 m a.s.l.) and i s su ounding ag icul u al egions (14°46'N,
85°46'W, ca 320 m a.s.l.) encompass he municipali ies o Ca acamas, Culmí, San-
a Ma ía del Real, and Gualaco, loca ed in he Olancho depa men , eas o Hondu-
as. The mean empe a u e anges om 25.2°C o 28°C, wi h an annual ain all o
1,368 mm (ICF 2002). The ege a ion in he pa k includes opical mois o es s in
he lowland a eas, g adually ansi ioning o lowe mon ane we o es s (cloud o -
es s) a highe ele a ions (Po illo e al. 2013). The ainy season ex ends om May
o Oc obe , wi h a mean mon hly ain all o 383 mm, while he d y season occupies
he emainde o he yea , wi h a mean mon hly ain all o 124 mm (da a ob ained
om Ca acamas Municipali y in Wea he and Clima e 2024). Al hough he po ion
o he Na ional Pa k is mode a ely conse ed, ea u ing small human se lemen s
(i.e., illages) wi hin i s bounda ies, he su ounding ag icul u al egion consis s o
small o es emnan s in e spe sed wi hin a li es ock-domina ed landscape.
B us Laguna is a municipali y loca ed in no heas e n Hondu as, wi hin he
Depa men o G acias a Dios (15°44'N, 84°29'W). I bo de s he Depa men s
o Olancho o he wes and Colón o he no h. B us Laguna, wi h an a ea o
321,234.82 hec a es, cons i u es 38.59% o he Rese a del Homb e y Río Plá a-
no Biosphe e Rese e (RHRP-BR), one o he mos impo an p o ec ed a eas
in he Mesoame ican Biological Co ido and one o he la ges in Hondu as,
co e ing 832,338.963 hec a es (ICF 2013b). The egion’s al i ude a ies om
0 o 710 m a.s.l., wi h an a e age empe a u e o 25 °C (minimums o 23 °C
132
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
and maximums o 33 °C). Annual p ecipi a ion anges om 2,000–4,000 mm,
wi h he ainy season spanning om May o No embe (ICF 2013b). The main
ecosys ems o he su eyed a eas include opical mois o es s and opical
looded o es s o pine sa annas. The o me is composed o a laye o ma u e
b oadlea ees up o 35 me e s all, wi h dominan species such as Swie enia
mac ophylla King, Ced ela odo a a L., and Co dia alliodo a (Ruiz & Pa .) Oken.
On he o he hand, he pine sa anna is domina ed by Pinus ca ibaea Mo ele ,
which g ows alongside g asses, palms, and some i e- esis an ees. The pines
can each a heigh o 15–20 m, and he canopy is co e ed by a dense laye o
he baceous ege a ion and g asses, wi h swamp ege a ion (ICF 2013b).
Dung bee le sampling
Dung bee les we e su eyed du ing June and July 2023, which comp ised he
ainy season. The su eyed egions we e a leas 130 km om each o he . In
each egion, sampling was pe o med in ou si es a leas 2 km apa , excep o
B us Laguna, whe e only h ee si es we e sampled, o aling n = 23 s udy si es.
To ob ain a us wo hy sample o dung bee les om each s udy si e, en pi all
aps bai ed wi h ~ 25 g o esh human eces we e ins alled in a linea ansec ,
wi h a dis ance o ca 40 m om he nea es ones (me hodology adap ed om
Ri e a and Fa ila 2022; Mo a-Aguila e al. 2023). Pi all aps consis ed o a
1,000-ml plas ic cup ha was illed wi h ~ 250 ml o a mix u e o wa e , neu al
de e gen (~ 5 ml), and sal (~ 10 g); such mix u e se ed o cap u e and con-
se e he dung bee les ha ell in he aps. Abo e he plas ic cup, a 50-ml ecip-
ien was a ached wi h he a ac i e bai ; a plas ic lid was a ached o he pi all
ap o a oid he en ance o ainwa e and li e . A e wo days, he pi all aps
we e checked, collec ing and s o ing dung bee les in ials wi h 70% e hanol.
Complemen a y samplings we e conduc ed a he Uni e sidad Nacional
Au ónoma de Hondu as (UNAH) be ween Ma ch and July 2024, co e ing he
end o he d y season and he beginning o he ainy season. Nine d y o es
pa ches we e selec ed, in which ou o i e pi all aps we e ins alled pe pa ch.
The aps we e a anged in linea ansec s, spaced 40 me e s apa , and bai ed
wi h 25 g o esh human eces. Finally, a single sampling e en was conduc -
ed in Sep embe 2024 a Los Olingos Lodge, loca ed wi hin a lowland opical
mois o es in he communi y o Agua Chiqui a in he Depa men o A lán ida
(15°47'1.74"N, 87°38'19.17"W), using en pi all aps bai ed wi h human eces.
Species iden i ica ions o he collec ed specimens we e pe o med wi h he
aid o axonomic keys (e.g., Solís and Kohlmann 2002, 2004, 2013; Edmonds
and Zídek 2012; Da ling and Génie 2018; Cupello e al. 2023; He nández e al.
2024a, b) and in o ma ion by wo o he coau ho s o his s udy (BK and ÁS).
Pho og aphy
Se e al habi us’ pho os we e aken by he co-au ho MM using a Canon EOS
Rebel T5i came a moun ed on a LEICA EZ4 s e eoscope. Ano he se o pho os
was aken by ÁS using an Olympus E-M10 Ma k III digi al came a moun ed on a
WeMac o ocus s acking ail, and hen p ocessed wi h Helicon Focus 8.2.7 Li e
s acking so wa e. Finally, BK ook some de ailed pho os using an Adons a
249S-M Digi al Mic oscope sys em.
133
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Resul s
We eco ded wo species o Bolboce a idae, one species o Geo upidae, and
95 o Sca abaeinae in Hondu as. Among hese, ou Sca abaeine species a e
endemic o he coun y, while wo species ha e been in oduced om he Old
Wo ld (Digi on hophagus gazella and Euoni icellus in e medius). All hese axa
belong o 25 gene a and en ibes. On hophagus is he mos species- ich ge-
nus, wi h 21 species, ollowed by Can hon and Cop is, wi h en and nine axa,
espec i ely (Fig. 2). Thi y-one species a e egis e ed o he i s ime in Hon-
du as. Fou species a e egis e ed as new and awai desc ip ion.
Checklis , axonomic no es, and ema ks
In he p esen lis , he ollowing ma ks a e used as app op ia e: * = endemic
species o he coun y, ** = in oduced species, NR = new eco d, CS = Spe-
cies collec ed in he ield samplings, RL = species epo ed in he e iewed
li e a u e o Hondu as, EZ = species wi h Hondu an specimens housed in
he collec ion o Escuela Ag ícola Paname icana Zamo ano. In he check-
lis , he classi ica ion o Bolboce a idae ollows Schol z and B owne (1996);
he Geo upinae ibal and gene ic classi ica ion ollows Zunino (1984); he
Sca abaeinae ibal classi ica ion ollows Cupello e al. (2023). The check-
lis includes e ised ma e ial om specimens collec ed du ing he 2023–
2024 su eys, wi h axonomic no es and addi ional ema ks on selec ed
species. The newly collec ed ma e ial is indica ed below by species; his
ma e ial is being housed in he Uni e idad Nacional Au ónoma de Hondu as
(UNAH) in Tegucigalpa.
Figu e 2. Rank-abundance cu e wi h he dung bee le gene a dis ibu ion acco ding o he numbe o species eco ded
o each genus.
134
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
BOLBOCERATIDAE
A hy eini
Neoa hy eus exca a us (Cas elnau, 1840) (RL)
Neoa hy eus quad iden a us Howden, 1964.
Mexico, Gua emala, Hondu as, El Sal ado , Cos a Rica, Panama, Colombia, Ven-
ezuela, Guyana, Su iname, F ench Guiana, Pe u, B azil, T inidad and Tobago.
Cas elnau 1840; Howden and Ma ínez 1978; Howden and Gill 1984; How-
den 1985; Ho gan 2002; Hielkema 2017; Pablo-Cea e al. 2023; Mo ales
Mo ales e al. 2024.
Bolboce a ini
Eucan hus (Bolbelasmus) a cua us (Ba es, 1887) (RL)
Bolboce as a cua us Ba es, 1887.
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica, Panama.
Ba es 1887; Howden 1964; Ho gan 2008; Pablo-Cea e al. 2023; Mo ales Mo-
ales e al. 2024.
GEOTRUPIDAE
Ce a o upini
Haplogeo upes gua emalensis iden a us Howden, 1974 (NR, CS, EZ)
Fig. 3A
Geo upes gua emalensis iden a us Howden, 1974.
Hondu as, El Sal ado .
Howden 1974; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (22): Lempi a: G acias, Pa que Nacional Mon-
aña de Celaque; 14°33'53"N, 88°37'54"W, 1622 m; 16.VI.2023; JD Ri e a, R Sa-
lomão and M Michel leg.; pi all ap, human dung; UNAH (4). Same locali y;
14°29'5"N, 88°40'11"W, 2030 m; 17.VI.2023; JD Ri e a, R Salomão and M Michel
leg.; pi all ap, human dung; UNAH (18).
Rema ks. This subspecies was ini ially desc ibed based on ou speci-
mens om a cloud o es in Mon e C is o, El Sal ado , collec ed in 1971
a 2300 m on ho se dung (Howden 1974). Fi y- h ee yea s la e , his ax-
on has been collec ed o a second ime and i is a i s o Hondu as. The
Hondu an specimens we e collec ed in bo h pine–oak and cloud o es s.
This subspecies is pa o a poly ypic species o med by H. g. gua emal-
ensis (Ba es, 1887) and H. g. uniden a us (Howden, 1974). Haplogeo upes
includes a second species ha is oglophile, eddelli (Howden, 1967), om
he Xóchi l ca e in Puebla, Mexico, li ing inside he dung galle ies o oden
nes s. Zunino (1984), in his genus-le el sys ema ics o he Geo upinae, con-
side s Haplogeo upes and i s sis e genus Hal e ius o be he i s b anch
135
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Figu e 3. Habi us. A. Haplogeo upes gua emalensis iden a us, Pa que Nacional Mon aña de Celaque, Depa amen o de
Lempi a; B. Can hon chiapas, Re ugio Hábi a /Especie Colib í Esme alda Hondu eño (Olanchi o), Depa amen o de Yo o;
C. Can hon sallei, Re ugio Hábi a /Especie Colib í Esme alda Hondu eño (Olanchi o), Depa amen o de Yo o; D. Can hon
sallei, B us Lagunas, Rio Pla ano Biosphe e Rese e; E. Can hon spinosus, Agua Chiqui a, Depa amen o de A lán ida;
F. Del ochilum scab iusculum, Pa que Nacional Sie a de Agal a, Depa amen o de Olancho. Scale ba : 2 mm.
o sepa a e om he Ame ican geo upid ee (Ce a o upini). This b anch-
ing canno be ega ded as ecen and could ha e s a ed al eady du ing he
Paleogene (Zunino 1984).
142
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
leg.; 29.VI.2023, 2.7 (1) 346 m, 3.1 (2) 252 m; 30.VI.2023, 3.1 (1) 252 m, 3.5 (3)
246 m, 3.7 (1) 243 m, 3.8 (1) 243 m, 4.3 (1) 251 m, 4.5 (1) 256 m, 4.7 (1) 258 m,
4.8 (3) 257 m. NICARAGUA (9). Managua. Rese a P i ada El Bajo, 649–709 m,
27-28.XI.2019, B. He nández leg., Sende o Húmedo (6), Sende o Ensueño, (2);
Dep. Chinandega. Es e o Real, Ce an es, 35 m, 22.VI.2019. B. He nández leg.
(1). COSTA RICA (580). Alajuela. Caño Neg o, en el Pueblo, 10.8938°N,
84.78884°W, 20 m, Ma ínez Ka ia leg., 10.X.1992, (1); Cebadilla, 9.95653°N,
84.34518°W, 540 m, Solís & Al a ado leg., 25.VIII.1985, (1); Gua uso, 10.6395°N,
84.90911°W, 300 m, Rod íguez Gladys leg., 23.II.1998, (2); Rio Ba anca, 450 m,
Asolis001582, A. Solís leg., 19.VI.2020, (1); Rio San Lo enci o, 5 km N de Colo-
nia Palma ena, 10.2266°N, 84.63249°W, 900 m, I Sca abaeidae cou se leg.,
13.VI.1993, (1); Guanacas e. 3 km NO de Nacaome, P. N. Ba a Honda,
10.1682°N, 85.37365°W, 100 m, Reyes Ma ielos leg., 5.VII.1992, (6); 3.VIII.1992,
(5); 14.IX.1992, (1); 3.V.1993, (5); 1.VII.1993, (1); 2.VII.1993, (1); 1.X.1993, (2);
1.XI.1993, (4); 1.VIII.1994, (1); 1.X.1994, (1); 1.XI.1994, (3); 1.IV.1995, (6); 1.
VII.1995, (3); 1.IX.1996, (1); A.C.A, La C uz La Ga i a Es Los Almend os,
11.0338°N, 85.52479°W, 280 m, López Elba leg., 1.VII.1992, (8); 28.VII.1992, (3);
24.VIII.1992, (4); 23.IX.1992, (6); 23.X.1992, (3); 21.XI.1992, (3); 11.VI.1993, (4);
7.IX.1993, (1); Ma ínez Ka ia leg., 13.X.1993, (1); 300 m, López Elba leg., 3.
VIII.1993, (1); 3.X.1993, (3); 7.VII.1994, (1); 4.IX.1994, (2); 2.X.1994, (2); 4.
XI.1994, (3); 16.VII.1995, (3); 22.VIII.1995, (13); 28.X.1995, (1); Bagaces,
10.3491°N, 85.35234°W, 10 m, Ace edo Daniel leg., 1.VI.1991, (1); 1.VIII.1991,
(1); Cha a ía Ulises leg., 1.VIII.1991, (1); 4.V.1992, (1); 10.3666°N, 85.38326°W,
25 m, Po as Wendy leg., 25.VIII.1999, (2); 30.IX.1999, (3); 29.XI.1999, (2);
Bagaces, Pque Nal Palo Ve de, Línea Vieja, 10.3609°N, 85.33640°W, 10 m, Po -
as Wendy leg., 30.X.1999, (1); 6.V.2000, (2); Cañas, 10.3477°N, 85.35036°W, 50
m, Solís Ángel leg., 6.VII.1985, (1); Ce o El Hacha, 12 km SE de La C uz,
10.9831°N, 85.54106°W, 300 m, Solís Ángel leg., 23.VII.1986, (3); 27.VII.1986,
(4); Ce o Lo os., 9.92947°N, 85.38518°W, 500 m, Po as Wendy leg., 8.XI.2002,
(1); Es Los Almend os, 12 Km Ca e e a a San a Cecilia, 11.0342°N, 85.52754°W,
280 m, López Elba leg., 8.V.1994, (3); 15.VIII.1994, (6); 1.VI.1995, (2); 4.I.1996,
(15); 14.V.1996, (5); 15.VII.1996, (3); 10.X.1996, (9); 9.XI.1996, (3); 23.XI.1996,
(4); Es ación Expe imen al Ho izon es, 10.7054°N, 85.57930°W, 125 m, Mo aga
Calix o leg., 27.VI.1993, (1); Es ación Lomas Ba budal, 10.4950°N, 85.37794°W,
30 m, Ace edo Daniel leg., 1.VII.1991, (2); Es ación Mu ciélago, 8Km S.W. de
Cuajiniquil, 10.9019°N, 85.73101°W, 100 m, Cano Ca olina leg., 15.V.1993, (2);
Quesada F eddy leg., 18.V.1993, (1); Es ación San a Rosa. Sende o Indio Desnu-
do., 10.8388°N, 85.62777°W, 300 m, B iceño Du alie leg., 24.X.2000, (1); Finca
Jenny, 30 Km N de Libe ia, 10.8740°N, 85.57997°W, 240 m, A aya Elda leg.,
10.VIII.1996, (2); 10.X.1996, (1); Finca Loaiciga, 6 Km S. de San a Cecilia,
11.0125°N, 85.42771°W, 500 m, Mo aga Calix o leg., 6.I.1992, (1); Ríos Pe ona
leg., 6.I.1992, (5); Finca YAFA, 11.0402°N, 85.50468°W, 320 m, Cano Ca olina
leg., 19.I.1992, (1); La C uz, 10.9831°N, 85.54106°W, 300 m, López Elba leg., 1.
IX.1991, (5); 1.X.1991, (3); 1.XI.1991, (3); 350 m, López Elba leg., 1.XI.1991, (1);
Solis & Al a ado leg., 15.I.1988, (3); 10.9891°N, 85.42583°W, 700 m, Mo aga
Calix o leg., 1.VI.1994, (8); 1.VIII.1994, (2); La C uz, Es ación Mu ciélago 8Km
S.W. de Cuajiniquil, 10.9019°N, 85.73101°W, 70 m, Cano Ca olina leg., 3.X.1994,
(5); Quesada F eddy leg., 3.X.1994, (4); La C uz, Pque Nal Guanacas e Lado
SSW Ce o El Hacha, 10.9931°N, 85.54293°W, 350 m, Chacón Isid o leg.,
143
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
26.VII.1986, (7); La C uz, Pque Nal San a Rosa, 10.8563°N, 85.61191°W, 290 m,
Janzen Daniel leg., 1.XII.1982, (1); 1.VI.1984, (3); La C uz, San a Elena, P.N. San-
a Rosa Es ación Mu ciélago, 10.9019°N, 85.73101°W, 100 m, Cano Ca olina
leg., 3.XII.1993, (1); Quesada F eddy leg., 16.VI.1993, (2); 70 m, Cano Ca olina
leg., 16.VI.1993, (4); 4.VIII.1994, (1); Quesada F eddy leg., 6.VI.1994, (1); 4.
VIII.1994, (1); 31.X.1994, (1); La Ga i a Es Los Almend os, 11.0338°N,
85.52479°W, 280 m, López Elba leg., 7.I.1993, (1); Libe ia, 10.8364°N,
85.61549°W, 300 m, Janzen Daniel leg., 15.IX.1975, (1); 22.VI.1981, (1); 1.
VII.1982, (1); 1.V.1985, (1); Mo aga Ros e leg., 1.VII.1992, (5); Pa a axónomos
III cu so leg., 3.VI.1992, (8); 13.VI.1992, (2); Pe ei a Guille mo leg., 1.VII.1992,
(1); Libe ia, P.N. San a Rosa, 10.8364°N, 85.61550°W, 300 m, Zumbado Manuel
leg., 25.VI.1999, (1); Libe ia, P.N.S.R. Playa Na anjo, 10.8027°N, 85.67479°W,
25.5 m, Alcáza Eliden leg., 1.I.1991, (8); 1.II.1991, (5); Nicoya, Es ación Ba a
Honda, 10.1698°N, 85.37913°W, 100 m, Po as Wendy leg., 22.VI.2000, (1);
3.VIII.2000, (2); Nicoya, Es ación Ba a Honda””, 10.1698°N, 85.37913°W, 100
m, Po as Wendy leg., 2.IX.2000, (1); Nicoya, San An onio, Es ación Co al de
Pied a, 10.2385°N, 85.33080°W, 16 m, Gamboa Billen leg., 3.VII.2005, (1); P. N.
Ba a Honda, 200 m, Asolis002536, A. Solís leg., 18.V.2021, (1); P. N. Ba a Hon-
da., 10.1701°N, 85.34719°W, 100 m, Zumbado Manuel leg., 30.V.1993, (1); P.N.
Ba a Honda, 10.1710°N, 85.34720°W, 150 m, Unknown leg., 8.VII.2000, (3); P.N.
Ba a Honda. Los Mesones, 10.1700°N, 85.35084°W, 350 m, Reyes Ma ielos
leg., 1.VII.1995, (3); P.N. Guanacas e. Finca Jenny., 10.8663°N, 85.57720°W,
240 m, A aya Elda leg., 20.V.1994, (1); P.N. Guanacas e. La C uz. Es ación Bi-
ológica Ma i za., 10.9625°N, 85.49524°W, 600 m, Pa a axónomos II cu so leg.,
1.VIII.1990, (7); P. N. San a Rosa, Bosque Húmedo, A. Solís leg., 6.VI.2016, (1);
P.N. San a Rosa, Á ea Adminis a i a.””, 10.8399°N, 85.62648°W, 300 m, Janzen
Daniel leg., 1.VIII.1984, (1); Pococí, Río Sa dinas, 10.6440°N, 83.74200°W, 10 m,
A aya Flo leg., 10.XI.1993, (1); Pococí, Río Sa dinas Ba a del Colo ado,
10.6440°N, 83.74200°W, 10 m, A aya Flo leg., 25.VIII.1992, (1); 1.IX.1992, (3);
18.VI.1993, (1); 16.VII.1993, (1); 15 m, A aya Flo leg., 18.IV.1993, (2); 2.I.1994,
(1); 16.III.1994, (1); Pococí, Río Sa dinas Ba a del Colo ado””, 10.6440°N,
83.74200°W, 10 m, A aya Flo leg., 11.XII.1992, (1); R.V.S. Bosque Di iá. San a
C uz. Poza Redonda., 10.1737°N, 85.59588°W, 200 m, Ba on Salaza Raúl leg.,
24.XI.1998, (1); San a Cecilia, 11.0645°N, 85.41737°W, 325 m, Mo aga Calix o
leg., 1.VI.1994, (5); San a C uz, Es B. N. Di iá, 10.1342°N, 85.63359°W, 970 m,
Cá denas Yow leg., 15.IX.2002, (1); Sec o Las Pailas, 4.5 Km SW del Volcán
Rincón de la Vieja, 10.7767°N, 85.35191°W, 800 m, Rod iguez Gladys leg.,
15.VII.1992, (1); Tila án, Tie as Mo enas, 10.5710°N, 85.02594°W, 685 m, Ro-
d iguez Gladys leg., 1.XII.1994, (1); He edia. Cinco Esquinas de Ca izal,
10.1092°N, 84.15359°W, 1800 m, Ba an es Gilbe leg., 8.V.1988, (1); Sa apiquí,
Es Biol La Sel a 3 Km S de Pue o Viejo, 10.4311°N, 84.00333°W, 75 m, OET
INBio ALAS p ojec leg., 1.XI.1992, (1); 1.VIII.1995, (2); Limón, A.C.L.A.C, San
Miguel Albe gue CASACODE Send Ce illos, 9.56835°N, 82.64647°W, 20 m, Lobo
Manuel leg., 23.II.1999, (1); 50 m, Zumbado Manuel leg., 23.II.1999, (1); Amub i,
Sende o Soki, 9.51482°N, 82.95537°W, 70 m, Galla do Ge a dina leg., 2.V.1994,
(2); 6.VI.1994, (3); 1.I.1995, (1); 4.IV.1995, (5); 9.VIII.1995, (1); 1.XI.1995, (1); 2.
III.1996, (1); 1.IX.1996, (1); 1.VI.1997, (1); Mo a And és leg., 3.IV.1995, (2); Ce o
To ugue o, 10.5848°N, 83.52920°W, 50 m, Cha a ía Ulises leg., 1.VII.1990, (1);
60.5 m, Delgado Robe o leg., 1.I.1992, (1); Es . Mi ama , R. B. Hi oy Ce e e,
144
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
9.63336°N, 83.00342°W, 500 m, Ca ballo Ge a do leg., 1.VIII.1994, (3); Limón.
Río Blanco, 9.99556°N, 83.10838°W, 34 m, Solís Manuel leg., 11.IV.2011, (1);
Limón, Valle de la Es ella, 9.67103°N, 83.02614°W, 160 m, A ana Wil edo leg.,
22.IV.1999, (1); Pococí, Colo ado Es ación Cua o Esquinas, 10.5395°N,
83.50648°W, 5 m, Delgado Robe o leg., 1.XI.1990, (3); 23.IV.1991, (2); 1.IX.1991,
(1); 1.I.1993, (1); Solano José leg., 1.VI.1990, (2); Pococí, Es ación Cua o Es-
quinas, 10.5395°N, 83.50648°W, 5 m, Aguila Reinaldo leg., 1.IV.1989, (1); Poco-
cí, Es ación Cua o Esquinas., 10.5395°N, 83.50648°W, 5 m, Delgado Robe o
leg., 1.IX.1992, (1); Pococí, Colo ado, 10.5942°N, 83.71651°W, 150 m, Rojas
Elías leg., 26.VII.1992, (1); 1.VI.1993, (1); Pococí, Sec o Ced ales de la Ri a,
10.5273°N, 83.72578°W, 10 m, Rojas Elías leg., 1.VIII.1995, (2); 1.XI.1995, (1);
1.I.1996, (1); 1.VIII.1996, (1); R.B. Hi oy Ce e e, 9.67537°N, 83.02677°W, 120 m,
Ca ballo Ge a do leg., 1.VII.1993, (2); R.V.S. Gandoca Manzanillo. Cen o Ope -
a i o Manzanillo., 9.63258°N, 82.65905°W, 50.5 m, Quesada F eddy leg.,
6.I.1993, (1); Taylo Ka la leg., 22.X.1992, (1); Talamanca, Amub i, 9.51934°N,
82.95627°W, 70 m, Galla do Ge a dina leg., 12.X.1992, (1); 2.IX.1993, (1); 1.
II.1994, (2); 24.VII.1994, (1); 1.VIII.1994, (1); 1.X.1994, (1); Talamanca, Amub i,
9.51934°N, 82.95627°W, 70 m, Galla do Ge a dina leg., 16.VIII.1992, (1);
12.XI.1992, (1); Talamanca, B a si, 9.51934°N, 82.95627°W, 70 m, Galla do Ge-
a dina leg., 5.I.1993, (1); Talamanca, Valle La Es ella, 9.67176°N, 83.02770°W,
100 m, Ca ballo Ge a do leg., 1.VI.1994, (1); Guzmán Rosa leg., 19.IV.1992, (1);
López Elba leg., 19.IV.1992, (1); 120 m, Quesada F eddy leg., 20.VI.1992, (2);
Pun a enas. Ca a a, sende o Laguna Meánd ica, 9.80544°N, 84.58399°W, 50
m, Guzmán Rosa leg., 1.V.1994, (2); Cóbano, R.N.A. Cabo Blanco, Es San Mi-
guel, 9.58892°N, 85.12397°W, 250 m, Po as Wendy leg., 18.V.2001, (2); 300 m,
Al a ado F ancisco leg., 27.X.1997, (1); Es . San Miguel, 3 Km. NW. de Cabo
Blanco, 9.58121°N, 85.13670°W, 100 m, Ramí ez Ma in leg., 1.IX.1993, (38);
1.XI.1993, (6); Es . San Miguel, Sende o Ma en, 9.63186°N, 85.12681°W, 300 m,
Al a ado F ancisco leg., 29.X.1997, (58); Ga abi o, Res Biol Ca a a, Es Queb a-
da Boni a, 9.76745°N, 84.60811°W, 50 m, Bello E ick leg., 1.VI.1990, (1); 1.
VII.1990, (3); Guzmán Rosa leg., 18.X.1992, (2); Sabo ío Juan Ca los leg., 1.
VII.1992, (1); 1.XI.1992, (1); Zúñiga Rodol o leg., 1.XI.1989, (1); 1.XII.1989, (1);
9.77423°N, 84.60812°W, 100 m, Guzmán Rosa leg., 1.I.1994, (1); 1.IV.1995, (1);
Ga abi o, Tá coles, C uce de queb adas, 9.77649°N, 84.60311°W, 0 m, Quesada
Eddie leg., 5.XI.1990, (1); Guacimal, Finca Buen Amigo Mon e e de 4Km S. de
la Rese a, 10.2767°N, 84.79661°W, 1040 m, Fuen es Zobeida leg., 1.IX.1992,
(1); Jacó, La Ca a a a, 9.57276°N, 84.59156°W, 75 m, Zumbado Manuel leg.,
24.XI.2008, (1); 26.XI.2008, (5); Los Ángeles. Paque a, 9.79028°N, 85.04518°W,
120 m, Al a ado F ancisco leg., 23.VI.1998, (10); Paque a, Rio G ande,
9.87133°N, 84.98519°W, 120 m, Al a ado F ancisco leg., 17.VI.1998, (4); Pun-
a enas, Isla San Lucas, Pues o Isla San Lucas, 9.94652°N, 84.90556°W, 50 m,
Azo ei a José leg., 13.IX.2005, (1); Pun a enas. Rese a Ka en Mogensen, Sen-
de o P incipal, 9.86709°N, 85.05995°W, 315 m, B iceño Du alie leg., 23.VI.2003,
(13); Po as Wendy leg., 22.VI.2003, (2); 335 m, Cá denas Yow leg., 3.VII.2003,
(1); 350 m, Cá denas Yow leg., 20.IX.2003, (2); 8.X.2003, (1); 400 m, Po as
Wendy leg., 24.VI.2003, (1); R. P i . Ka en Mogensen, Sende o T es Ríos,
9.86314°N, 85.06274°W, 315 m, B iceño Du alie leg., 26.VI.2003, (1); 400 m,
Po as Wendy leg., 26.VI.2003, (1); R.N.A. Cabo Blanco, 9.59256°N, 85.11487°W,
300 m, Al a ado F ancisco leg., 16.V.1997, (4); Rese a P i ada Ka en Mogensen,
145
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Queb ada Pé ez, 9.86437°N, 85.06496°W, 315 m, Cá denas Yow leg., 23.
VI.2003, (8); Sec o Laguna Meánd ica., 9.79821°N, 84.58125°W, 100 m, Zúñiga
Rodol o leg., 1.VI.1990, (2); San José, Es . Bijagual, 600 m. N de Bijaguali o,
9.74308°N, 84.54476°W, 500 m, Sabo ío Juan Ca los leg., 1.VII.1995, (1). Pan-
ama (15). Bocas del To o. Almi an e, 58 km no e de Boque e, 9.300°N,
-82.400°W; Changuinola Dis ic , 9.250°N, -82.650°W. Colón. Alhajuela, en la
mi ad del ayec o en e Panamá y Colón, 9.2166°N, -79.6333°W; Mon u a Ca-
noa, Ex emo no e del Lago Ga ún, 9.350°N, -79.800°W; F ijoles, 12 km WNW
de Gamboa, 9.1666°N, -79.7916°W; Pipeline Road, 5 km NW de Gamboa,
9.150°N, -79.7166°W; Río T inidad, Panamá, 9.0166°N, -79.9666°W. Panamá.
Co ozal, 3.5 km NW de Ancón Hill, 8.9833°N, -79.5666°W; Cu undú, 3 km N de
Ancón Hill, 8.9833°N, -79.55°W; Madden Fo es , 9 km E de Gamboa, 9.100°N,
-79.6166°W; Summi Pa k, 8 km SE de Gamboa, 9.0666°N, -79.650°W. Panamá
Oes e. La Campana, 5.5 km WSW de Ce meno, 8.7166°N, -79.900°W; Fo Kob-
be A my Rese a ion, 6.5 km SW de Ancón Hill, 8.900°N, -79.5833°W; Isla Ba o
Colo ado, Panamá, 9.150°N, -79.850°W. Ve aguas. 1.74 km al es e de Paso
Real, Panamá, Leg: A. Solís and C. Víquez, 8.1996°N, -81.2343°W.
Diagnosis. Do sal su ace anges om comple ely glossy o ange-yellow,
o o ange-yellow and g een, o comple ely g een, some imes wi h blackish
maculae on p ono um and/o ely a (Fig. 3C, D); sombe head; pygidium o -
ange-yellow; las abdominal en i e bicolo ed, cen e black and la e al bo de s
o ange-yellow (Fig. 8A, B). Body leng h 6–10 mm. Clypeus wi h ou ee h, wo
cen al clypeal ee h long and slende , sligh ly cu ed a hei base (Fig. 3C, D).
Ven al oo h o p ono al an e ola e al bo de absen . P ono al su ace smoo h,
inely punc a e. Eigh h ely al s ia wi h basal ca ina a iable in leng h. P es-
cu ella imp ession a iable in dep h. Pygidium o ange-yellow wi h ine punc-
ua ion, bea ing no se ae o e y sho and ine se ae; wi h s ong basal ca ina
(Fig. 8C, D). Male pygidial leng h/wid h a io 0.66 (Fig. 8).
Rema ks. This species was o iginally desc ibed by on Ha old (1863) om
G anada, Nica agua. La e , Schmid (1922) eclassi ied i as a a ie y (ambiguous
e m o classical Linnean axonomy o a he e ogeneous g oup o phenomena,
including nongene ic a ia ion o he pheno ype; May and Ashlock 1991: 432) o
Can hon cyanellus LeCon e, 1859, wi hou p o iding any explana ion o his ax-
onomic decision. Such ea men was main ained by Robinson (1948), Hal e
(1961, 2003), Howden (1966), and subsequen au ho s a a subspeci ic le el.
Robinson (1948: 96–97) conside ed ha C. cyanellus eached B azil and Pe u and
ha he dis inc subspecies could only be ecognized by di e ences in colo a ion
because no disc epancies in geni alia we e appa en . Hal e (1961) p oposed
he ollowing subspecies and a ie ies wi hin Can hon cyanellus: C. c. cyanellus
LeCon e, 1859; C. c. iole ae Hal e , 1961; C. c. sallei Ha old, 1863, C. c. sal-
lei a . iangula us Schmid , 1922; and C. c. sallei a . gu icollis Schmid , 1922.
Hal e (1961: 262), ollowing Robinson (1948), ecognized ha he subspecies
we e delimi ed on how colo a ian s we e geog aphically s uc u ed. Hal e
(1961) ecognized h ee subspecies o C. cyanellus as geog aphical aces. Tha
is, allopa ic popula ions dis inguishable on he basis o gene ically based phe-
no ypic di e ences. Quo ing Hal e (1961: 262): “changes in colou a ion co e-
spond o de ined geog aphical egions, wi h a g adual inc ease in he da k g een
pa s om he sou h o he no h”. In o he wo ds, Hal e ’s poly ypic classi i-
ca ion o C. cyanellus was based on he assump ion ha colo a ia ion in he
146
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
species was geog aphically s uc u ed; ha is, he di e en colo phena we e no
ound sympa ically (o a leas only o e a na ow zone o in e g ada ion). This
was wha he claimed his da a showed in 1961. To complemen his iew, Hal e
also enume a ed subspecies-le el (and, he e o e, supposedly geog aphical) di -
e ences in he le el o p ono al and ely al punc a ion, in he p esence o he
hume al ca ina, in he p esence o he scu ella imp ession (in he key, he said
ins ead he p escu ella imp ession), and he shape o he me a ibiae. Acco ding
o his axonomy, he subspecies iole ae was in e media e bo h geog aphically
and pheno ypically be ween cyanellus, o he no h, and sallei, o he sou h.
This iew s a ed o collapse wi h he obse a ion by Solís and Kohlmann
(2002) ha he colo phenon ypi ying ssp. cyanellus, he no he n (Mexican)
subspecies, was also ound sympa ically wi h he colo phenon ypi ying ssp.
sallei, he sou he n (Cen al Ame ican) subspecies, in some locali ies in Cos a
Rica, well beyond he supposed zone o in e g ada ion in sou he n Mexico. The
sou h-no h pheno ypic di ide p oposed by Hal e (1961) was shown o be a
leas pa ly inco ec — pa ly because Hal e (1961) was s ill co ec in no ing
ha o ange, he sallei phenon, is, by a , he dominan phenon in Cen al Ame ica.
So, i was w ong o ea he subspecies as monomo phic, bu co ec abou he
geog aphical di ide in ela i e p eponde ance. Fu he mo e, because he cyanel-
lus phenon was supposedly p esen in wo disjunc po ions o he ange o he
species, Mexico, o he no h, and Cos a Rica, o he sou h, Solís and Kohlmann
(2002) co ec ly conside ed i poly opic. Finally, Hal e (2003) main ained his
axonomic poin o iew in his las axonomic ea men o C. cyanellus.
In his doc o al hesis, Nunes (2019) p ima ily elied on colo and gloss cha -
ac e is ics o alida e C. sallei as a dis inc species. Vaz-de-Mello labeled wo C.
sallei specimens, housed in he Na u al His o y Museum, London (Fig. 9), as lec-
o ype and pa alec o ype in 2014. Howe e , he lec o ype designa ion is in alid
because i has no been published. Nunes (2019) also indica ed ha Can hon
den ige Ha old, 1868 is a synonym o C. sallei and designa ed a lec o ype om
Ma a uin, Colombia, which is housed a he MFNB – Museum ü Na u kunde
Be lin, Leibniz-Ins i u ü E olu ions- und Biodi e si ä s o schung, Be lin, Ge -
many. This lec o ype designa ion is also in alid because he wo k was no pub-
lished. Nunes (2019) gi es a de ailed desc ip ion o he ex e nal mo phology
and male geni alia o his species and conside s C. sallei o be dis ibu ed om
sou he n Chiapas in Mexico o no hwes e n Colombia, which is basically he
same dis ibu ion asc ibed o he subspecies sallei by Hal e (1961) (Fig. 10).
In e es ingly, C. sallei is no p esen in he Paci ic opical ain o es ha spans
sou h Cos a Rica and no he n Panama. I s niche may be aken by Can hon mu-
abilis, a species ha li es in his a ea and also eeds on ca ion. Nolasco-So o
e al. (2020, 2023) ound ha he mo phological a ia ion o he aedeagus does
no align wi h he gene ic-phylogeog aphic s uc u e o his axon, no wi h he
subspecies and a ie ies axonomy based on colo p oposed by Hal e (1961).
Do sal colo analysis by Solís and Kohlmann (2002) o C. sallei in Cos a Rica
indica ed ha he same popula ion could p esen di e en colo pa e ns in
a single locali y a he same ime, and ha hese same colo di e ences had
been used as a diagnos ic cha ac e o iden i ying di e en axa by Hal e
(1961) (C. sallei, C. iole ae). Solís and Kohlmann (2002: 12) did no ecognize
he exis ence o subspecies o C. cyanellus in hei s udy based mainly on colo
di e ences as had been p oposed by Hal e (1961) and co ec ly ecognized
147
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
a he ime ha he popula ion o sallei was poly opic ollowing he ecommen-
da ion made by May and Ashlock (1991: 100) ha :
When subspecies o a species di e only in a single diagnos ic cha ac e
in ol ing colo , size, o pa e n, se e al un ela ed and mo e o less widely sepa-
a ed popula ions may independen ly de elop an iden ical pheno ype. The e o-
lu ionis knows ha such popula ions a e no iden ical gene ically, bu since he
subspecies is no an e olu iona y concep , axonomis s some imes combine
such isually iden ical popula ions in o a single subspeci ic axon. A geog aph-
ically he e ogeneous subspecies o his ype is called a poly opic subspecies.
Usually, i is p e e able no o ecognize poly opic subspecies. In he absence
o diagnos ic di e ences, he e is no legi ima e excuse o ecognizing se e al
subspecies me ely on he basis o locali y.
The p esen s udy has shown ha C. sallei is no longe poly opic, bu poly-
mo phic. Polych oma ism is no limi ed o Cos a Rica, bu o he es o Cen al
Ame ica and sou he n Mexico. The eason he colo phena do no quali y as
subspecies unde May ’s de ini ion is no longe because hey a e poly opic, bu
because hey a e now known no o ep esen geog aphical aces. Being poly o-
pic does no p e en a phenon om ep esen ing a geog aphical ace. Ac ually,
be o e 1969, May a gued o he ecogni ion o poly opic ace as subspecies
in all o his w i ings, including in his 1963 seminal book Animal Species and
E olu ion (May 1963: 388–389; 1969: 193). The e o e, i one adop s a de ini ion
o subspecies as geog aphical aces wi h no polymo phisms, Hal e ’s (1961)
colo -based poly ypic classi ica ion is in alid, and he subspeci ic names mus
be synonymized. The di e ence be ween he Hal e (1961) and he Solis and
Kohlmann iews (2002) does no conce n he species le el o he exis ence o
a ia ion, bu how o ea his a ia ion axonomically wi hin he species: Hal e
(1961) saw he a ia ion as he basis o a poly ypic classi ica ion, whe eas Solís
and Kohlmann (2002) saw i as he basis o a poly opic classi ica ion.
A e u he s udy and analysis o se e al dis ibu ion a eas, ex e nal mo pho-
logical di e ences (p ono al punc ua ion, pygidial leng h, punc ua ion, and se ae;
pygidial and las en i e colo a ion) ha e been ecognized by he p esen au ho s
(AS, BK), allowing he sepa a ion o di e en axa wi hin he C. cyanellus species
complex. Specimens om Panama, Cos a Rica, Nica agua, and Hondu as ha e
been examined and ound o be dis inc species om he popula ions common-
ly dis ibu ed in Mexico. The popula ions o hese Cen al Ame ican a eas co e-
spond o wha was desc ibed as C. sallei by on Ha old (1863). We do no ha e
specimens om Sou h Ame ica, p e en ing mo phological compa isons wi h axa
om ha egion. Howe e , we analyzed m DNA di e ences among he Cen al and
Sou h Ame ican popula ions which a e also colo polymo phic (unpublished da a).
In hei analysis o ype specimens o Sou h Ame ican dung bee les, Vaz-de-
Mello and Cupello (2018) include a pho og aph o he lec o ype o Can hon sal-
lei a . iangula us Schmid , 1920 (Columbien, La Ga i a Gebi ge, W. F ische),
which Schmid (1920) also ci ed (e oneously) o be p esen in Hondu as and
Cos a Rica. The ypical in e ed iangula macula o he p ono um clea ly seen
in he pho og aph by Vaz-de-Mello and Cupello (2018: ig. 45) has no been
ound by us in Cen al Ame ica and is p obably ci cumsc ibed in i s dis ibu ion
o Sou h Ame ica. The same seems o hold o Can hon sallei a . gu icollis
Schmid , 1920 ci ed only om Colombia, which Nunes conside s o be a new
synonym o he ele a ed C. iangula us. In his doc o al hesis, Nunes (2019)
148
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
ecognized he exis ence o C. iangula us Schmid , 1920, om B azil, Colom-
bia, T inidad and Tobago, and Venezuela and sepa a ed i om C. sallei using
he p ono al ma k and gloss in his iden i ica ion key.
Rega ding common axa eco ded o Mexico, Nunes (2019) lis s C. cyanel-
lus, C. spinosus (as speciosus), C. iole ae, and a new species and subspecies,
as well as a inge pene a ion o C. sallei in o sou hwes e n Chiapas. Can hon
sallei can be dis inguished om C. spinosus (see he axonomic his o y o
C. spinosus below in he nex species ea men ). Can hon cyanellus spinosus
has been ea ed as a junio synonym o Can hon cyanellus cyanellus since
Hal e (1961) and is e alida ed he e), as al eady men ioned by on Ha old
(1868b), by i s yellow pygidium wi h ine punc a ion and ei he absen o e y
sho , ine se ae (Fig. 8). In con as , C. spinosus has a g een o blue pygidium
wi h an e iden punc ua ion bea ing sho and hick se ae (Fig. 11). Can hon
sallei has he las abdominal en i e black wi h an o ange-yellow a ea o a i-
able leng h a he ex emes o he segmen and none o e y small se ae (Fig.
8); in con as , C. spinosus has a unicolo da k (no black) las abdominal en-
i e wi h sho hick se ae (Fig. 11). The male pygidial leng h/wid h a io o
C. sallei is 0.66, whe eas C. spinosus has a longe pygidium wi h a a io o 0.72
(Figs 8, 11). Because o hese di e ences in mo phological cha ac e s, C. sallei
is ele a ed back o i s o me species s a us. The cha ac e s used p e iously
by Hal e (1961) o hume al ely al ca ina, scu ella imp ession, and p ono al
punc ua ion a e a iable and as such do no make good axonomic cha ac e s.
Nunes (2019: 64) p esen s in his doc o al hesis a key o dis inguishing
axa wi hin he C. cyanellus species g oup. Howe e , in he i s couple , he mis-
akenly sepa a es C. cyanellus and C. spinosus (as speciosus) om C. sallei,
s a ing ha he do sal colo a ion o he i s wo species is uni o mly g een o
blue. This is inaccu a e, as C. sallei—as no ed by Solís and Kohlmann (2002:
31)—exhibi s a wide ange o do sal colo a ion, om yellow- eddish o en i ely
g een. This a ian can be easily sepa a ed because o i s yellow pygidium e -
sus a blue o g een one in C. cyanellus o C. spinosus. In couple 2 o his key,
Nunes (2019: 64) mis akenly indica ed ha C. sallei does no ha e a ca ina a
he eigh h ely al s ia; C. sallei has his ca ina, which a ies in leng h.
Can hon cyanellus iole ae was desc ibed by Hal e in 1961 based on one
emale specimen om he Paci ic slope o Chiapas. Hal e (1961) based his
subspecies on g een colo ed ely a, a eddish-o ange p ono um and pygidium,
excep a i s blackish-g een pygidial bo de . To y o cla i y he ela ionship
be ween bo h axa, one o us (BK) ied o s udy he holo ype, looking o i in
he Hal e Collec ion ha has now been inco po a ed in o he IEXA Insec Col-
lec ion “D . Miguel Ángel Mo ón Ríos”, Xalapa, Mexico. Howe e , nei he he cu-
a o no BK could ind he holo ype o o he specimens belonging o his axon.
Sadly, he cu a o had no in o ma ion abou he whe eabou s o his holo ype o
addi ional ma e ial o i i had been loaned.
Nunes (2019) ci es he h ee species (C. sallei, C. iole ae, C. sp. no .) coex-
is ing in he same ype locali y o C. iole ae o El Ve gel, Mo ozin la, on he Pa-
ci ic slope o Chiapas. Howe e , he axonomic si ua ion be ween C. sallei and
C. iole ae has been esol ed by Nolasco-So o e al. (2023). In hei ig. 9, hey
p esen ed he esul s o colo a ia ion in ela ion o he o ma ion o clades an-
alyzed by a Bayesian consensus ee. In his analysis, Nolasco-So o e al. (2023)
ob ained he SPS (sou he n Paci ic slope) clade, comp ising popula ions om
149
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
El Ve gel and Raymundo En íquez. These popula ions, cha ac e ized by speci-
mens wi h yellow pygidia, exhibi he ull ange o do sal colo a ion— om g een
ely a, which Hal e (1961) and Nunes (2019) associa ed wi h C. iole ae, o
o ange ely a, p e iously a ibu ed o C. sallei. The coexis ence o all hese col-
o o ms in one locali y a he same ime had al eady been epo ed by Solís
and Kohlmann (2002) o Cos a Rica. We ha e ound he same si ua ion o
Hondu as in his s udy. Nolasco-So o e al. (2023) in e p e ed his si ua ion in
Chiapas o be a possible hyb idiza ion zone be ween C. sallei and C. iole ae,
o ha se e al codominan genes (sic, i should be epis asis, codominance is
p esen in alleles) con ol colo a ia ion.
Ou in e p e a ion o he exis ence o he g ea colo a ia ion epo ed o
he locali ies o C. iole ae in El Ve gel and Raymundo En íquez by Nolasco-So-
o e al. (2023: igs 3, 4, 9; able 2) is simple . I is also consis en wi h one
o he explana ions in oked o Bolbi es by Cupello e al. (2022: 1009), o a
single polymo phic locus in an undi ided deme (a single species). Fi s , a p e i-
ous s udy (Solís and Kohlmann 2002) and his one conce ning Hondu as ha e
ound he coexis ence o specimens wi h g een and o ange ely a ( ep esen ing
C. iole ae and C. sallei, espec i ely) in he same locali y, a he same ime.
Second, he gene ic-phylogeog aphic analysis by Nolasco-So o e al. (2023:
igs 3, 4, 9) shows ha specimens wi h g een and o ange ely a om he same
locali y o m he same gene ic-phylogeog aphic clade (SPS). Conside ing ha
Nunes (2019) indica ed he colo o he ely a as he only di e ence be ween
C. sallei and C. iole ae, his leads o he conclusion ha hei wo-subspecies/
species axonomy is w ong. We ha e a single polymo phic mono ypic species.
We ag ee wi h he commen made by Cupello e al. (2022: 1012):
Fo phanaeines a leas , W. D. Edmonds’ wo ks, which ini ially used he subspe-
cies ca ego y bu la e abandoned i , ha e shown ha a ple ho a o subspecies
names p e iously es ablished o e e o local colou a ian s ha e li le expli-
ca i e o ope a ional alue. They a e, hence, bes ea ed as junio synonyms o
olde names. This should be likely ue o many o he g oups o dung bee les, as
exempli ied by Solís and Kohlmann (2002) and Nolasco-So o e al. (2017, 2020)
who independen ly demons a ed he a i iciali y o Hal e ’s (1961) subspecies
classi ica ion o he colou a ian s o Can hon cyanellus LeCon e, 1859.
We hus conside ha C. iole ae is he junio synonym o C. sallei, and ec-
ommend ha wo k conduc ed and published unde he name o C. iole ae
should be eassigned o C. sallei.
Specimens om he Paci ic slope s emming om he Hal e collec ion
we e also analyzed om Gue e o and Mo elos in he Balsas Ri e Dep ession
and ound o be a new axon esembling C. spinosus and belonging o he C. cy-
anellus species g oup, as Nunes (2019) had al eady sugges ed. I awai s de-
sc ip ion as a new species.
Can hon spinosus Ha old, 1863, eins a ed (NR)
Fig. 3E
Can hon speciosus Ha old, 1868, syn. no .
Mexico, Gua emala, Belize, Hondu as.
150
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Can hon spinosus on Ha old 1863, 1868, 1869, Ba es 1887, Gille 1911, Nunes
2019, Nolasco-So o e al. 2023.
Can hon spinosus Ha old, 1863: 174 (desc ip ion, ca alogue); Ha old 1868: 42
(gene ic key); Gemminge and on Ha old , 1869: 994 (ca alogue); Ba es
1887: 27 (ca alogue); Gille 1911: 29 (ca alogue); Bal hasa 1939a ( axono-
my); Hal e 1961 (synonymy); Howden 1966 (synonymy); Nolasco-So o e
al. 2023: able 2 (phylogeog aphy).
Can hon speciosus Ha old 1868b: 41 (desc ip ion and gene ic key); Ha -
old 1869: 994 (ca alogue); Ba es 1887: 27 ( e ision o Cen al Ame ican
Sca abaeinae); Gille 1911: 29 (ca alogue), Nolasco-So o e al. 2023: able
2 (phylogeog aphy).
Can hon cyanellus speciosus — Bal hasa 1939b: 211 (new s a us); Bal hasa
1951: 328 (ca alogue); Hal e 1961: 262 (synonymy); Howden 1966: 727
(species lis ).
Ma e ial examined (254). Hondu as (4). A lán ida. Tela, Los Olingos Lodge,
9/XI/2024, Cebo heces humanas, Bosque opical húmedo, 20 m al u a,
15°46'59" N, 87°38'9" W, José Daniel Ri e a y Mau o Za ala cols. (4). Mex-
ico (250). Campeche. Calakmul. Rese a de la Biós e a Calakmul. VII-14
18°19'49.13"N, 89°49'20.16"W Capello, M.V. & Escoba Hdez., F., col. (25).
Chiapas. Lacanjá-Chansayab, 300 m, 6/VIII/1977, ca oña en sel a, col. B.
Kohlmann (1); Yaxchilán 12.VII.81. E. Fuen es, col. Al . 140 m. Sel a B osim-
um Nec o ampa (4). Palenque. Casa a quólogos. 17/VI/1993 G y V Hal e
Col. Bo de cebo-exc emen o 13–18 ho as (13). Oaxaca. La Nue a Espe an-
za. 3-VIII-1980 C. Hue a, G. Quin e o, E. Ri e a Cols. (2), 14-V-80 140 m g
Quin e o, C. Cas illo, M.L. Cas illo, E. Ri e a, Col. (2); San a Ma ía Chimalapa,
San F ancisco La Paz. 21.VI. 2016. cop o ampa. 17°5'33.3"N, 94°9'52.6"W
po e o 245 m Víc o Moc ezuma Col. (51); 29,VI.2016 88 m. 17°5'29.9"N,
94°6'21.8"W (37). Ve ac uz. Palma Sola, 18–22/VIII/1976, D. Edmonds y B.
Kohlmann cols. (7), 22/VI/1972, G. Hal e y P. Reyes cols. (7), 29/V/1975, P.
Reyes, R. Blume, C. Hue a col., pas izal con es os de sel a subcaduci olia
al. 320 m, cebo pescado noche-día (1); Palma Sola, Ba anca del Río Pla ana ,
22–31/VIII/1976 (3); Los Tux las, 31/VIIII/1978, Es . Biol. T op. UNAM, ampa
pescado, M.A. Mo ón col. (7); Ca e e a a Ac opan, Bocana pasando la des .
Ídolos, 20/IX/1994, L. A ellano y R. Sánchez cols., al . 240 m., sel a baja sub-
caduci olia, T ampa nec o pe manen e (30); P esidio, F. Islas, VIII/1940 (6),
IX/1940 (2); Tezonapa, F. Islas, 9/IV/1941(1). 700 m al NE de Apazapan NTP-
80 Al . 340 m 30.VII.92 Sel a Baja cadu. Luis Qui oz, col. (45); Jalcomulco 17-
Jul-90 Col. L. A ellano Sel a Baja Al 450 m NTP 80 (6). Rancho El Benjamín
Nanchi al IV-V-2016 Pas izal - B. opical pe enni olio J. Pelayo col. (76).
Diagnosis. Do sal su ace glossy g een o glossy blue; sombe head; pygid-
ium da k blue o da k g een; las abdominal en i e unicolo ed, con inuously
sombe (Fig. 3E). Body leng h 6–10 mm. Clypeus wi h ou ee h, he wo cen-
al clypeal ee h a e sligh ly cu ed a hei base (Fig. 3E). The en al oo h o
he p ono al an e ola e al bo de absen . P ono al su ace smoo h and inely
punc u ed. Eigh h ely al s ia wi h basal ca ina almos eaching he basal ely-
al hal . P escu ella imp ession a iable in dep h. Pygidium g een o blue wi h
e iden punc ua ion, bea ing sho and hick se ae; wi h a s ong basal ca ina
(Fig. 11). Male pygidial leng h/wid h a io 0.72.
151
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Rema ks. Can hon spinosus was ini ially desc ibed by on Ha old (1863)
om sou he n Mexico. In 1868 Ha old desc ibed C. speciosus based on a
single emale om Mexico oo and indica ed a ce ain closeness o his spe-
cies o C. sallei. La e , Ba es (1887: 27) men ioned C. spinosus om sou he n
Mexico and C. speciosus om Gua emala and concluded ha hey a e mos
p obably he same species. In e es ingly, Schmid (1922) makes no men ion
o hese species a all in his wo k. Bal hasa (1939a: 211) conside ed C. spe-
ciosus (Mexico, Gua emala) o be a a ie y o C. cyanellus (USA) bu made no
men ion o C. spinosus. Subsequen ly, Hal e (1961) synonymized C. specio-
sus and C. spinosus wi h C. cyanellus cyanellus. La e , Howden (1966: 727)
ollowed sui and also conside ed C. speciosus and C. spinosus as synonyms
o C. c. cyanellus. Finally, Nunes (2019: 73), in his doc o al hesis, conside ed
C. speciosus a alid axon a he species le el and C. spinosus o be a senio
synonym o C. speciosus wi hou men ioning any speci ic eason.
Nunes (2019) sepa a ed C. spinosus (as speciosus) om he o he axa be-
longing o he C. cyanellus species g oup using do sal colo , he absence o a
me a emo al pos e io ma gin, and he p ono al su ace. Fu he , C. spinosus
has a smoo h p ono al do sal su ace, while C. cyanellus has a ugous do sal
su ace. Nunes (2019) ga e a de ailed desc ip ion o he ex e nal mo phology
and male geni alia o his species and conside ed C. spinosus (as speciosus) o
be dis ibu ed om cen al Ve ac uz, Mexico o Pe én, Gua emala. Lec o ypes
we e labeled o C. speciosus and C. spinosus by Vaz-de-Mello in 2014 a he
Muséum Na ional d’His oi e Na u elle, Pa is, bu as hey ha e no been o mal-
ly designa ed hey emain syn ypes. The mo phological di e ences be ween
C. sallei and C. spinosus a e de ailed in he sec ion abo e ea ing C. sallei.
Because o hese di e ences in mo phological cha ac e s, we ele a e Can hon
spinosus back o i s o me species s a us.
We disag ee wi h Nunes (2019: ig. 16), howe e , ha he bee le popula-
ions o he Yuca an Peninsula belong o C. spinosus. This is inco ec as hey
belong o a new ela ed species ha is awai ing desc ip ion, and i di e s
om C. spinosus by ha ing a longe pygidium and a smalle size. Capello
and Hal e (2019) commen ed ha he popula ions collec ed in Calakmul,
Campeche (a he base o he Yuca an Peninsula), which hey conside ed (e -
oneously, in ou opinion) o pe ain o C. cyanellus, p esen ed wo di e en
size o ms, one big and a small one, each ac ually co esponding, espec i ely,
o C. spinosus and he new species. Bo h species mee in his locali y, wi h
he small species being he only one o ad ance u he No h in o he Yuca -
an Peninsula. Nolasco-So o e al. (2023) made he gene ic-phylogeog aph-
ic analysis o hese popula ions and eco e ed wo gene ic-phylogeog aph-
ic lines om Calakmul. One b anch o ms one lineage wi h specimens om
Calakmul, Los Chimalapas, Oaxaca, and Palenque, Chiapas, in sou heas e n
Mexico (C. spinosus conside ed as C. speciosus by Nolasco-So o e al. 2023).
In con as , ano he lineage includes indi iduals om Calakmul and Chame-
la, Jalisco, loca ed on he Paci ic coas . This makes sense because opical
deciduous o es s a e p esen in Jalisco and Yuca an, and bo h a eas likely
had a common bee le and ege a ion- ype o igin. Nolasco-So o e al. (2023)
epo ed a conside able numbe o gene ic-phylogeog aphic lineages in hei
s udy o he C. cyanellus species complex om Mexico o Colombia; how-
e e , i is impo an o in e p e hem wi h cau ion, pa icula ly in ligh o he
158
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Dicho omius enioi Mon oya & Vaz-de-Mello, 2021 (NR, CS, EZ)
Fig. 4B
Mexico, Gua emala, Hondu as.
Mon oya and Vaz-de-Mello 2021; Solís and Kohlmann 2022.
Ma e ial examined. Hondu as (58): Comayagua: Sigua epeque, Pa que Nacio-
nal Ce o Azul Meámba ; 14°52'50"N, 87°54'59"W, 712 m; 16.VII.2023; JD Ri e a
leg.; pi all ap, human dung; UNAH (15). Olancho: Ca acamas, Pa que Nacio-
nal Sie a de Agal a; 14°54'6"N, 85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and
R Salomão leg.; pi all ap, human dung; UNAH (7). Same locali y; 14°55'1"N,
85°52'57"W, 586 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human
dung; UNAH (9). Ca acamas, Aldea La Concepción; 14°45'57"N, 85°48'6"W, 320
m; 4.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH
(1). Same locali y; 14°47'2"N, 85°45'32"W, 321 m; 4.VII.2023; JD Ri e a and
R Salomão leg.; pi all ap, human dung; UNAH (6). Yo o: Olanchi o, Re ugio
Hábi a /Especie Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m;
29.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (20).
Dicho omius pisuicas Solís & Kohlmann, 2025 (CS, EZ)
Dicho omius sa anas (Ha old, 1867)
Hondu as, Nica agua, Cos a Rica, Panama.
Reyes Ma ínez and Suazo Euceda 2016; Solís and Kohlmann 2025.
Ma e ial examined. Hondu as (22): Olancho: Ca acamas, Pa que Nacional
Sie a de Agal a; 14°54'6"N, 85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R
Salomão leg.; pi all ap, human dung; UNAH (5). Same locali y; 14°55'1"N,
85°52'57"W, 586 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap,
human dung; UNAH (16). Ca acamas, Aldea La Concepción; 14°47'2"N,
85°45'32"W, 321 m; 4.VII.2023; JD Ri e a and R Salomão leg.; pi all ap,
human dung; UNAH (1).
Dicho omius yuca anus (Ba es, 1887) (CS, RL, EZ)
Fig. 4C
Pino us yuca anus Ba es, 1887.
Mexico, Hondu as, El Sal ado , Nica agua, Cos a Rica.
Ba es 1887; Howden and Young 1981; Kohlmann and Solís 1997; Kohlmann
2003; Solís and Kohlmann 2012; Maes e al. 2020.
Ma e ial examined. Hondu as (36): F ancisco Mo azán: Dis i o Cen al, Uni-
e sidad Nacional Au ónoma de Hondu as – Ciudad Uni e si a ia; 14°5'2"N,
87°11'32"W, 990 m; 20.VII.2024; JD Ri e a leg.; pi all ap, human dung; UNAH
(20). Olancho: Ca acamas, Aldea La Concepción; 14°45'57"N, 85°48'6"W, 320
m; 4.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH
(5). Same locali y; 14°47'2"N, 85°45'32"W, 321 m; 4.VII.2023; JD Ri e a and
159
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
R Salomão leg.; pi all ap, human dung; UNAH (2). Yo o: Olanchi o, Re ugio
Hábi a /Especie Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m;
29.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (9).
Eu ys e nini
Eu ys e nus angus ulus Ha old, 1869 (NR, CS, EZ)
Mexico, Gua emala, Belize, Hondu as.
on Ha old 1869; Jessop 1985; Génie 2009.
Ma e ial examined. Hondu as (3): Olancho: Ca acamas, Pa que Nacional Sie -
a de Agal a; 14°54'6"N, 85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R Salomão
leg.; pi all ap, human dung; UNAH (1). Same locali y; 14°55'1"N, 85°52'57"W;
3.VII.2023; 586 m; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (2).
Eu ys e nus ca ibaeus (He bs , 1789) (CS, RL, EZ)
Sca abaeus ca ibaeus He bs , 1789.
Eu ys e nus pe uanus Ha old, 1875; Eu ys e nus nebulosus Ki sch, 1871; Eu ys-
e nus planus Dalman, 1824.
Mexico, Gua emala, Belize, Hondu as, Nica agua, Cos a Rica, Panama, Colom-
bia, Venezuela, Guyana, Su iname, F ench Guiana, Ecuado , Pe u, B azil, Bo-
li ia, Pa aguay, A gen ina, T inidad and Tobago.
He bs 1789; Howden and Young 1981; Jessop 1985; Génie 2009; Solís and
Kohlmann 2012; Maes e al. 2020.
Ma e ial examined. Hondu as (11): Olancho: Ca acamas, Pa que Nacional
Sie a de Agal a; 14°54'6"N, 85°52′25"W, 483 m; 3.VII.2023; JD Ri e a and R
Salomão leg.; pi all ap, human dung; UNAH (5). Same locali y; 14°55'1"N,
85°52'57"W, 586 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, hu-
man dung; UNAH (6).
Eu ys e nus oedus Gué in-Méne ille, 1830 (RL, EZ)
Eu ys e nus claudicans Ki sch, 1871.
Mexico, Gua emala, Belize, Hondu as, Nica agua, Cos a Rica, Panama, Colom-
bia, Venezuela, F ench Guiana, Ecuado , Pe u, B azil, Boli ia.
Gué in-Méne ille 1844; Howden and Young 1981; Jessop 1985; Génie 2009;
C eedy and Mann 2012; Maes e al. 2020.
Eu ys e nus magnus Cas elnau, 1840 (CS, RL, EZ)
Mexico, Gua emala, Belize, Hondu as, El Sal ado , Nica agua, Cos a Rica, Panama.
Cas elnau 1840; Jessop 1985; Génie 2009; Ri e a and Can a e o 2011; C eedy
and Mann 2012; Solís and Kohlmann 2012.
160
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Ma e ial examined. Hondu as (9): Comayagua: Sigua epeque, Pa que Na-
cional Ce o Azul Meámba ; 14°52'21"N, 87°54'7"W, 993 m; 16.VII.2023; JD
Ri e a leg.; pi all ap, human dung; UNAH (1). F ancisco Mo azán: Dis-
i o Cen al, Pa que Nacional La Tig a; 14°11'38"N, 87°8'34"W, 1439 m;
22.VI.2023; JD Ri e a, R Salomão and M Michel leg.; pi all ap, human
dung; UNAH (1). Same locali y; 14°12'42"N, 87°8'51"W, 1464 m; 22.VI.2023;
JD Ri e a, R Salomão and M Michel leg.; pi all ap, human dung; UNAH
(3). Lempi a: G acias, Pa que Nacional Mon aña de Celaque; 14°33'53"N,
88°37'54"W, 1271 m; 16.VI.2023; JD Ri e a, R Salomão and M Michel leg.;
pi all ap, human dung; UNAH (2). Same locali y; 14°29'5"N, 88°40'11"W,
2030 m; 17.VI.2023; JD Ri e a, R Salomão and M Michel leg.; pi all ap,
human dung; UNAH (2).
Eu ys e nus mexicanus (Ha old, 1869) (CS, RL, EZ)
Mexico, Gua emala, Belize, Hondu as, El Sal ado , Nica agua, Cos a Rica, Pan-
ama, Colombia, Venezuela, Guyana, T inidad and Tobago.
on Ha old 1869; Jessop 1985; Génie 2009; C eedy and Mann 2012; Solís and
Kohlmann 2012; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (4): Olancho: Ca acamas, Pa que Nacional
Sie a de Agal a ; 14°54'6"N, 85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R
Salomão leg.; pi all ap, human dung; UNAH (1). Same locali y; 14°55'1"N,
85°52'57"W, 586 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap,
human dung; UNAH (1). Yo o: Olanchi o, Re ugio Hábi a /Especie Colib í
Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m; 29.VI.2023; JD Ri-
e a and R Salomão leg.; pi all ap, human dung; UNAH (1). Same locali y;
15°25'41"N, 86°53'38"W, 243 m; 30.VI.2023; JD Ri e a and R Salomão leg.;
pi all ap, human dung; UNAH (1).
Eu ys e nus obli e a us Génie , 2009 (RL)
Mexico, Gua emala, Belize, Hondu as.
Génie 2009; C eedy and Mann 2012.
Oni icellini
Euoni icellus in e medius (Reiche, 1850)** (RL, EZ)
Oni icellus in e medius Reiche, 1850; Oni icellus speciosus Cos a, 1853; Oni i-
cellus cla a us Ro h, 1851; Oni icellus pallens Lapo e, 1840.
Uni ed S a es, Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a
Rica, Panama, A ica, Aus alia.
Reiche 1852; Mon es de Oca e al. 1994; Solís and Kohlmann 2012; Ri e a and
Schlein 2014; Pablo-Cea e al. 2023.
161
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
On hophagini
Digi on hophagus gazella (Fab icius, 1787)** (RL, EZ)
Sca abaeus gazella Fab icius, 1787; On hophagus in e medius Reiche, 1840
(no es ablished; ambiguous); Cop is an ilope Fab icius, 1798; Cop is me al-
licus Fab icius, 1798; Sca abaeus do cas Oli ie , 1789.
Uni ed S a es, Hawai, Mexico, Gua emala, El Sal ado , Hondu as, Nica agua,
Panama, Colombia, Venezuela, B azil, Pe u, Boli ia, A gen ina, Pa aguay, U u-
guay, Dominican Republic, Guadeloupe, Jamaica, Angola, Bo swana, Como os,
Congo, Kenya, Madagasca , Malawi, Mozambique, Namibia, Sou h, A ica, Swa-
ziland, Tanzania, Uganda, Zambia, Zimbabwe, Aus alia, New Caledonia, Papua,
New Guinea, Vanua u.
Fab icius 1787; Bal hasa 1959; Kohlmann 1994; Ri e a and Can a e o 2011;
Génie and Mo e o 2017.
On hophagus acumina us Ha old, 1880 (NR, CS, EZ)
Mexico, Hondu as, El Sal ado , Nica agua, Cos a Rica, Panama, Colombia, Ec-
uado , Pe u, B azil.
Ha old 1880; Howden and Young 1981; Kohlmann and Solís 2001; Pulido and
Zunino 2007; Maes e al. 2020.
Ma e ial examined. Hondu as (136): Comayagua: Sigua epeque, Pa que Na-
cional Ce o Azul Meámba ; 14°50'40"N, 87°57'12"W, 748 m; 15.VII.2023; JD
Ri e a leg.; pi all ap, human dung; UNAH (1). Same locali y; 14°52'21"N,
87°54'7"W, 993 m; 16.VII.2023; JD Ri e a leg.; pi all ap, human dung; UNAH
(5). Same locali y; 14°52'50"N, 87°54'59"W, 712 m; 16.VII.2023; JD Ri e a leg.;
pi all ap, human dung; UNAH (29). G acias a Dios: B us Lagunas, Rese a
de Bios e a y el Homb e Río Plá ano; 15°41'40"N, 84°33'43"W, 9 m; 8.VII.2023;
JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (69). Same lo-
cali y; 15°42'45"N, 84°33'35"W, 13 m; 8.VII.2023; JD Ri e a and R Salomão leg.;
pi all ap, human dung; UNAH (27). Same locali y; 15°42'28"N, 84°30'22"W, 2
m; 9.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH
(1). Olancho: Ca acamas, Aldea La Concepción; 14°45'57"N, 85°48'6"W, 320 m;
4.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (4).
On hophagus an h acinus Ha old, 1873 (NR, CS, RL)
Mexico, Gua emala, Hondu as, Nica agua, Cos a Rica, Panama.
Ha old 1873; Ba es 1887; Pi ie and Biolley 1895; Howden and Ca w igh 1963;
Kohlmann and Solís 2001; Pulido and Zunino 2007; He nández e al. 2024b.
Ma e ial examined. Hondu as (1): Lempi a: G acias, Pa que Nacional Mon-
aña de Celaque; 14°33'53"N, 88°37'54"W, 1271 m; 16.VI.2023; JD Ri e a, R Sa-
lomão and M Michel leg.; pi all ap, human dung; UNAH (1).
162
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
On hophagus ba esi Howden & Ca w igh , 1963 (CS, RL, EZ)
Uni ed S a es, Mexico, Gua emala, Belize, Hondu as, El Sal ado , Nica agua,
Cos a Rica, Panama, Colombia.
Howden and Ca w igh 1963; Kohlmann and Solís 2001; Medina e al. 2001;
Pulido and Zunino 2007; Ri e a and Can a e o 2011; Solís and Kohlmann 2012.
Ma e ial examined. Hondu as (165): A lán ida: Tela, Pa que Nacional Jean-
ne e Kawas; 15°47'48"N, 87°33'21"W, 12 m; 18.VII.2024; JD Ri e a leg.; pi all
ap, human dung; UNAH (114). F ancisco Mo azán: Dis i o Cen al, Uni e sidad
Nacional Au ónoma de Hondu as – Ciudad Uni e si a ia; 14°5'2"N, 87°11'32"W,
990 m; 20.VII.2024; JD Ri e a leg.; pi all ap, human dung; UNAH (35). G acias
a Dios: B us Lagunas, Rese a de Bios e a y el Homb e Río Plá ano; 15°42'45"N,
84°33'35"W, 13 m; 8.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, hu-
man dung; UNAH (3). Olancho: Ca acamas, Pa que Nacional Sie a de Agal a;
14°54'6"N, 85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all
ap, human dung; UNAH (2). Ca acamas, Aldea La Concepción; 14°45'57"N,
85°48'6"W, 320 m; 4.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human
dung; UNAH (2). Same locali y; 14°47'2"N, 85°45'32"W, 321 m; 4.VII.2023; JD
Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (9).
On hophagus b e iconus Génie & Howden, 1999* (RL, EZ)
Gua emala, Hondu as.
Génie and Howden 1999; C eedy and Mann 2012.
On hophagus championi Ba es, 1887 (CS, RL, EZ)
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica.
Ba es 1887; Howden and Ca w igh 1963; Kohlmann and Solís 2001; Pulido
and Zunino 2007; Ri e a and Can a e o 2011; Solís and Kohlmann 2012; Moc-
ezuma and Hal e 2021b.
Ma e ial examined. Hondu as (103): Yo o: Olanchi o, Re ugio Hábi a /Especie
Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m; 29.VI.2023; JD
Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (15). Same locali y;
15°27'9"N, 86°53'41"W, 338 m; 29.VI.2023; JD Ri e a and R Salomão leg.; pi all
ap, human dung; UNAH (34). Same locali y; 15°25'41"N, 86°53'38"W, 243 m;
30.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (16).
Same locali y; 15°25'57"N, 86°52'31"W, 255 m; 30.VI.2023; JD Ri e a and R Sa-
lomão leg.; pi all ap, human dung; UNAH (38).
On hophagus c ini us Ha old, 1869 (NR, CS, EZ)
Mexico, Gua emala, Belize, Hondu as, El Sal ado , Nica agua, Cos a Rica, Pan-
ama, Colombia.
163
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
on Ha old 1869; Howden and Young 1981; Kohlmann and Solís 2001; Medina
e al. 2001; Pulido and Zunino 2007; Solís and Kohlmann 2012; Maes e al.
2020; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (68): G acias a Dios: B us Lagunas, Rese a de
Bios e a y el Homb e Río Plá ano; 15°41'40"N, 84°33'43"W, 9 m; 8.VII.2023; JD
Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (64). Same locali y;
15°42'45"N, 84°33'35"W, 13 m; 8.VII.2023; JD Ri e a and R Salomão leg.; pi all
ap, human dung; UNAH (4).
On hophagus cyanellus Ba es, 1887 (CS, RL, EZ)
On hophagus mesoame icanus Zunino & Hal e , 1988.
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica, Panama.
Ba es 1887; Pi ie and Biolley 1895; Kohlmann and Solís 2001; Pulido and Zun-
ino 2007; C eedy and Mann 2012; Kohlmann and Solís 2001, 2012; Maes e al.
2020; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (2): F ancisco Mo azán: Dis i o Cen al, Pa que
Nacional La Tig a; 14°12'42"N, 87°8'51"W, 1464 m; 22.VI.2023; JD Ri e a, R Sa-
lomão and M Michel leg.; pi all ap, human dung; UNAH (2).
On hophagus gazellinus Ba es, 1887 (NR, EZ)
Hondu as, Nica agua, Cos a Rica.
Ba es 1887; Kohlmann and Solís 2001, 2012; Maes e al. 2020.
On hophagus hoep ne i Ha old, 1869 (CS, RL, EZ)
On hophagus a izonensis Schae e , 1909.
Uni ed S a es, Mexico, El Sal ado , Hondu as, Nica agua, Cos a Rica.
on Ha old 1869; Howden and Ca w igh 1963; Kohlmann and Solís 2001; Ri e-
a and Can a e o 2011; Maes e al. 2020; Pablo-Cea e al. 2023.
On hophagus incensus Say, 1835 (CS, RL, EZ)
Uni ed S a es, Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a
Rica, Panama, Colombia, Venezuela, Ecuado .
Say 1835; Ba es 1887; Pi ie and Biolley 1895; Howden and Ca w igh 1963;
Kohlmann and Solís 2001; Pulido and Zunino 2007; Ri e a and Can a e o 2011;
Solís and Kohlmann 2012; Maes e al. 2020; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (13): F ancisco Mo azán: Dis i o Cen al,
Pa que Nacional La Tig a; 14°11'38"N, 87°8'34"W, 1439 m; 22.VI.2023; JD Ri-
e a, R Salomão and M Michel leg.; pi all ap, human dung; UNAH (5). Same
164
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
locali y; 14°12'42"N, 87°8'51"W, 1464 m; 22.VI.2023; JD Ri e a, R Salomão and
M Michel leg.; pi all ap, human dung; UNAH (6). Lempi a: G acias, Pa que
Nacional Mon aña de Celaque; 14°33'53"N, 88°37'54"W, 1271 m; 16.VI.2023; JD
Ri e a, R Salomão and M Michel leg.; pi all ap, human dung; UNAH (1). Same
locali y; 14°29'5"N, 88°40'11"W, 2030 m; 17.VI.2023; JD Ri e a, R Salomão and
M Michel leg.; pi all ap, human dung; UNAH (1).
On hophagus landol i Ha old, 1880 (CS, RL, EZ)
On hophagus exanus Schae e , 1914.
Uni ed S a es, Mexico, Gua emala, Belize, Hondu as, El Sal ado , Nica agua,
Cos a Rica, Panama, Colombia, Venezuela.
Ha old 1880; Pi ie and Biolley 1895; Howden and Ca w igh 1963; Howden and
Young 1981; Medina e al. 2001; Pulido and Zunino 2007; Ri e a and Can a e o
2011; Solís and Kohlmann 2012; Capello and Hal e 2019; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (888): F ancisco Mo azán: Dis i o Cen al, Uni-
e sidad Nacional Au ónoma de Hondu as – Ciudad Uni e si a ia; 14°5'2"N,
87°11'32"W, 990 m; 20.VII.2024; JD Ri e a leg.; pi all ap, human dung; UNAH
(20). Lempi a: G acias, Pa que Nacional Mon aña de Celaque; 14°29'5"N,
88°40'11"W, 2030 m; 17.VI.2023; JD Ri e a, R Salomão and M Michel leg.; pi all
ap, human dung; UNAH (1). Yo o: Olanchi o, Re ugio Hábi a /Especie Colib í
Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m; 29.VI.2023; JD Ri e-
a and R Salomão leg.; pi all ap, human dung; UNAH (220). Same locali y;
15°27'9"N, 86°53'41"W, 338 m; 29.VI.2023; JD Ri e a and R Salomão leg.; pi -
all ap, human dung; UNAH (309). Same locali y; 15°25'41"N, 86°53'38"W, 243
m; 30.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH
(114). Same locali y; 15°25'57"N, 86°52'31"W, 255 m; 30.VI.2023; JD Ri e a and
R Salomão leg.; pi all ap, human dung; UNAH (224).
On hophagus longimanus Ba es, 1887 (RL)
Mexico, Gua emala, Belize, Hondu as.
Ba es 1887; Pulido and Zunino 2007; C eedy and Mann 2012.
On hophagus nyc opus Ba es, 1887 (NR, EZ)
Hondu as, Nica agua, Cos a Rica, Colombia, Pe ú.
Ba es 1887; Kohlmann and Solís 2001; Medina e al. 2001; He nández e al.
2024b.
On hophagus pe enensis Howden & Gill, 1993 (NR, RL)
Mexico, Gua emala, Belize, Hondu as.
Howden and Gill 1993; Pulido and Zunino 2007; C eedy and Mann 2012.
165
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
On hophagus p aecellens Ba es, 1887 (NR, CS)
Gua emala, Hondu as, Nica agua, Cos a Rica, Panama, Colombia, T inidad and
Tobago.
Ba es 1887; Pi ie and Biolley 1895; Kohlmann and Solís 2001; Medina e al.
2001; Pulido and Zunino 2007; Solís and Kohlmann 2012; Maes e al. 2020.
Ma e ial examined. Hondu as (6): Comayagua: Sigua epeque, Pa que Nacio-
nal Ce o Azul Meámba ; 14°52'21"N, 87°54'7"W, 993 m; 16.VII.2023; JD Ri e a
leg.; pi all ap, human dung; UNAH (6).
On hophagus hinolophus Ha old, 1869 (CS, RL, EZ)
Mexico, Gua emala, Hondu as, Nica agua.
on Ha old 1869; Ba es 1887; Pulido and Zunino 2007; Capello and Hal e
2019; Maes e al. 2020.
Ma e ial examined. Hondu as (22): G acias a Dios: B us Lagunas, Rese a
de Bios e a y el Homb e Río Plá ano; 15°41'40"N, 84°33'43"W, 9 m; 8.VII.2023;
JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (1). Olancho:
Ca acamas, Pa que Nacional Sie a de Agal a; 14°54'6"N, 85°52'25"W, 483 m;
3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (6).
Same locali y; 14°55'1"N, 85°52'57"W, 586 m; 3.VII.2023; JD Ri e a and R Sa-
lomão leg.; pi all ap, human dung; UNAH (15).
On hophagus sal ado ensis Zunino & Hal e , 1988 (NR, CS)
Fig. 4D
Hondu as, El Sal ado .
Zunino and Hal e 1988; Pulido and Zunino 2007; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (2): Lempi a: G acias, Pa que Nacional Mon-
aña de Celaque; 14°33'41"N, 88°39'8"W, 1622 m; 16.VI.2023; JD Ri e a, R Sa-
lomão and M Michel leg.; pi all ap, human dung; UNAH (2).
Rema ks. As in he case o Haplogeo upes gua emalensis iden a us,
Howden collec ed his species o he i s ime a 2300 m a.s.l. in Ce o
Mon e C is o in El Sal ado in 1971. Fi y- h ee yea s elapsed be o e we col-
lec ed i again. I is also he i s ime his species has been collec ed in
Hondu as. As in he case o he geo upids, his species was collec ed in he
Pa que Nacional de Mon aña Celaque a 1560 m a.s.l. This is he sou he n-
mos known eco d o bo h species.
Acco ding o Zunino and Hal e (1988), his species belongs o he O. cya-
nellus species line, which is conside ed an elemen o ancien no he n o igin.
I s di e si ica ion is belie ed o ha e occu ed in ancien geological imes, as
e idenced by i s dis inc mo phology, dis ibu ion, and ecological equi emen s
(Zunino and Hal e 1988). This p oposal aligns wi h he o igin o he cloud
o es habi a whe e his species esides. The cloud o es is belie ed o ha e
166
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
an ancien Oligocene-Miocene o igin (Rzedowski 1996). This ege a ion ype
has also been ecognized as an impo an cen e o plan and animal di e si i-
ca ion, as well as an endemism cen e (Flo es and Na a o 1993).
On hophagus sp.* (RL)
Hondu as.
C eedy and Mann 2012.
On hophagus s ockwelli Howden & Young, 1981 (NR, EZ)
Hondu as, Nica agua, Cos a Rica, Panama, Colombia, Ecuado .
Howden and Young 1981; Kohlmann and Solís 2001; Pulido and Zunino 2007;
Solís and Kohlmann 2012; He nández e al. 2024b.
On hophagus i idi inosus Kohlmann & Solis, 2001 (NR, CS)
Hondu as, Nica agua, Cos a Rica.
Kohlmann and Solís 2001; Pulido and Zunino 2007; Solís and Kohlmann 2012;
He nández e al. 2024b.
Ma e ial examined. Hondu as (11): G acias a Dios: B us Lagunas, Rese a de
Bios e a y el Homb e Río Plá ano; 15°42'45"N, 84°33'35"W, 13 m; 8.VII.2023;
JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (4). Olancho:
Ca acamas, Pa que Nacional Sie a de Agal a; 14°55'1"N, 85°52'57"W, 586 m;
3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (7).
On hophagus yuca anus Delgado, Pe aza & Deloya, 2006 (NR, CS)
Fig. 4E
Mexico, Gua emala, Belize, Hondu as.
Delgado e al. 2006; Pulido and Zunino 2007; Capello and Hal e 2019.
Ma e ial examined. Hondu as (23): A lán ida: Tela, Los Olingos Lodge;
15°46'59"N, 87°38'9"W, 20 m; 9.XI.2024; JD Ri e a leg.; pi all ap, human dung;
UNAH (4). Comayagua: Sigua epeque, Pa que Nacional Ce o Azul Meámba ;
14°52'50"N, 87°54'59"W, 712 m; 16.VII.2023; JD Ri e a leg.; pi all ap, human dung;
UNAH (9). Olancho: Ca acamas, Pa que Nacional Sie a de Agal a; 14°54'6"N,
85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human
dung; UNAH (1). Same locali y; 14°55'1"N, 85°52'57"W, 586 m; 3.VII.2023; JD Ri-
e a and R Salomão leg.; pi all ap, human dung; UNAH (5). Yo o: Olanchi o, Re-
ugio Hábi a /Especie Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412
m; 29.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (4).
167
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Phanaeini
Cop ophanaeus bouca di (Ne inson, 1891) (RL, EZ)
Phanaeus bouca di Ne inson, 1891; Cop ophanaeus plu o bouca di (Ne inson);
A naud 2002.
Mexico, Hondu as, El Sal ado , Nica agua, Cos a Rica.
Ne inson 1891; Edmonds and Zídek 2010; Sánchez-He nández e al. 2019.
Cop ophanaeus co y hus (Ha old, 1863) (CS, RL, EZ)
Phanaeus co y hus Ha old, 1863; Phanaeus pe seus Ha old, 1880; Phanae-
us elamon co y hus (Ha old); Cop ophanaeus elamon co y hus (Ha old);
Cop ophanaeus elamon ne insoni A naud and Gámez 2002; Pe ei a and
Ma ínez 1956; A naud 2002.
Mexico, Gua emala, Belize, Hondu as, El Sal ado , Nica agua, Cos a Rica, Pan-
ama, Colombia, Venezuela.
Ha old 1863; Edmonds and Zídek 2010; C eedy and Mann 2012; Solís and Kohl-
mann 2012; Maes e al. 2020; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (50): Comayagua: Sigua epeque, Pa que Na-
cional Ce o Azul Meámba ; 14°50'40"N, 87°57'12"W, 748 m; 15.VII.2023; JD
Ri e a leg.; pi all ap, human dung; UNAH (2). Same locali y; 14°52'50"N,
87°54'59"W, 712 m; 16.VII.2023; JD Ri e a leg.; pi all ap, human dung;
UNAH (2). Same locali y; 14°51'45"N, 87°55'48"W, 730 m; 15.VII.2023; JD
Ri e a leg.; pi all ap, human dung; UNAH (4). G acias a Dios: B us Lagu-
nas, Rese a de Bios e a y el Homb e Río Plá ano; 15°41'40"N, 84°33'43"W,
9 m; 8.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung;
UNAH (9). Same locali y; 15°42'45"N, 84°33'35"W, 13 m; 8.VII.2023; JD Ri e-
a and R Salomão leg.; pi all ap, human dung; UNAH (12). Same locali y;
15°42'28"N, 84°30'22"W, 2 m; 9.VII.2023; JD Ri e a and R Salomão leg.; pi -
all ap, human dung; UNAH (1). Same locali y; 15°42'26"N, 84°32'33"W, 8
m; 8.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH
(14). Olancho: Ca acamas, Pa que Nacional Sie a de Agal a; 14°55'1"N,
85°52'57"W, 586 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap,
human dung; UNAH (4). Ca acamas, Aldea la Concepción; 14°47'2"N,
85°45'32"W, 321 m; 4.VII.2023; JD Ri e a and R Salomão leg.; pi all ap,
human dung; UNAH (2).
Cop ophanaeus gilli A naud, 1997 (RL, EZ)
Cop ophanaeus hen yi Malý & Poko ný, 2008.
Mexico, Gua emala, Belize, Hondu as.
A naud 1997; A naud 2002b; Edmonds and Zídek 2010; C eedy and Mann 2012;
Kohlmann and Solís 2012.
174
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Va ia ion. Males. Leng h 20.2–23.6 mm. Wid h 11.9–14.9 mm. Females.
Leng h 20.6–22.1 mm. Wid h 12.4–14.1 mm.
E ymology. The name is dedica ed in hono o he Hondu an na ional he o
and symbol o libe y and cul u al esis ance, who was a cap ain o he Lenca
ibes ha ough he Spanish in asion and conques .
Habi a . This species is dis ibu ed be ween 1528 and 2060 m a.s.l. Pine
and pine-oak associa ion a e dis ibu ed in his pa k be ween 1000 and 1500
m a.s.l. on he A lan ic slope, and be ween 1800 and 2300 m a.s.l. on he
Paci ic slope (Mejía O dóñez and House 2002). Six o he se en pine species
o he coun y occu he e, Pinus ooca pa Schiede ex Schl dl., 1838 being he
Figu e 12. A. Pho o o he emale me abasi a sus o Phanaeus melampus. Texas A&M
Uni e si y Insec Collec ion, ex F. Pso a collec ion, ex Field Museum, ex Da id Edmonds
collec ion. Pho o cou esy o Ma io Cupello, no locali y label. B. Pho o o he emale
me abasi a sus o Phanaeus lempi a sp. no . Hondu as. Pa que Nacional de Mon aña
Celaque. 14,484138°N, -88,668799°W.
Figu e 11. Las abdominal s e ni e o Can hon spinosus A. Male; B. Female, which a e
sombe and unicolo ous. Pygidium o C. spinosus showing se ae C. Male; D. Female.
175
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
mos common one. Mixed o es (pine, Liquidamba s y aci lua L., 1753, Cle-
h a sp., Nec and a sp.) is eco ded highe han 1500 m a.s.l. and be ween
1800 and 2000 m a.s.l. he cloud o es begins on he A lan ic slope o he
pa k, while on he Paci ic slope i s a s a ca 2300 m a.s.l. The cloud o es
is cha ac e ized by a signi ican plan species ichness (Abies gua emalensis
Rehde , 1939, Alnus sp., Pinus pseudos obus Lindl., 1839, P unus sp., Que -
cus lanci olia Schl dl. & Cham, 1830, Taxus globosa Schl dl., 1838), whe e ee
e ns a e e y common. I is he highes moun ain ange in Hondu as. This
pa k is home o se e al endemic species o plan s, amphibians, sh ews, a
sca ab gold bee le (Ch ysina pas o i Cu oe, 1994), and a sca ab lowe bee le
(A chedinus an oshkai Seidel and A iaga-Va ela, 2018).
Dis ibu ion. Phanaeus lempi a is closely ela ed o he Mexican endemic Ph.
melampus Ha old, 1863. Edmonds (1994) eco ded he la e om h ee local-
i ies in Ve ac uz and wo in Chiapas, and ano he Chiapas locali y was added
by Na a e e-Gu ié ez and Edmonds (2006). The nine Na a e e-Gu ié ez and
Edmonds specimens we e collec ed a 145 m a.s.l. du ing Feb ua y, Augus ,
and Oc obe in aps bai ed wi h human eces in he Mon es Azules Biosphe e
Rese e in all and medium e e g een opical o es s. Upon eques , Na a e-
e-Gu ié ez p o ided us wi h pho os o hese specimens. Upon close exam-
ina ion, hese indi iduals showed all he cha ac e is ics o he new Phanaeus
species desc ibed in his pape . E en hough we did no see Edmonds’s (1994)
Chiapas specimens, hey also come om places co e ed in we opical e -
e g een o es , El Bosque and Tenejapa, a 1460 m a.s.l., indica ing hey also
mos p obably belong o his new species. Finally, among he ma e ial pho o-
g aphed by Ma io Cupello om he Texas A&M Uni e si y collec ion, he e is
one specimen wi h he cha ac e is ics o Ph. lempi a ha bea s a label say-
ing only “Tabasco, Messico”. This specimen migh ha e been collec ed a ound
Teapa, whe e Tabasco eaches he piedmon o he Chiapas moun ains. All his
sugges s ha Ph. lempi a is dis ibu ed om Chiapas o Hondu as (Fig. 13),
whe eas Ph. melampus is dis ibu ed yonde he Is hmus o Tehuan epec di ide
– cu en ly known exclusi ely om Ve ac uz – and no sou h o i .
Liza do e al. (2022: suppl. ig. 1), on p edic ed and eco ded dis ibu ions o
Phanaeus p esen s a map o he expec ed dis ibu ion o he Phanaeus ame-
hys inus species g oup o which his species is a membe . These clima ically
p edic ed dis ibu ions a e somewha a odds wi h he ac ual eco ded dis ibu-
ions. Fo example, hei model p edic s a possible dis ibu ion o Ph. melam-
pus (p obable sis e species o Ph. lempi a) in he ma shy opical sa annah
o no heas e n Hondu as, whe e no species o Phanaeus has been egis e ed
a p esen and whe e he majo i y o he collec ed species belong o he olle
guild and some On hophagus axa as well. These clima ic models usually make
dis ibu ion p edic ions ha a e way o he ma k in opical moun ainous a -
eas. As Kohlmann e al. (2007) ha e al eady discussed, ine -scale sys ems, like
he Li e Zones o Hold idge (1967), a e be e sui ed o hese opical a eas.
Phenology. This species was collec ed in June, July, and No embe using
aps bai ed wi h human eces.
Taxonomic ema ks. This species is closely ela ed o Ph. melampus, epo -
ed om a moun ain we opical e e g een o es om Ve ac uz in Mexico col-
lec ed a 1500 m a.s.l., and o Ph. gua emalensis Ha old, 1871, epo ed om
o es s and adjacen open habi a s in he Gua emalan Highlands, dis ibu ed
176
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
om 1500 o 2400 m (Edmonds 1994). The nea es eco ded species g oup
specimen o he ype locali y o his new species is Ph. gua emalensis, 230 km
away, sugges ing a g ea geog aphical sepa a ion.
The new species Ph. lempi a can be dis inguished om Ph. melampus
by he shape o he smoo h, ans e se basal a ea o he male p ono um,
which ea u es a sulcus a he mid-base o he p ono um. In Ph. lempi a, his
a ea is sligh ly iangula ly a ched (Fig. 6), esembling he shape obse ed
in Ph. ame hys inus. In con as , Ph. melampus exhibi s a s ongly iangula ,
punc a e a ea ha lacks he sulcus a he base o he medial oo h (Fig.
14A). Addi ionally, he in e s iae o Ph. melampus a e dull and a ely show
a bluish cas , whe eas Ph. lempi a has shiny in e s iae wi h coppe y- ed e-
lec ions. Female specimens o Ph. melampus ha e a semici cula p ono al
ans e se ca ina, while Ph. lempi a ea u es a mo e open ans e se ca ina
(Figs 7, 14B). The emale o Ph. lempi a has a me abasi a sus widened api-
cally, and he ou e idge o he en al su ace is dis inc ly mo e sinuous
han he inne idge, simila o Ph. ame hys inus (Fig. 12B). Con e sely, he
emale o Ph. melampus has a na owe me abasi a sus wi h bo h en al
idges unning mo e o less s aigh (Fig. 12A).
Phanaeus lempi a can be sepa a ed om Ph. gua emalensis by he in e s i-
ae, la o weakly con ex in Ph. lempi a, aised mid-longi udinally, appea ing
cos a e in Ph. gua emalensis. The p ono oum o Ph. lempi a emales ha e as-
pe a e punc u es pos e omedially, whe eas in Ph. gua emalensis he a ea is
coa sely punc a e o ugo-punc a e, appea ing oughened.
Wi h his new species, he numbe o axa in he ame hys inus species g oup
(ame hys inus, blackalle i, gua emalensis, melampus) inc eases o i e. In he
wo k o Liza do e al. (2022) Phanaeus genie i, a membe o he ame hys inus
species g oup, is conside ed in hei dis ibu ion p edic ion analysis, al hough
his species has no ye been eins a ed as a alid species. Gille and Toussain
Figu e 13. Map o he dis ibu ion o Phanaeus lempi a sp. no . (s a s) and Ph. gua emalensis (do s). T iangles ep esen
specimens ha we e conside ed ini ially o belong o Ph. melampus by Na a e e-Gu ié ez and Edmonds (2006), bu ha
on pho og aphic inspec ion show he cha ac e is ics o Ph. lempi a.
177
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
(2020), based on molecula gene ic analysis, conside ed ha his species
g oup appea ed du ing he Miocene a ound 17 Ma. Howe e , mode n species
o ma ion seems o ha e been ela i ely ecen du ing he Pleis ocene, ~ 1 Ma.
Key o species o he ame hys inus g oup (modi ied om Edmonds
and Zídek 2012)
1 Punc u es o pos e omedial po ion o emale p ono um simple o weakly
aspe a e, weakes in on o basal ossae, a ea appea ing smoo h o un-
aided eye. Ely a da k, sombe ; me allic colo , i any, es ic ed o ma gins;
p ono um usually da k excep o me allic blue la e al ma gins ................2
– Pos e omedial po ion o emale p ono um coa sely punc a e o ugo-punc-
a e, a ea appea ing oughened o unaided eye. Ely a in mos specimens
wi h me allic colo ma ching ha o p ono um and pygidium .................... 4
2(1) Pos e omedial po ion o emale p ono um wi h simple punc u es (×10).
Female p ono um bea ing a weakly i- ube cula e, s aigh ans e se
idge whose leng h clea ly exceeds in e ocula dis ance; idge no ol-
lowed by a dis inc conca i y. Head ho n o la ge male ape ing g adually
om base o apex, base no dis inc ly swollen. Ely al in e s iae dis inc ly
con ex, weakly shiny. Smoo h, ans e se, iangula a ea along pos e i-
o ma gin o male p ono um lacking a ube cle on medial angle. Smalle ,
leng h 14–20 mm, usually < 20 mm. Paci ic slope o Sie a Mad e del Su
in Gue e o and Oaxaca ........... Phanaeus (P. ) blackalle i Delgado-Cas illo
– Pos e omedial po ion o emale p ono um wi h inely aspe a e punc u es
(×10). Head ho n o la ge male dis inc ly swollen basally. Ely al in e s iae
la o only weakly con ex, dull. T ans e se, iangula a ea along pos e io
Figu e 14. Habi us Phanaeus melampus A. Male; B. Female. Ve ac uz, O izaba, Mexico.
Ins i u oyale des Sciences na u elles de Belgique, h ough Roge Damoiseau. Pho os
cou esy o W. D. Edmonds.
178
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
ma gin o male p ono um wi h an acu e ube cle on medial angle. La ge ,
leng h 18–28 mm, usually well o e 20 mm ................................................3
3(2) Smoo h, sligh ly iangula ly a ched basal a ea o male p ono um ea u es
a sulcus a base o medial oo h (Fig. 6), esembling he shape obse ed
in Ph. ame hys inus. In e s iae wi h coppe y- ed e lec ions. Female spec-
imens ea u e an open ans e se ca ina (Fig. 7). Female o wi h me ab-
asi a sus widened apically, ou e idge o en al su ace dis inc ly mo e
sinuous han inne idge, simila o Ph. ame hys inus (Fig. 12B) .................
....................................................................... Phanaeus (P. ) lempi a sp. no .
– Punc a e a ea o highly iangula basal a ea o male p ono um lacking sul-
cus a base o medial oo h (Fig. 14A). In e s iae dull and a ely showing
bluish cas . Female specimens ha e a semici cula p ono al ans e se ca i-
na (Fig. 14B). Female wi h na owe me abasi a sus wi h bo h en al idges
unning mo e o less s aigh (Fig. 12A) ......Phanaeus (P. ) melampus Ha old
4(1) A leas in e s iae 2 and 3, bu in mos specimens 2–5, highly shiny and
aised mid-longi udinally, dull adjacen o s iae, appea ing cos a e o
unaided eye. Sides o p ono um densely g anula e- ugose. Highlands o
Gua emala and adjacen Chiapas ..... Phanaeus (P. ) gua emalensis Ha old
– All ely al in e s iae e enly and in mos specimens b igh ly shiny, e enly
con ex. Sides o p ono um only mode a ely densely g anulo- ugose. High-
lands o cen al Chiapas, moun ains o eas e n Oaxaca, Sie a Mad e O i-
en al om Ve ac uz o sou he n Tamaulipas ................................................
............................................................... Phanaeus (P. ) ame hys inus Ha old
Phanaeus po ioni A naud, 2001 (CS, RL, EZ)
Fig. 5B, C
Belize, Hondu as.
A naud 2001; Edmonds and Zídek 2012; Moc ezuma and Hal e 2021a.
Ma e ial examined. Hondu as (11): F ancisco Mo azán: Dis i o Cen al,
Pa que Nacional La Tig a; 14°12'42"N, 87°8'51"W, 1464 m; 22.VI.2023; JD Ri e-
a, R Salomão and M Michel leg.; pi all ap, human dung; UNAH (1). Lempi a:
G acias, Pa que Nacional Mon aña de Celaque; 14°33'53"N, 88°37'54"W, 1271
m; 16.VI.2023; JD Ri e a, R Salomão and M Michel leg.; pi all ap, human dung;
UNAH (3). Olancho: Ca acamas, Pa que Nacional Sie a de Agal a; 14°54'6"N,
85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, hu-
man dung; UNAH (5). Same locali y; 14°55'1"N, 85°52'57"W, 586 m; 3.VII.2023;
JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (2).
Phanaeus wagne i Ha old, 1863 (CS, RL, EZ)
Fig. 5D
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica.
Ha old 1863; Ba es 1887; Pi ie and Biolley 1895; Edmonds 1994; A naud
2002a; Edmonds and Zídek 2012; Solís and Kohlmann 2012; Maes e al. 2020;
Pablo-Cea e al. 2023.
179
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Ma e ial examined. Hondu as (6): Lempi a: G acias, Pa que Nacional Mon aña
de Celaque; 14°29'5"N, 88°40'11"W, 2030 m; 17.VI.2023; JD Ri e a, R Salomão
and M Michel leg.; pi all ap, human dung; UNAH (2). Yo o: Olanchi o, Re ugio
Hábi a /Especie Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m;
29.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (4).
Sulcophanaeus ch yseicollis (Ha old, 1863) (NR, CS, EZ)
Fig. 5E
Phanaeus ch yseicollis Ha old, 1863.
Mexico, Hondu as.
Ha old 1863; Ba es 1887; Edmonds 2000.
Ma e ial examined. Hondu as (9): Olancho: Ca acamas, Pa que Nacional Sie a
de Agal a; 14°54'6"N, 85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R Salomão
leg.; pi all ap, human dung; UNAH (4). Same locali y; 14°55'1"N, 85°52'57"W, 586
m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (5).
Sisyphini
Sisyphus mexicanus Ha old, 1863 (NR, CS, RL)
Fig. 5F
Mexico, Hondu as, El Sal ado , Nica agua, Cos a Rica.
Ha old 1863; Ba es 1887; Haa 1955; Howden 1966; Hal e 2003; Ri e a and
Can a e o 2011; Solís and Kohlmann 2012; Capello and Hal e 2019; Maes e
al. 2020; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (535): Yo o: Olanchi o, Re ugio Hábi a /Especie
Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m; 29.VI.2023; JD
Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (232). Same locali y;
15°27'9"N, 86°53'41"W, 338 m; 29.VI.2023; JD Ri e a and R Salomão leg.; pi all
ap, human dung; UNAH (199). Same locali y; 15°25'41"N, 86°53'38"W, 243 m;
30.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (61).
Same locali y; 15°25'57"N, 86°52'31"W, 255 m; 30.VI.2023; JD Ri e a and R Sa-
lomão leg.; pi all ap, human dung; UNAH (43).
Rema ks. This species has been conside ed in he pas o be a ypical
inhabi an o opical ain o es s. Fo ins ance, Hal e (2003: 44) indica es
ha i is no an abundan species and ha i inhabi s e e g een and semide-
ciduous opical o es s and seconda y communi ies in Mexico. Howe e ,
one o he coau ho s (BK) collec ed i in 1976 in Palma Sola, Ve ac uz, olling
cow dung balls, in e es ingly only on pa hs de oid o ege a ion, a e sing
in oduced opical g ass. Recen wo k con i ms he la e obse a ion and
sugges s he ea lie iew is inco ec . Al a ado e al. (2018) epo a sig-
ni ican numbe o S. mexicanus in hei analysis (12,628 specimens o he
169,372 o al), his species being he i h mos nume ous one a e (in de-
c easing o de ) Can hon (C.) chiapas (ci ed as C. (C.) indigaceus che ola i),
On hophagus landol i, Pseudocan hon pe plexus, and Can hon (Gl.) eu yscelis.
180
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Mos o he specimens we e ound in d y opical o es agmen s in he
no h-cen al egion o he Yuca an Peninsula.
The p esen wo k a i es a a simila conclusion. Sisyphus mexicanus has been
caugh in signi ican numbe s in Olanchi o, an a ea co e ed by opical d y o es .
This species is he hi d mos nume ous, a e On hophagus landol i and Can hon
(C.) chiapas. The ou h and i h mos nume ous species a e Can hon (Gl.) champi-
oni and On hophagus ba esi. As in he p e ious example, i is an ecosys em domina -
ed mainly by ball- olling dung bee les. All his sugges s ha Sisyphus mexicanus is
a denizen o he land-locked Cen al Ame ican opical d y o es . This species has
been epo ed as a e in all o he a eas (Sou h-Eas e n Mexico, El Sal ado , Nica a-
gua, Cos a Rica) (Solís and Kohlmann 2012; Maes e al. 2020; Pablo-Cea e al. 2023).
Haa (1955), in his s udy o he genus Sisyphus, sugges ed ha S. c ispa us Go y,
1833 is o med by h ee subspecies, S. c. c ispa us om Sou h and Wes A ica,
S. c. hi us Wiedemann, 1823 om India and S i Lanka, and. S. c. mexicanus Ha old,
1863 om Mexico. Haa (1955) conside ed hem all membe s o one species be-
cause all h ee lack any spines o p ojec ions on he me a emo a and me a ibiae in
he male specimens. Pablo-Cea e al. (2023) ollowed his axonomic a angemen
in hei ca alog o he dung bee le auna o El Sal ado , a axonomic use which
we deem inco ec . Haa (1955) sepa a ed S. c. c ispa us om S. c. mexicanus
because he las subspecies ha e much s onge punc ua ion and mo e poin -
ed clypeal ee h han he i s ones. Paschalidis (1974), in he M.Sc. hesis, syn-
onymized S. c ispa us wi h S. cos a us (Thunbe g, 1818), a Sou h A ican species
li ing in mois sa annas and upland g asslands. Howe e , Paschalidis (1974) made
no men ion o any o he subspecies, as Haa (1955) had conside ed in his s udy.
Subsequen ly, Daniel e al. (2018) conside ed S. c ispa us as a nomen dubium. On
he o he hand, S. mexicanus has been ea ed as a alid species in all Ame ican
sys ema ic s udies o his axon (Howden 1965; Hal e 2003). Mo eo e , Daniel e
al. (2018) a e no con iden in s a ing which o he wo species (S. ca e Boheman,
1857, S. so didus Boheman, 1857) migh be he possible junio synonym o S. c is-
pa us. As Daniel e al. (2018: 21) indica e ha he ype specimen o S. c ispa us
has been los and as he o iginal desc ip ion and image p esen ed a e imp ecise
and insu icien o de e mine he iden i y o he species, we hen conside he e Si-
syphus mexicanus as a alid axon a he species le el and no as a subspecies o a
nomen dubium. Finally, his decision is suppo ed by he pho og aphic compa ison
o he A ican and Ame ican species (Daniel e al. 2018: ig. 5B, F). Sisyphus ca e
and S. so didus ha e dense and longe do sal se ae co e han S. mexicanus; bo h
A ican species ha e a ounde ho ax and comple ely ounded an e io p ono al
hume i, whe eas he Ame ican species has a less ounded ho ax and e y poin ed
an e io p ono al hume i; he A ican species ha e den ed me a ibiae, whe eas he
Ame ican species is uly se a ed. These mo phological di e ences make i clea
ha he A ican and Ame ican species a e no conspeci ic, as Haa (1955) had
sugges ed, and as such S. mexicanus canno be a subspecies o S. c ispa us.
Sca abaeinae ince ae sedis
Can hidium a dens Ba es, 1887 (CS, RL)
Mexico, Gua emala, Hondu as, Nica agua, Cos a Rica, Panama.
181
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Ba es 1887; C eedy and Mann 2012; Pi ie and Biolley 1895; Howden and
Young 1981; Solís and Kohlmann 2004; Kohlmann and Solís 2006; Solís and
Kohlmann 2012; Cupello 2018.
Ma e ial examined. Hondu as (9): Comayagua: Sigua epeque, Pa que Nacio-
nal Ce o Azul Meámba ; 14°52'21"N, 87°54'7"W, 993 m; 16.VII.2023; JD Ri e a
leg.; pi all ap, human dung; UNAH (3). Olancho: Ca acamas, Pa que Nacional
Sie a de Agal a; 14°55'1"N, 85°52'57"W, 586 m; 3.VII.2023; JD Ri e a and R
Salomão leg.; pi all ap, human dung; UNAH (6).
Can hidium cen ale Boucomon , 1928 (CS, RL)
Can hidium ma inezi Edmonds & Hal e , 1978.
Mexico, Gua emala, Hondu as, Nica agua, Cos a Rica, Panama, Colombia, Su i-
name, F ench Guiana, Ecuado .
Boucomon 1928; Howden and Young 1981; Medina e al. 2001; Solís and Kohl-
mann 2004; Kohlmann and Solís 2006; C eedy and Mann 2012; Solís and Kohl-
mann 2012; Cupello 2018.
Ma e ial examined. Hondu as (22): Comayagua: Sigua epeque, Pa que Na-
cional Ce o Azul Meámba ; 14°53'33"N, 87°51'21"W, 720 m; 15.VII.2023; JD
Ri e a leg.; pi all ap, human dung; UNAH (8). Same locali y; 14°50'40"N,
87°57'12"W, 748 m; 15.VII.2023; JD Ri e a leg.; pi all ap, human dung;
UNAH (3). Same locali y; 14°52'50"N, 87°54'59"W, 712 m; 16.VII.2023; JD Ri-
e a leg.; pi all ap, human dung; UNAH (10). Yo o: Olanchi o, Re ugio Háb-
i a /Especie Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m;
29.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (1).
Can hidium lae um Ha old, 1867 (CS, RL, EZ)
Can hidium g ani o um Hal e & Hal e , 1978.
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica.
Ha old 1867a; Ba es 1887; Solís and Kohlmann 2004; Kohlmann and Solís
2006; Solís and Kohlmann 2012; Cupello 2018; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (6): F ancisco Mo azán: Dis i o Cen al, Uni-
e sidad Nacional Au ónoma de Hondu as – Ciudad Uni e si a ia; 14°5'2"N,
87°11'32"W, 990 m; 20.VII.2024; JD Ri e a leg.; pi all ap, human dung;
UNAH (5). Yo o: Olanchi o, Re ugio Hábi a /Especie Colib í Esme alda Hon-
du eño; 15°27'53"N, 86°52'52"W, 412 m; 29.VI.2023; JD Ri e a and R Sa-
lomão leg.; pi all ap, human dung; UNAH (1).
Can hidium mo oni Kohlmann & Solís, 2006 (EZ)
Kohlmann 2003, 2004, included as pa o C. espe inum.
182
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Mexico, Gua emala, Hondu as.
Kohlmann and Solís 2006; Cupello 2018; Capello and Hal e 2019; Sán-
chez-He nández e al. 2022.
Can hidium ube i ons Howden & Young, 1981 (NR, EZ)
Hondu as, Nica agua, Cos a Rica, Panama.
Howden and Young 1981; Ma ínez and Hal e 1986; Solís and Kohlmann
2004; 2012, Cupello 2018; He nández e al. 2020.
Can hidium espe inum Howden & Young, 1981 (EZ)
Mexico, Hondu as, Cos a Rica, Panama.
Howden and Young 1981; Solís and Kohlmann 2004, 2012; Cupello 2018.
C yp ocan hon sp.* (RL)
Hondu as.
C eedy and Mann 2012.
U oxys bone i Pe ei a & Hal e , 1961 (RL)
U oxys biden is Howden & Young, 1981.
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica, Panama, Co-
lombia, Ecuado .
Pe ei a and Hal e 1961; Delgado and Kohlmann 2007; Solís and Kohlmann
2012; Solís and Kohlmann 2013.
U oxys bonplandi Kohlmann, Solís & Al a ado, 2019 (NR, EZ)
Hondu as, Cos a Rica.
Kohlmann, Solís and Al a ado 2019.
U oxys dea ilai Delgado & Kohlmann, 2007 (NR, CS, EZ)
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica.
Delgado and Kohlmann 2007; Solís and Kohlmann 2012, 2013; Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (77): Yo o: Olanchi o, Re ugio Hábi a /Especie
Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m; 29.VI.2023; JD
Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (41). Same locali y;
15°27'9"N, 86°53'41"W, 338 m; 29.VI.2023; JD Ri e a and R Salomão leg.; pi all
ap, human dung; UNAH (36).
183
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
U oxys mic ocula is Howden & Young, 1981 (NR, CS, EZ)
Mexico, Gua emala, Hondu as, El Sal ado , Nica agua, Cos a Rica, Panama.
Howden and Young 1981; Delgado and Kohlmann 2007; Solís and Kohlmann
2012; 2013, Pablo-Cea e al. 2023.
Ma e ial examined. Hondu as (65): A lán ida: Tela, Pa que Nacional Jean-
ne e Kawas; 15°52'0"N, 87°37'0"W, 100 m; 11.X.2024; JD Ri e a leg.; pi all ap,
human dung; UNAH (15). Comayagua: Sigua epeque, Pa que Nacional Ce o
Azul Meámba ; 14°50'40"N, 87°57'12"W, 748 m; 15.VII.2023; JD Ri e a leg.; pi -
all ap, human dung; UNAH (3). Same locali y; 14°52'21"N, 87°54'7"W, 993 m;
16.VII.2023; JD Ri e a leg.; pi all ap, human dung; UNAH (24). Same locali y;
14°52'50"N, 87°54'59"W, 712 m; 16.VII.2023; JD Ri e a leg.; pi all ap, human
dung; UNAH (4). F ancisco Mo azán: Dis i o Cen al, Pa que Nacional La Tig a;
14°11'38"N, 87°8'34"W, 1439 m; 22.VI.2023; JD Ri e a, R Salomão and M Michel
leg.; pi all ap, human dung; UNAH (2). Same locali y; 14°12'42"N, 87°8'51"W,
1464 m; 22.VI.2023; JD Ri e a, R Salomão and M Michel leg.; pi all ap, human
dung; UNAH (2). Dis i o Cen al, Uni e sidad Nacional Au ónoma de Hondu as –
Ciudad Uni e si a ia; 14°5'2"N, 87°11'32"W; 20.VII.2024; 990 m; JD Ri e a leg.; pi -
all ap, human dung; UNAH (7). Olancho: Ca acamas, Pa que Nacional Sie a de
Agal a; 14°54'6"N, 85°52'25"W, 483 m; 3.VII.2023; JD Ri e a and R Salomão leg.;
pi all ap, human dung; UNAH (6). Same locali y; 14°55'1"N, 85°52'57"W, 586
m; 3.VII.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (2).
U oxys mic os Ba es, 1887 (CS, EZ)
Mexico, Gua emala, Hondu as, Nica agua, Cos a Rica, Panama, Colombia.
Ba es 1887; Howden and Young 1981; Delgado and Kohlmann 2007; Solís and
Kohlmann 2012, 2013.
Ma e ial examined. Hondu as (54): Comayagua: Sigua epeque, Pa que Nacio-
nal Ce o Azul Meámba ; 14°50'40"N, 87°57'12"W, 748 m; 15.VII.2023; JD Ri e a
leg.; pi all ap, human dung; UNAH (3). Same locali y; 14°52'21"N, 87°54'7"W,
993 m; 16.VII.2023; JD Ri e a leg.; pi all ap, human dung; UNAH (21). Same
locali y; 14°52'50"N, 87°54'59"W, 712 m; 16.VII.2023; JD Ri e a leg.; pi all ap,
human dung; UNAH (6). F ancisco Mo azán: Dis i o Cen al, Pa que Nacional
La Tig a; 14°11'38"N, 87°8'34"W, 1439 m; 22.VI.2023; JD Ri e a, R Salomão and
M Michel leg.; pi all ap, human dung; UNAH (6). Lempi a: G acias, Pa que
Nacional Mon aña de Celaque; 14°33'53"N, 88°37'54"W, 1271 m; 16.VI.2023; JD
Ri e a, R Salomão and M Michel leg.; pi all ap, human dung; UNAH (4). Same
locali y; 14°29'5"N, 88°40'11"W, 2030 m; 17.VI.2023; JD Ri e a, R Salomão
and M Michel leg.; pi all ap, human dung; UNAH (1). Olancho: Ca acamas,
Pa que Nacional Sie a de Agal a; 14°55'1"N, 85°52'57"W, 586 m; 3.VII.2023;
JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (1). Ca acamas,
Aldea la Concepción; 14°45'57"N, 85°48'6"W, 320 m; 4.VII.2023; JD Ri e a and
R Salomão leg.; pi all ap, human dung; UNAH (2). Yo o: Olanchi o, Re ugio
Hábi a /Especie Colib í Esme alda Hondu eño; 15°27'53"N, 86°52'52"W, 412 m;
29.VI.2023; JD Ri e a and R Salomão leg.; pi all ap, human dung; UNAH (10).
190
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Figu e 20. Reco ded locali ies o A. Dicho omius pisuicas; B. Dicho omius yuca anus; C. Digi on hophagus gazella; D. Euo-
ni icellus in e medius; E. Eu ys e nus angus ulus; F. Eu ys e nus ca ibaeus; G. Eu ys e nus oedus; H. Eu ys e nus magnus;
I. Eu ys e nus mexicanus in Hondu as.
Figu e 21. Reco ded locali ies o A. Eu ys e nus obli e a us; B. Haplogeo upes gua emalensis iden a us; C. Malagoniella
yuca eca; D. Mega hoposoma candezei; E. Neoa hy eus exca a us; F. On hophagus acumina us; G. On hophagus an h aci-
nus; H. On hophagus ba esi; I. On hophagus b e iconus in Hondu as.
191
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Figu e 22. Reco ded locali ies o A. On hophagus championi; B. On hophagus c ini us; C. On hophagus cyanellus;
D. On hophagus gazellinus; E. On hophagus hoep ne i; F. On hophagus incensus; G. On hophagus landol i; H. On hophagus
longimanus; I. On hophagus nyc opus in Hondu as.
Figu e 23. Reco ded locali ies o A. On hophagus pe enensis; B. On hophagus p aecellens; C. On hophagus hinolophus;
D. On hophagus sal ado ensis; E. On hophagus sp.; F. On hophagus s ockwelli; G. On hophagus i idi inosus; H. On hoph-
agus yuca anus; I. Phanaeus endymion in Hondu as.
192
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Figu e 24. Reco ded locali ies o A. Phanaeus excelsus; B. Phanaeus eximius; C. Phanaeus lempi a sp. no .; D. Phanaeus
po ioni; E. Phanaeus wagne i; F. Pseudocan hon pe plexus; G. Sca imus o a us; H. Sisyphus mexicanus; I. Sulcophanaeus
ch yseicollis in Hondu as.
Figu e 25. Reco ded locali ies o A. Scybalocan hon monilia us; B. Syl ican hon aequinoc ialis; C. U oxys bone i; D. U oxys
bonplandi; E. U oxys dea ilai; F. U oxys mic ocula is; G. U oxys mic os; H. U oxys nebulinus in Hondu as.
193
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Acknowledgemen s
We wan o exp ess ou hea el g a i ude o all he indi iduals and o ganiza-
ions in Hondu as who suppo ed us h oughou his p ojec . Ou hanks go
o Ma yu i Alemán A ilez, Nicolás Zúñiga, He mes Vega, and he MAPANCE
PROCELAQUE non-p o i o ganiza ion in Celaque; Jo ge Luis Palma and F an-
cisco Vásquez om AMITIGRA; and C is ian Wild om he En omological
Museum a he Uni e sidad Nacional Au ónoma de Hondu as (UNAH-CU) o
hei assis ance in La Tig a Na ional Pa k. We also app ecia e he in aluable
ield suppo p o ided by Ke in He nandez, Ulises So o, Nahum Munguía,
and Se gio Unau o ASIDE Hondu as in Yo o, as well as Diego, Ca he ine, and
Cin hia in Olancho. Special hanks go o Ma ía José Bú om he Hyd obi-
ology Labo a o y (UNAH-CU), Nohemi Wood Michel, Odai Rod íguez Wood,
Alex Suazo and Eudo o Po illo Cu ly o hei dedica ed help du ing ou ield-
wo k in B us Laguna. We u he hank Napoleón Mo azán and he Wildli e
Conse a ion Socie y (WCS – Hondu as-Nica agua) o hei logis ical sup-
po du ing ou s ay in B us Laguna. We ex end ou hea el g a i ude o he
Te i o ial Council o he Mosqui ia, D apa Ta a Ra Uplika Nani Asla Takanka
(DIUNAT), o g an ing us pe mission o wo k on hei ances al lands. We a e
g a e ul o G ecia Daniela Na a o, Gene al Héc o Benjamín Vale io A dón,
and he mili a y pe sonnel om he P ime Ba allón de P o ección Ambien al
who accompanied us in Olancho, as well as o I o Al a ado, Gus a o Cas o,
Edga do O ellana, Ke in He nández, and Key i Díaz om Aldea SOS o hei
suppo in PANACAM. We also hank he Ins i u o de Conse ación Fo es al
(ICF) o hei aluable logis ical assis ance wi h he esea ch. Addi ionally,
we hank F ançois Génie om he Canadian Museum o Na u e, O awa, o
pho og aphing he holo ype specimen o On hophagus sal ado ensis. A u o
González kindly p o ided us wi h pho og aphs o Can hon cyanellus species
g oup specimens om Colombia in he collec ion o Alejand o Lope a. We
a e also indeb ed o Da ío Na a e e o sha ing pho os o he new Phanaeus
species housed in he insec collec ion a ECOSUR, Chiapas, Mexico, and o
W. D. Edmonds o p o iding se e al pho og aphs o Phanaeus melampus and
Phanaeus gua emalensis om his p e ious publica ions. Ou since e hanks
go o Ma io Cupello, cu a o a he Texas A&M Uni e si y Insec Collec ion, o
sha ing pho og aphs and label in o ma ion o Phanaeus melampus. We also
app ecia e he gene osi y o Jesús O ozco and Ka e in Aguila in allowing us
o e iew he insec collec ion a Escuela Ag ícola Paname icana El Zamo a-
no and lending he pho og aphic equipmen . We a e e y g a e ul o Vi idiana
Vega and Ede Fa id Mo a om he IEXA Insec Collec ion “D . Miguel Ángel
Mo ón Ríos” in Xalapa, Mexico, o hei in aluable assis ance in acili a ing
BK’s s udy o specimens in hei collec ion. Finally, we ex end ou deepes
hanks o Ha y B ailo sky and C is ina Mayo ga om he Na ional Insec Col-
lec ion (CNIN), Ins i u e o Biology, UNAM, Mexico, o hei in aluable sup-
po du ing BK’s isi and s udy o ma e ial housed in hei collec ion. Las bu
no leas , we a e in deb o Ma io Cupello and And ey F olo o hei excellen
and hough ul e iews o his ex ha g ea ly imp o ed i . JDRD g a e ully
acknowledges he Consejo Nacional de Humanidades, Ciencia y Tecnología
(CONAHCYT) and he Di ección Gene al de Asun os del Pe sonal Académico
(DGAPA-UNAM) o hei suppo h ough a pos doc o al schola ship.
194
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
No unding was epo ed.
Au ho con ibu ions
Concep ualiza ion: JDRD. Da a cu a ion: ÁS, MM, JDRD, BK. Funding acquisi ion: JDRD.
In es iga ion: MM, BK, MIM, RPS. P ojec adminis a ion: JDRD. Resou ces: MIM.
Valida ion: MM, ÁS. Visualiza ion: BK, MIM, JDRD, MM. W i ing – o iginal d a : RPS, BK,
MM, ÁS, JDRD, MIM.
Au ho ORCIDs
José D. Ri e a-Dua e h ps://o cid.o g/0000-0001-7610-8446
Rena o P. Salomão h ps://o cid.o g/0000-0001-9826-7472
Ángel Solís h ps://o cid.o g/0000-0001-8622-4555
Mau icio Michel h ps://o cid.o g/0009-0009-2331-6810
Ma cela I. Ma amo os h ps://o cid.o g/0000-0002-2928-206X
Be Kohlmann h ps://o cid.o g/0000-0002-3441-3933
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex .
Re e ences
Al a ado F, Escoba F, Williams DR, A oyo-Rod íguez V, Escoba -He nández F (2018) The ole
o li es ock in ensi ica ion and landscape s uc u e in main aining opical biodi e si y.
Jou nal o Applied Ecology 55(1): 185–194. h ps://doi.o g/10.1111/1365-2664.12957
Al a ado López FM, Oso io Ka an FJ (2020) Coleóp e os nec ó agos, cop ó agos y
ugí o os en la co dille a El Me endón, Hondu as. BSc Thesis, Escuela Ag ícola Pan-
ame icana.
Amadon D (1967) The supe species concep . Sys ema ic Zoology 15(3): 245–249.
h ps://doi.o g/10.2307/2411398
And esen E, U ea-Galeano LA (2022) E ec s o dung bee le ac i i y on opical o -
es plan s. F on ie s in Ecology and E olu ion 10: 979676. h ps://doi.o g/10.3389/
e o.2022.979676
A naud P (1997) Desc ip ion de nou elles espèces du gen e Cop ophanaeus Ols. Be-
soi o 4: 4–8.
A naud P (2001) Desc ip ion de nou elles espèces de Phanaeides. Besoi o 6: 2–8.
A naud P (2002a) Phanaeini. Vol. 28. Hillside Books, Can e bu y, 151 pp.
A naud P (2002b) Desc ip ion de nou elles espèces de Phanaeides. Besoi o 7: 2–9.
195
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
A iaga-Jiménez A, Zunino M, Hal e G, Escoba -He nández F, Rossini M (2023) A
axonomic e iew o he endemic Mexican genus Ce a o upes Jekel, 1865 (Coleop-
e a: Geo upidae: Ce a o upini), wi h he desc ip ion o an unexpec ed new species
om he Sie a Mad e del Su . Eu opean Jou nal o Taxonomy 885: 1–20. h ps://doi.
o g/10.5852/ej .2023.885.2183
Aseso a S de RL (2009) Plan de Manejo del Á ea Hábi a / Especie Colib í Esme alda
Hondu eño Amazilia Luciae, 347 pp.
Bal hasa V (1939a) Eine Vo s udie zu Monog aphie de Ga ung Can hon H sg. Folia
Zoologica e Hyd obiologica 9: 179–238.
Bal hasa V (1939b) Neue A en de Ga ung Sca imus E . Fes sch i zum 60. Gebu -
s age on P o esso D . Emb ik S and 5: 87–91. [Riga.]
Bal hasa V (1951) Sca abaeidae Lapa os ic ae: Bei ag zu Fauna Pe us. S udien-Ins i-
u , Jena 2: 322–344.
Bal hasa V (1959) Bei ag zu Kenn nis de Ga ung On hophagus La . Sbo ník en omo-
logického Oddelení Ná odního Musea P aze 33: 461–471.
Ba es HW (1887) Insec a. Coleop e a. Vol. II. Pa 2. Pec inico nia and Lamellico nia.
25–160. In: Biologia Cen ali-Ame icana. Taylo and F ancis, London, 432 pp.
Ba es HW (1889) Insec a. Coleop e a. Vol. II. Pa 2. Pec inico nia and Lamellico nia.
337–416. In: Biologia cen ali Ame icana. Taylo and F ancis, London, 432 pp.
Blackwelde R (1944) Checklis o he Coleop e ous insec s o Mexico, Cen al Ame ica
he Wes Indies and Sou h Ame ica. Pa 2. Bulle in - Uni ed S a es Na ional Museum
185: 189–306. h ps://doi.o g/10.3406/bse .1928.28008
Boheman CH (1858) Coleop e a, in Kongliga S enska F ega en Eugenies Resa Om-
k ing Jo den unde Be al a C.A.Vi gin, A en 1851–53. Ve enskapliga Iak agelse
Pa H.M.K. Osca den Fo s es. Kongliga S enska Ve enskapsakademien Handlinga ,
S ockholm, 1–617.
Boucomon A (1928) Cop ophages d’Ame ique du Sud nou eaux ou peu connus. Bulle-
in de la Socié é En omologique de F ance 13(13): 202–207. h ps://doi.o g/10.3406/
bse .1928.28014
B owne D, Schol z CH (2002) A phylogeny o he amilies o Sca abaeoidea (Coleop e a). Sys-
ema ic En omology 24(1): 51–84. h ps://doi.o g/10.1046/j.1365-3113.1999.00067.x
Bu meis e HCC (1848) Die En wicklungsgeschich e de Ga ung Del ochilum. Zei ung
ü Zoologie. Zoo omie und Palaeozoologie 1: 133–136.
Capello V, Hal e G (2019) Lis ado ilus ado de las especies de Sca abaeinae (Cole-
op e a: Sca abaeidae) de la Rese a de la Biós e a de Calakmul, Campeche, México.
Dugesiana 26(2): 103–131. h ps://doi.o g/10.32870/dugesiana. 26i2.7080
Cas elnau FL (1840) His oi e Na u elle des Insec es Coléop è es A ec une in oduc-
ion en e man l’Ana omie e la Physiologie des Animaux A iculés, pa M. B ullé. P.
Duménil, Pa is, 21–564.
Ce i os Ga cía DG, Ochoa Cadena LA (2017) Ca ac e ización de la comunidad de es-
ca abajos (Coleop e a: Sca abaeidae) asociados al es ié col en Zamo ano, Hondu-
as. BSc Thesis, Escuela Ag ícola Paname icana Zamo ano.
C eedy TJ, Mann DJ (2012) Iden i ica ion Guide o he Sca abaeinae Dung Bee les o Cu-
suco Na ional Pa k, Hondu as. Ope a ion Wallacea, Conse a ion esea ch h ough
academic pa ne ship, Lincolnshi e, Uni ed Kingdom, 52 pp.
Cupello MJ (2018) On he ype species o he New Wo ld dung bee le genus Can hid-
ium E ichson, 1847 (Coleop e a: Sca abaeidae: Sca abaeinae), wi h an anno a ed
checklis o species. Zoo axa 4388(4): 451–486. h ps://doi.o g/10.11646/zoo -
axa.4388.4.1
196
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Cupello MJ (2022) A euchus dung bee les and hei e olu ion (Coleop e a: Sca abaei-
dae): a new model in speciology. PhD Thesis, Pa aná, Cu i iba.
Cupello M, Ribei o-Cos a CS, Vaz-de-Mello FZ (2022) The e olu ion o Bolbi es oni oides
(Coleop e a: Sca abaeidae: Phanaeini): i s phylogene ic signi icance, geog aphical
polych oma ism and he subspecies p oblem. Zoological Jou nal o he Linnean So-
cie y 194(3): 973–1034. h ps://doi.o g/10.1093/zoolinnean/zlab015
Cupello M, Sil a FAB, Vaz-de-Mello FZ (2023) The Taxonomic Re olu ion o New Wo ld
dung bee les (Coleop e a: Sca abaeidae: Sca abaeinae). F on ie s in Ecology and
E olu ion 11: 1168754. h ps://doi.o g/10.3389/ e o.2023.1168754
Ću čić S, Pa iće ić D, Rado ić D, Veso ić N, Bekchie R, Ću čić N, Gueo guie B (2019)
Cu en and p edic ed dis ibu ion o he a e and h ea ened bee le Bolbelasmus
(Bolbelasmus) unico nis (Coleop e a: Geo upidae) in Se bia. Eu opean Jou nal o
En omology 116: 413–424. h ps://doi.o g/10.14411/eje.2019.042
Daniel GM, Da is ALV, Sole CL, Schol z CH (2018) Taxonomic e iew o he ibe Sisyph-
ini sensu s ic o Mulsan , 1842 (Coleop e a: Sca abaeidae: Sca abaeinae) in sou h-
e n A ica, including new species desc ip ions. Insec Sys ema ics & E olu ion 51(1):
1–6. h ps://doi.o g/10.1163/1876312X-00002195
Da ling JDG, Génie F (2018) Re ision o he axonomy and dis ibu ion o he Neo-
opical Cop is ince us species complex. Canadian En omologis 150(5): 539–577.
h ps://doi.o g/10.4039/ ce.2018.32
Delgado L, Kohlmann B (2001) A new species and wo subspecies o Cop is om Mex-
ico and Cen al Ame ica (Coleop e a: Sca abaeidae: Sca abaeinae). Jou nal o he
New Yo k En omological Socie y 109(3 & 4): 344–353. h ps://doi.o g/10.1664/0028
-7199(2001)109[0344:ANSATS]2.0.CO;2
Delgado L, Kohlmann B (2007) Re isión de las especies del géne o U oxys Wes wood de
México y Gua emala. Folia En omologica Mexicana 46: 1–35.
Delgado L, Pe aza L, Deloya C (2006) On hophagus yuca anus, a new species o he
clypea us g oup om Mexico and Gua emala. The Flo ida En omologis 89(1): 6–9.
h ps://doi.o g/10.1653/0015-4040(2006)89[6:OYANSO]2.0.CO;2
Edmonds WD (1994) Re ision o Phanaeus Macleay, a New Wo ld genus o Sca abaeine
dung bee les (Coleop e a: Sca abaeidae, Sca abaeinae). Con ibu ions in Science
443: 1–105. h ps://doi.o g/10.5962/p.208079
Edmonds WD (2000) Re ision o he Neo opical dung bee le genus Sulcophanaeus
(Coleop e a: Sca abaeidae: Sca abaeinae). Folia Hey o skyana (Supplemen um
6): 1–60.
Edmonds WD, Zídek J (2010) A axonomic e iew o he Neo opical genus Cop ophanae-
us Olsou ie , 1924 (Coleop e a: Sca abaeidae, Sca abaeinae). Insec a Mundi 129:
1–111.
Edmonds WD, Zídek J (2012) Taxonomy o Phanaeus e isi ed: Re ised keys o and
commen s on species o he New Wo ld dung bee le genus Phanaeus MacLeay, 1819
(Coleop e a: Sca abaeidae: Sca abaeinae: Phanaeini). Insec a Mundi 274: 1–108.
Emlen DJ, Philips TK (2006) Phylogene ic e idence o an associa ion be ween unnel-
ing beha io and he e olu ion o ho ns in dung bee les (Coleop e a: Sca abaeidae:
Sca abaeinae). Coleop e is s Socie y Monog aph Numbe 5: 47–56. h ps://doi.o g/
10.1649/0010-065X(2006)60[47:PEFAAB]2.0.CO;2
Engle EA (2020) Composi ion o dung bee le communi ies in a opical mon ane o es
al e s he a e o dung emo al mo e han species di e si y alone. W igh S a e Uni-
e si y, 57 pp.
197
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
Fab icius JC (1787) Man issa insec o um sis ens species nupe de ec as adiec is syn-
onymis, obse a ionibus, desc ip ionibus, emenda ionibus. Tome II. Ch is ian Go lieb
P o , Copenhagen, Denma k, 382 pp.
Fa ila ME, Hal e G (1997) The use o indica o g oups o measu ing biodi e si y as
ela ed o communi y s uc u e and unc ion. Ac a Zoologica Mexicana Nue a Se ie
0: 1–25. h ps://doi.o g/10.21829/azm.1997.72721734
Fa ila ME, No iega JA, da Sil a PG, Ko asaki V (2023) Edi o ial: Neo opical dung bee le
di e si y: ecological, his o ical, and an h opogenic pe spec i es. F on ie s in Ecology
and E olu ion 11: 1294471. h ps://doi.o g/10.3389/ e o.2023.1294471
Finche GT, S ewa TB, Hun e JS III (1983) The 1981 dis ibu ion o On hophagus ga-
zella Fab icius om eleases in Texas and On hophagus au us Sch ebe om an
unknown elease in Flo ida (Coleop e a: Sca abaeidae). Coleop e is s Bulle in 37:
159–163. h ps://www.js o .o g/s able/4008007
Flo es O, Na a o A (1993) Un análisis de los e eb ados e es es endémicos de Me-
soamé ica en México. Re is a de la Sociedad Mexicana de His o ia Na u al XLIV:
387–395. [edición especial]
Gacha ná GC (1951) Ca álogo de los coleóp e os colombianos. Re is a de la Academia
Colombiana de Ciencias Exac as, Físicas y Na u ales 8: 221–229.
Gemminge M, on Ha old E (1869) Ca alogus coleop e o um hucusque desc ip o um
synonymicus e sys ema icus. Tome IV: Sca abaeidae. Sump u E. H. Gummi, Mona-
co, 380 pp. h ps://doi.o g/10.5962/bhl. i le.9089
Génie F (2009) Genus Eu ys e nus Dalman, 1824 (Sca abaeidae: Sca abaeinae: Oni i-
cellini), axonomic e ision and iden i ica ion keys illus a ed (Vol. 85). Penso Pub-
lishe s, So ia-Moscow, 430 pp.
Génie F (2012) A new species and no es on he subgenus Del ochilum (Del ochilum)
Eschschol z, 1822 (Coleop e a: Sca abaeidae: Sca abaeinae: Del ochilini). Zoo axa
3357(1): 25–36. h ps://doi.o g/10.11646/zoo axa.3357.1.2
Génie F, Howden HF (1999) Two new Cen al Ame ican On hophagus La eille o he
mi abilis species g oup (Coleop e a: Sca abaeidae, Sca abaeinae). Coleop e is s Bul-
le in 53: 130–144.
Génie F, Mo e o P (2017) Digi on hophagus Bal hasa , 1959: axonomy, sys ema ics,
and mo phological phylogeny o he genus e ealing an A ican species complex
(Coleop e a: Sca abaeidae: Sca abaeinae). Zoo axa 4248(1): 1–110. h ps://doi.
o g/10.11646/zoo axa.4248.1.1
Gill BD (1991) Dung bee les in opical Ame ican o es s. 211–229. In: Hanski I, Cambe-
o Y (Eds) Dung bee le ecology. P ince on Uni e si y P ess; P ince on, NJ, 481 pp.
Gille JJE (1910) Espèces nou elles du gen e Cop is e ele é synonymique des espèces
déc i es. No es Leyden Museum 32: 1–31.
Gille JJE (1911) Sca abaeidae: Cop inae I. Coleop e o um Ca alogus. Pa s 38. Ku ü s-
endamm, Be lin, 100 pp.
Gille CP, Toussain EF (2020) Mac oe olu ion and shi s in he eeding biology o he
New Wo ld sca ab bee le ibe Phanaeini (Coleop e a: Sca abaeidae: Sca abaeinae).
Biological Jou nal o he Linnean Socie y. Linnean Socie y o London 130(4): 661–
682. h ps://doi.o g/10.1093/biolinnean/blaa058
González A, Vaz-de-Mello FZ (2015) Taxonomic e iew o he subgenus Hybomidium
Shipp 1897 (Coleop e a: Sca abaeidae: Sca abaeinae: Del ochilum). Annales de la
Socié é En omologique de F ance 50(3–4): 431–476. h ps://doi.o g/10.1080/0037
9271.2014.989178
198
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
González-Tokman D, Esqui el-Román A, Ma ínez MI (2024) Physiology, ecology, and
e olu ion o a success ul colonize : The ho ned dung bee le, Euoni icellus in e -
medius. En omologia Expe imen alis e Applica a 172(7): 581–589. h ps://doi.
o g/10.1111/eea.13439
Go y HL (1833) Monog aphie du gen e Sisyphus. Méquignon-Ma is Pè e e Fils, Pa is,
15 pp.
Gué in-Méne ille FE (1844) Iconog aphie du Règne Animal de G. Cu ie . Pa 7. Insec es.
Balliè e, Pa is (1829–1844) 7: 1–576.
Haa E (1955) Übe die Ga ung Sisyphus La . (Col. Sca ab.). En omologische A bei en
aus dem Museum G. F ey 6: 341–381.
Hal e G (1961) Monog a ia de las especies no eame icanas del géne o Can hon.
Ciencia. Re is a Hispano-Ame icana de Ciencias Pu as y Aplicadas 20: 22–320.
Hal e G (2003) Sub ibu Can honina. 22–43. In: Mo ón MA (Ed.) A las de los esca a-
bajos de México. Coleop e a: Lamellico nia (Vol. II) Familias Sca abaeidae, T ogidae,
Passalidae y Lucanidae. A gania Edi io, Ba celona, 226 pp.
Hal e G, Edmonds WD (1982) The Nes ing Beha io o Dung Bee les (Sca abaeibae).
An Ecological and E olu i e App oach. Publicación 10. Ins i u o de ecología, México,
D.F., 176 pp.
Hal e G, Ma ínez A (1977) Re isión monog á ica de los Can honina Ame icanos IV.
Cla e pa a géne os y subgéne os. Folia En omologica Mexicana 38: 29–107.
Hanski I, Cambe o Y (1991) Dung Bee le Ecology. P ince on: P ince on Uni e si y
P ess, 514 pp. h ps://doi.o g/10.1515/9781400862092
on Ha old E (1869) No e su quelques Cop ides de Mexique. Annales de la Socié é
En omologique de F ance. Pa is (Sé ie 4) 9: 49–51.
He bs JFW (1789) Na u sys em alle bekann en in und ausländischen Insek en, als eine
Fo se zung de on Bü onschen Na u geschich e. Nach dem Sys em des Ri e s
on Linné und Fab icius zu bea bei en ange angen on Ca l Gus a Jablonsky on Jo-
hann F ied ich Wilhem He bs . De Kä e zwey e Theil. Joachim Pauli, Be lin, 330 pp.
He nández B, Maes J-M, Solís A (2020) Sca abaeinae (Sca abaeidae) de Ce o de Jesús
(Nue a Sego ia, Nica agua) y edescub imien o de Cop is maesi Ra cli e. Re is a
Nica aguense En omologica 176: 1–48.
He nández B, Solís Á, Zeas L, Lemus J (2024a) Las especies de A euchus de Nica agua
con el egis o nue o de A euchus e e i Kohlmann, 1996 (Sca abaeidae: Sca abaein-
ae). Re is a Nica aguense En omologica 316: 1–13.
He nández B, Solís Á, Lemus J, Rod íguez O, Núñez A, Pé ez S, Owa d P (2024b) Las
especies de On hophagus de Nica agua (Sca abaeidae: Sca abaeinae) con cua o
nue os egis os pa a la en omo auna nica aguense. Re is a Nica aguense de En o-
lología 319: 1–23.
Hielkema AJ (2017) Some co ec ions and ema ks ega ding he nomencla u e o Neo-
opical A hy eini, Passalini, Phanaeini, Ru elini, Cyclocephalini, Dynas ini and O yc i-
ni. Insec a Mundi 561: 1–18. h ps://doi.o g/10.5281/zenodo.5168803
Hold idge LR (1967) Li e Zone Ecology. T opical Science Cen e , San José, Cos a Rica,
149 pp.
Ho gan F (2002) Shady ield bounda ies and he coloniza ion o dung by cop ophagous
bee les in Cen al Ame ican pas u es. Ag icul u e, Ecosys ems & En i onmen 91(1–
3): 25–36. h ps://doi.o g/10.1016/S0167-8809(01)00261-4
Ho gan F (2008) Dung bee le assemblages in o es s and pas u es o El Sal ado : A
unc ional compa ison. Biodi e si y and Conse a ion 17(12): 2961–2978. h ps://
doi.o g/10.1007/s10531-008-9408-2
199
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
House PR (2004) In o me sob e el colib í esme alda en Queb ada Seca, A enal. Moni-
o eo ecológico colib í esme alda. Unidad de Cuencas SANAA.
Howden HF (1964) The Geo upinae o No h and Cen al Ame ica. Memoi s o he En-
omological Socie y o Canada 39(S39): 1–91. h ps://doi.o g/10.4039/en m9639
Howden HF (1965) A second New Wo ld species o Sisyphus La eille (Coleop e a: Sca abaei-
dae). Canadian En omologis 97(8): 842–844. h ps://doi.o g/10.4039/En 97842-8
Howden HF (1966) No es on Can honini o he “Biologia Cen ali Ame icana” and de-
sc ip ions o new species. Canadian En omologis 98(7): 725–741. h ps://doi.
o g/10.4039/En 98725-7
Howden HF (1974) Addi ional eco ds and desc ip ions o No h and Cen al Ame ica Geo-
upinae. Canadian Jou nal o Zoology 52: 567–573. h ps://doi.o g/10.1139/z74-073
Howden HF (1982) La al and adul cha ac e s o F ickius Ge main, i s ela ionship o he
Geo upini, and a phylogeny o some majo axa in he Sca abaeoidea (Insec a: Cole-
op e a). Canadian Jou nal o Zoology 60(11): 2713–2724. h ps://doi.o g/10.1139/
z82-347
Howden HF (1985a) A e ision o he Aus alian bee le gene a Bolboleaus Howden
and Coope , Blackbolbus Howden and Coope , and Bolbo hachium Boucomon
(Sca abaeidae: Geo upinae). Aus alian Jou nal o Zoology. Supplemen a y Se ies
111(111): 1–179. h ps://doi.o g/10.1071/AJZS111
Howden HF (1985b) A e ision o he Sou h Ame ican species in he genus Neoa hy eus
Howden and Ma ínez (Coleop e a: Sca abaeidae: Geo upinae). Con ibu ions o he
Ame ican En omological Ins i u e 21: 1–95.
Howden H, Ca w igh L (1963) Sca ab bee les o he genus On hophagus La eille no h
o Mexico (Coleop e a: Sca abaeidae). P oceedings o he Uni ed S a es Na ional
Museum 114(3467): 1–133. h ps://doi.o g/10.5479/si.00963801.114-3467.1
Howden HF, Gill BD (1984) Two new species o Neoa hy eus Howden and Ma ínez om
Cos a Rica wi h dis ibu ion no es on o he A hy eini om Mexico and Cen al Ame -
ica (Coleop e a: Geo upinae). Canadian En omologis 116(12): 1637–1641. h ps://
doi.o g/10.4039/En 1161637-12
Howden HF, Gill BD (1987) New species and new eco ds o Panamanian and Cos a
Rican Sca abaeinae (Coleop e a: Sca abaeidae). Coleop e is s Bulle in 41: 201–224.
Howden H, Gill BD (1993) Mesoame ican On hophagus La eille in he dic anius and mi-
abilis species g oups (Coleop e a: Sca abaeidae). Canadian En omologis 125(6):
1091–1114. h ps://doi.o g/10.4039/En 1251091-6
Howden HF, Ma ínez A (1978) A e iew o he New Wo ld genus A hy eus. Con ibu ions
o he Ame ican En omological Ins i u e 15: 1–70.
Howden HF, Young OP (1981) Panamanian Sca abaeinae: Taxonomy, dis ibu ion, and
habi s (Coleop e a, Sca abaeidae). Con ibu ions o he Ame ican En omological In-
s i u e 18: 1–204.
Hue a C, C uz-Rosales M, González-Vaine P, Chamo o-Flo escano I, Ri e a JD, Fa ila
ME (2023) The ep oduc i e beha io o Neo opical dung bee les. F on ie s in Ecolo-
gy and E olu ion 11: 1102477. h ps://doi.o g/10.3389/ e o.2023.1102477
Huijb eg s J (1984) Bdely us geijskesi, a new sca ab (Coleop e a: Sca abaeidae) om
Su iname associa ed wi h B omeliaceae. Zoölogische Mededeelingen 59(6): 61–67.
Iannuzzi L, Libe al CN, De Souza TB, Pelleg ini TG, Da Cunha JCS, Ko oi a R, De Albuque -
que LSC, Cos a FC, Salomão RP, Maia ACD, Lei as FWT (2021) Sampling Me hods o
Bee les (Coleop e a). In: San os, JC, Fe nandes GW (Eds) Measu ing A h opod Bio-
di e si y. Sp inge , Cham, 125–185. h ps://doi.o g/10.1007/978-3-030-53226-0_6
ICF (2002) Plan de Manejo del Pa que Nacional Sie a de Agal a, 140 pp.
206
ZooKeys 1255: 127–206 (2025), DOI: 10.3897/zookeys.1255.150586
José D. Ri e a-Dua e e al.: Dung bee le o Hondu as
on Ha old E (1880) Ve zeichniss de on E. S einheil in Neu-G anada gesammel en
cop ophagen Lamellico nien. S e ine En omologische Zei ung. S e in 41: 13–46.
V eugdenhil ID, House PR, Ce a o CA, Ma ínez RA, Pe ei a AC (2002) Ra ionalisa ion
o he p o ec ed a eas sys em o Hondu as. Vol. 1: Main s udy. P epa ed by WICE.
Financed by PROBAP/Wo ld Bank/UNDP/GEF. Tegucigalpa, Hondu as.
Wea he and Clima e (2024) A e age mon hly ain all, empe a u e, and sunshine in Ca -
acamas, Hondu as. h ps://wea he -and-clima e.com/a e age-mon hly-Rain all-Tem-
pe a u e-Sunshine,Ca acamas,Hondu as
Wiedemann CKW (1823) Zweihunde neue Kä e on Ja a, Bengalen und dem Vo ge-
bi ge de gu en Ho nung. Zoologisches Magazin 2: 3–133.
Wille A (1973) Coleop e a: Sca abaeidae. Obse a ions on he beha io o a opical ain
o es dung bee le, Mega hoposoma candezei (Ha old). Re is a de Biología T opical
21: 41–57. h ps://a chi o. e is as.uc .ac.c /index.php/ b /a icle/ iew/27322
Zunino M (1984) Sis ema ica gene ica dei Geo upinae (Coleop e a, Sca abaeoidea:
Geo upidae), ilogenesi della so o amiglia e conside azione biogeog a iche. Bolle -
ino Museo Regionale di Scienze Na u ale To ino 2: 9–162.
Zunino M, Hal e G (1988) Análisis axonómico, ecológico y biogeog á ico de un g upo
ame icano de On hophagus. Museo Regionale di Scienze Na u ale To ino Monog a ia
9: 1–211.
