Curculionidae

Ocladius salicorniae, copyright K. V. Makarov.

Belongs within: Phytophaga.
Contains: Alcidides, Acicneminae, Brachycerini, Tanysphyrini, Platypodinae, Dryophthorinae, Bagoinae, Erirhinina, Sitonini, Hyperinae, Cyclominae, Curculioninae.

The Curculionidae are the largest family of weevils, distinguished by geniculate (elbowed) antennae with an elongate scape. Members of the family typically possess an elongate rostrum though this is relatively short in some subfamilies such as the Entiminae and absent in others such as Scolytinae. Most curculionids are phytophagous with the larvae feeding internally in plant tissues or subterraneously on roots (Lawrence & Britton 1991). A smaller proportion of species are saprophagous, feeding on dead plant matter, particularly wood. Some species have larvae feeding externally on foliage or reproductive structures (Anderson 2002). Many curculionids are economically significant, particularly as pests of crops, and have become widely distributed outside their native ranges. Some species have been deliberately introduced outside their places of origin to control plant pests.

Perhaps not surprisingly for such a large group, the division of the Curculionidae into subfamilies has varied significantly between authors. Recent molecular phylogenetic studies have further confused the issue by supporting few recognised subfamilies as monophyletic. Even such groups as the Scolytinae, distinctive enough to have historically been recognised as a distinct family, may not be resolved as a single clade (Mugu et al. 2018).

A basal grade of taxa, including the subfamilies ‘Erirhininae’ or ‘Brachycerinae’, Platypodinae and Dryophthorinae, retains features of the male genitalia in common with other non-curculionid weevils, such as separation of the pedon and tectum within the aedeagus. As such, they have been excluded from the Curculionidae sensu stricto by some authors. Molecular analyses have commonly supported a clade uniting Platypodinae and Dryophthorinae whereas Erirhininae/Brachycerinae are not monophyletic (Li et al. 2023).

The Myrtonymini and Raymondionymini are eyeless weevils with only four tarsomeres. Myrtonymini have the mesocoxal cavities connate, and both head and rostrum taper towards the antennal insertions. In Raymondionymini, the mesocoxal cavities are separated, and the head tapers to the rostrum base but the rostrum is subparallel (Legalov 2020). Within Raymondionymini, Alaocybites has contiguous front coxae and a deep suture between ventrites 4 and 5. Gilbertiola and Schizomicrus have the front coxae narrowly separated by the prosternum and the suture between ventrites 4 and 5 is more shallow. Gilbertiola has just five segments in the antennal funicle whereas Alaocybites and Schizomicrus have seven segments (Anderson 2002).

The rostrum is reduced in Cryptolaryngini and Perieges. The third tarsomere is bilobed in Cryptolaryngini, scales are not continuous over the body, the forehead is flat, and the ovipositor styles are weakly sclerotised. In Perieges the third tarsomere is slender, not bilobed, the scales form a continuous covering, the forehead is depressed, and the ovipositor styles are strongly sclerotised with a dentate sclerite at the apex (Legalov 2020).

A ventral channel is present on the prosternum in Arthrostenini, Himasthlophallus, TadiusAonychusOcladius, Paleocryptorhynchini and Desmidophorini. This channel extends onto the mesosternum in many of these taxa but not in Arthrostenini and Himasthlophallus. The procoxal cavities are separated in Arthrostenini. Himasthlophallus has reduced eyes and lacks a scutellum. Postocular lobes are well developed in Ocladius, Paleocryptorhynchini and Desmidophorini but weak in Tadius and Aonychus (Legalov 2020). Tadius is an Indo-Australian genus with externally lobed mandibles (Zimmerman 1993). Aonychus is an Australian genus with reduced tarsi, having only three segments and lacking terminal claws (Zimmerman 1993). Ocladius, a genus found in Africa and western Eurasia, has a spherical body form and no visible scutellum. Long teeth are present on the femora in Paleocryptorhynchini but only shorter ones in Desmidophorini, which also have a short metasternum (Legalov 2020).

The Curculionidae sensu stricto have the dorsum of the aedeagus fused with the sides and venter to form a cylinder or flattened cylinder (Zimmerman 1993). Molecular data support a division of most Curculionidae sensu stricto between two clades. One clade includes Curculioninae and allies, with most species having longer rostra and larvae that are internal feeders on above-ground plant tissues (Gillett et al. 2014). The other clade, including the Cyclominae, Entiminae and related subfamilies, includes mostly species with relatively short and broad rostra and larvae that are external feeders on roots. The Phrynixini, an Australo-Pacific tribe whose larvae (where known) develop in leaf litter, may represent the sister lineage to other members of this clade (Haran et al. 2023). The Styphlini, despite having endophytic larvae found in aerial parts of their hosts, may also belong to this clade (Haran et al. 2023).

The Coptonotinae are a small group of tropical weevils with a reduced rostrum, very similar in overall habitus to the Scolytinae and Platypodinae (Wood 1986). The Mecopelmini are small pale species, less than two millimetres in length, with relatively stout fore legs whereas the Coptonotinae are larger and darker with relatively slender fore legs. Galloisia is an eastern Asian genus that was assigned to its own subfamily by Morimoto (1962) on the basis of its lack of labial palpi, large maxillary palpiger, short fourth tarsal segment, and swollen rostrum in the female.

Characters (from Lawrence & Britton 1991): Usually moderately to strongly convex, robust, heavily sclerotised, often clothed with scales or bristles. Head more or less produced in front of eyes to form rostrum, which is usually much longer than broad; antennae geniculate with long scape and more or less compact club; labrum absent; maxillae reduced with short rigid palps; gular sutures fused; penultimate tarsal segment usually minute and concealed at base of lobed third segment. Larvae more or less C-shaped, very lightly sclerotised, with minute antennae and robust mandibles.

<==Curculionidae (see below for synonymy)L20
|--Raymondionymini [Raymondionyminae]LE23
| |--Raymondionymus perrisiB14
| |--BordoniolaB14
| |--Gilbertiola Osella 1982 [incl. Gilbertia Osella 1977 non Cossman 1889]A02
| |--Schizomicrus Casey 1905 [incl. Schizonotus Casey 1892 nec Ratzeburg 1852 nec Thorell 1888]A02
| | `--S. caecus (Casey 1892)A02
| `--Alaocybites Gilbert 1956A02
| |--A. californicus Gilbert 1956B14
| |--A. dubatolovi Legalov 2020L20
| |--A. egorovi Grebennikov 2010L20
| `--A. rothiB14
`--+--Ocladius [Ocladiinae, Ocladiini]LE23
| |--O. bifasciatus Tourn. 1875 [=Oecladius bifasciatus; incl. Oc. setipes Ancey 1881]G89
| |--O. holomelas Fairmaire 1877L20
| |--O. salicorniae [=Rhynchaenus salicorniae]LC40
| |--O. sapeti Gestro 1889G89
| `--O. sharpiG89
`--+--BrachyceriniLE23
`--+--+--TanysphyriniLE23
| `--+--PlatypodinaeLE23
| `--DryophthorinaeLE23
`--+--BagoinaeLE23
`--+--+--TheanellusHL23
| `--+--ErirhininaHL23
| `--Thryogenes festucaeHL23
`--+--+--Phrynixini [Phrynixinae]L20
| | | i. s.: SyagriusHL23
| | | |--S. fulvitarsis Pascoe 1875Mas86
| | | `--S. intrudens Waterhouse 1903Z93
| | |--Phrynixus Pascoe 1875HL23
| | | `--P. terreusHL23
| | `--GeochinaL20
| | |--Geochus Broun 1882L20
| | `--Afrogeochus Rheinheimer 1998L20
| `--+--SitoniniHL23
| `--+--HyperinaeHL23
| `--+--+--Hypsomus Schoenherr 1836HL23
| | | `--H. albosuturalis Marshall 1921HL23
| | `--StyphliniHL23
| | |--Styphlus penicillusSKR96
| | |--Orthochaetes Germar 1824HL23
| | | `--O. setigerHL23
| | `--Pseudostyphlus Tournier 1874HL23
| | `--P. pilumnusHL23
| `--CyclominaeHL23
`--CurculioninaeLE23

Curculionidae incertae sedis:
Camptoscapus planiusculus (Broun 1880)E02
Microtribus huttoniE02
Hydramphus tuberculatusWH02
Ectopsis ferrugalisWH02
Megacolabus garviensisWH02
Tychanus verrucosusWH02
TychanopaisWH02
|--T. fougeriWH02
`--T. tuberosusED04
Didymus impexusWH02
Echinopeplus insolitusWH02
EpitimetesWH02
|--E. cuprealisWH02
|--E. grisealisWH02
`--E. lutosusWH02
Sargon suturalisWH02
SympediusWH02
RachidiscodesED04
GermainiellusMC94
Sericotrogus subaenescensWFS04
Acanthothorax Gaede 1832BG06
Tigones caudataT27
Hoplocneme hookeriT27
Empaeotes censoriusT27
Xanthochelus postumus Faust 1883G89
CorigetusG89
Odoiporus longicollisMHG04
OodemasO94
RhyncogonusO94
RhynchodesG84
Homalinotus Schoenherr 1826WL09, C92 [=Homalonotus Agassiz 1846 nec Koenig 1825 nec Agassiz 1846C92]
|--H. coriaceusWL09
|--H. jamaicensis [=Rhynchaenus jamaicensis, Curculio jamaicensis, Dionychus (Homalonotus) jamaicensis]LC40
`--H. lherminieri Chv. 1878FS90
AlophusC01
|--A. quadriguttatusC01
|--A. rudis [incl. A. rudis var. gibbulosus]C01
`--A. triguttatus [=Curculio triguttatus]LC40
Cnemargus Schönherr 1837CW92a
Merodontus Jekel 1854CW92b
Neocnemis Crotch 1867CW92c
AlcididesMas86
EremnidesMas86
|--Platytrachelus chloris Pascoe 1870Mas86
|--Oops pistor Germ. 1848Mas86
|--PephricusMas86
| |--P. echymis Pascoe 1870Mas86
| `--P. rattulus Pascoe 1882Mas86
`--Eremnus setulosusLC40
Hipporrhinus [Hipporhinides, Hipporhininae]Z93
|--H. bulbiferB35
|--H. duponti Boisduval 1835B35
|--H. heeriB14
`--H. sexvittatusLC40
RhyparosomidesMas86
|--Zephryne sordida Pascoe 1870Mas86
`--DysostinesMas86
|--D. cellaris Pascoe 1873Mas86
|--D. fuligineus Pascoe 1870Mas86
|--D. hoplostethus Pascoe 1870Mas86
|--D. pilipes Pascoe 1870Mas86
|--D. pustulosus Pascoe 1870Mas86
`--D. valgus Pascoe 1870Mas86
CylindrorhinidesMas86
|--Pantopoeus cervinusMas86
|--Peripagis rufipes Pascoe 1870Mas86
|--Lycosura bispinosa Pascoe 1875Mas86
|--Decienus sphasodes Pascoe 1873Mas86
`--OcynomaMas86
|--O. antennata Pascoe 1873Mas86
`--O. cordipennis Pascoe 1873Mas86
TanyrhynchidesMas86
|--Synatonyx ovatus Wollast. 1853Mas86
|--Xynaea saginata Pascoe 1865Mas86
`--Tanyrhynchus porifer [=Curculio porifer]LC40
BronchusZ93
Dinomorphus [Dinomorphinae]Z93
BrotheinaeZ93
ByrsopinaeZ93
LagostomusB35
|--L. australisB35
`--L. turritusLC40
ElytrurusB35
|--E. cinctus Dupont in Boisduval 1835B35
`--E. lapeyrousei Boisduval 1835B35
Brachyrhinus Latreille 1802L02 [BrachyrhininaeH02]
|--B. lineatus (Linnaeus 1758)L02, L58 [=Curculio lineatusL02]
`--B. niger [=Curculio niger]L02
CylindrocopterusG05
Penestes tigrisZ93, LC40 [=Rhynchaenus tigrisLC40]
Anathymus singularis Pascoe 1885Mac86
AmpharthropelmaP92
AnchorthorrhinusP92
ArchimetrioxenaP92
ElectrotribusP92
ErirhinoidesP92
IsalcidodesP92
NecrodryophthorusP92
PaleopissodesP92
SuccinacallesP92
ThryogenosomaP92
Pantomerus galapagoensisPP72
Phyllolytus Fairmaire 1889Fa89
`--*P. longicornis Fairmaire 1889Fa89
Trymatoderus Fairmaire 1889Fa89
`--*T. spongiicollis Fairmaire 1889Fa89
Oxyderces Schönh. 1833FS90
|--O. cretaceus Fab. 1792FS90 [=Curculio cretaceusLC40, Cyphus cretaceusLC40; incl. O. scutellaris Chv. 1880FS90]
`--O. lacteus Fab. 1781 [incl. O. religiosus]FS90
Neocyphus Bed. 1883FS90
|--N. gentilis Ol. 1807FS90
|--N. pudens Boh. 1833 [incl. N. gilvipes, N. pudens var. leucocephalus Chv. 1880]FS90
`--N. strangulatus Chv. 1880 [incl. N. sanguinipes]FS90
Prepodes Schönh. 1826FS90
|--P. marmoreus Gyll. 1840 [incl. P. marmoreus var. marmoratus Chv. in Fleutiaux & Sallé 1890]FS90
|--P. novemdecimpunctatus [=Curculio novemdecimpunctatus, Diaprepes novemdecimpunctatus]LC40
`--P. quadritaenia Chv. 1880FS90
Ophthalmoborus testaceusFS90
Cylindrocorynus Schönh. 1837FS90
`--C. thoracicus Chv. 1880FS90
Pseudoscythropus apollinisDL90
MetallitesDL90
|--M. ambiguusLC40
`--M. globosus [incl. M. javeti, Cneorhinus pubescens]DL90
Holcorrhinus metallicusDL90
Strabus Gerstaecker 1860V02
Plaesiorhinus notatusA89
Epizeuxis lyterioidesFl89
Galloisia Hustache 1920 [Galloisiinae]M62
`--G. inflataM62
Endaenidius Morimoto 1962M62
`--*E. ikezakii Morimoto 1962M62
Gryphorrhynchus Roelofs 1875M62
Imerodes Marshall 1926M62
Heterendaeus Morimoto 1962M62
`--*H. miyamotoi Morimoto 1962M62
Nesendaeus Marshall 1931M62
Euphyllobiomorphus Morimoto 1962M62
`--*E. kurosawai Morimoto 1962M62
Brachyonyx Schönherr 1836M62
AcicneminaeM62
Pachyops incertus Gahan 1900JG19
Rhyncolosoma dubium (Gahan 1900) [=Phloeophagosoma dubium]JG19
RhyncholobusJG19
|--R. andrewsi Gahan 1900JG19
|--R. discoidalis (Waterhouse 1887) [=Piezonotus discoidalis]JG19
|--R. rossi Gahan 1900JG19
`--R. vittatus Gahan 1900JG19
Arariperhinus monneiLB20
CoptonotinaeMPJ18
|--Mecopelmus Blackman 1944 [Mecopelmini]MPJ18
| `--M. zeteki Blackman 1944MPJ18
`--CoptonotiniGC-P14
|--Protohylastes annosusW86
|--Protoplatypus vetulusW86
|--Schedlarius mexicanusW86
`--Coptonotus Chapuis 1873MPJ18
`--C. cyclopusMPJ18
*Sophrorhinus duvernoyiM78
AmphialodesM78
Ypsilepidus thisoidesM78
Protacallinus Morimoto 1962M78, M62
`--*P. uenoi Morimoto 1962M62
Protacalles Voss 1957M78, M62
EpriasM78
MicrapriesM78
Catabonops monachusM78
SystaltopezusM78
BrachycolobodesM78
ColobodellusM78
NannocolobodesM78
DinapriesM78
AmphialusM78
EucolobodesM78
Exapries lophonotusM78
Cyphomidica megacallesM78
Dystropicus squalidusM78
Acallinus Morimoto 1962M78, M62
`--*A. tuberculatus Morimoto 1962M62
Colobodes billbergiM78
Thisus biguttatusM78
Perrhaebius ephippigerM78
Phrygena ephippiataM78
SolobrachisM78
OcolobodesM78
Apries eremitaM78
ParapriesM78
DeretiosomimusM78
Deiradocranus Marshall 1953 [incl. Deiradocranoides Morimoto 1962]M78
|--D. latebrisM78
`--*Deiradocranoides’ setosus Morimoto 1962M62
MecocorynusM78
|--M. humerosus Fairmaire 1889Fa89
|--M. varipesFa89
`--M. westermanniM78
Clisis modesta Pascoe 1875Mas86
Diethusa fervida Pascoe 1873Mas86
Eniopea amoena Pascoe 1873Mas86
Nedyleda semiusta Pascoe 1872Mas86
Oenochroma rubeta Pascoe 1872Mas86
Hedyopsis selligera Pascoe 1873Mas86
Methone ornata Pascoe 1875Mas86
Paryzeta musiva Pascoe 1873Mas86
Olanaea nigricollis Pascoe 1873Mas86
Cyttalia griseipila Pascoe 1873Mas86
Glaucopela unicolor Pascoe 1874Mas86
Empira variegata Pascoe 1874Mas86
Trechia Pascoe 1873Mas86
NemestraMas86
|--N. incerta Pascoe 1872Mas86
`--N. vibrata Pascoe 1882Mas86
XedaMas86
|--X. amplipennis Pascoe 1873Mas86
`--X. bilineata Pascoe 1873Mas86
OrphaMas86
|--O. flavicornis Pascoe 1870Mas86
`--O. persimilis Pascoe 1872Mas86
PhaeodicaMas86
|--P. fulvicornis Pascoe 1874Mas86
`--P. scutellaris Pascoe 1874Mas86
PhrenozemiaMas86
|--P. lunata Pascoe 1873Mas86
`--P. lyproides Pascoe 1872Mas86
EnideMas86
|--E. aestuans Pascoe 1873Mas86
|--E. porphyrea Pascoe 1873Mas86
`--E. saniosa Pascoe 1873Mas86
BrexiusMas86
|--B. angusticollis Pasoce 1870Mas86
|--B. lineatus Pascoe 1873Mas86
`--B. murinus Pascoe 1870Mas86
AntyllisMas86
|--A. aurulenta Pascoe 1873Mas86
|--A. griseola Pascoe 1873Mas86
|--A. setosa Pascoe 1873Mas86
`--A. togata Pascoe 1875Mas86
RhachiodesMas86
|--R. bicaudatus (Boisduval 1835) [=Myllorhinus bicaudatus; incl. R. spinicollis]Mas86
|--R. conformis Chev. 1879Mas86
|--R. dentifer [=Pteroporus dentifer]Mas86
|--R. granulifer Chev. 1879Mas86
|--R. multidentatus Chev. 1879Mas86
|--R. nigropunctatus Chev. 1879Mas86
|--R. posticus Chev. 1879Mas86
`--R. signaticollis Chev. 1879Mas86
AopolocnemisMas86
|--A. alboguttatus Chev. 1879Mas86
|--A. dorsonotatus Chev. 1879Mas86
|--A. fulvus Chev. 1879Mas86
|--A. guttiger Pascoe 1882Mas86
|--A. lineata Pascoe 1872Mas86
|--A. phaleratus Erichs. 1842Mas86
|--A. rufipesMas86
|--A. semicinctus Chev. 1879Mas86
`--A. suturalis Pascoe 1882Mas86
MyrtonyminiL20
CryptolarynginiL20
ArthrostenusLC40 [ArthrosteniniL20]
`--A. fulloLC40
PaleocryptorhynchiniL20
DesmidophoriniL20
Perieges [Periegini]L20
`--P. bardus Boheman 1842L20
Himasthlophallus Zherikhin & Egorov 1990MPJ18 [HimasthlophalliniL20]
`--H. flagellifer Egorov & Zherikhin 1991L20
Tadius Pascoe 1885Z93 [Tadiinae, TadiiniL20]
|--*T. erirhinoides Pascoe 1885 (see below for synonymy)Z93
|--T. barbatus Lea 1928 [=T. barbatulus (l. c.)]Z93
`--T. laticollis Faust 1898Z93
Aonychus Schoenherr 1844Z93 [AonychusiniL20]
|--*A. hopei Boheman 1844Z93
|--A. barbatus Lea 1927Z93
|--A. bicruciatus Lea 1905 [=A. hopei var. bicruciatus]Z93
|--A. lineatus Pascoe 1870Z93
|--A. lituratus Lea 1927Z93
|--A. luctuosus Pascoe 1872 [incl. A. picatus Lea 1927]Z93
|--A. pachypus Lea 1905Z93
`--A. striatus Lea 1905Z93
Ita [Itini]HL23
`--I. chavanoniHL23
Rhigus schuppeliLC40
Polydius steveniLC40
Epirrhynchus argusLC40
Nebalis squalidaLC40
Psalidium maxillosumLC40
Syzygops cyclopsLC40
Herpisticus laevicollis [incl. Curculio erimita]LC40
Hadropus albicerisLC40
CyphusLC40
| i. s.: C. schoenherriL90
|--C. (Cyphus)LC40
| |--C. (C.) gibber [=Curculio gibber]LC40
| `--C. (C.) juvencus [=Curculio juvenus]LC40
`--C. (Clatyamus) nodipennisLC40
Eustales auronitensLC40
Polycomus lanuginosusLC40
Eurilia lateralis [=Apotomus lateralis]LC40
Anaemerus fuscus [=Curculio fuscus]LC40
Siderodactylus sagittarius [=Curculio sagittarius]LC40
Pandeleticus pauperculusLC40
Eugnathus viridanusLC40
Entyus albicinctusLC40
Eudius quadrisignatusLC40
Tanaos sanguineusLC40
Pachycerus albariusLC40
Gronops lunatus [=Curculio lunatus]LC40
HypsonotusLC40
|--H. (Hypsonotus) clavulusLC40
`--H. (Eurylobus) cingulatusLC40
Lophotus eschscholtziLC40
Minyops carinatus [=Curculio carinatus; incl. C. senes, C. variolosus]LC40
Cepurus torridus [=Curculio torridus]LC40
Byrsops bucellariusLC40
Eupages tuberculosusLC40
Rhytirrhinus inequalis [=Curculio inequalis]LC40
Brachipus lixoidesLC40
Eudocinus mannerheimiiLC40
Aorus spadiceusLC40
Cnemidophorus fasciculatusLC40
Phytophilus cruciferusLC40
Anthobius testaceus [=Rhynchaenus testaceus]LC40
Adelus cupreusLC40
Brachonyx indigena [=Curculio indigena, Rhynchaenus indigenus]LC40
Botanebius tuberculatusLC40
Rachiodes spinicollisLC40
LoncophorusLC40
|--L. obliquusLC40
`--L. parasitaLC40
Megarhinus firmusLC40
Phiternus farinosusLC40
Anchylorhynchus variabilisLC40
Tylomus gonopterus [=Orobitis gonopterus]LC40
Euderes lineicollis [=Rhynchaenus lineicollis]LC40
Solenopus cacicusLC40
Amerhinus dufresnii [=Rhynchaenus dufresnii]LC40
Pinarus spiculum [=Paecilma spiculum]LC40
Athrotomus depressusLC40

Curculionidae [Brachyceridae, Brachycerinae, Calandraeides, Calandridae, Calandrides, Calandrina, Calendridae, Calendrinae, Cryptopygi, Dryophthoridae, Erirhinidae, Erirhinides, Erirhininae, Erirhinini, Gonatoceri, Ithyporides, Ithyporini, Phanerognatha, Rhynchophoridae, Rhynchophorides, Sophrorhinini, Sophrorhinides, Sophorrhini]L20

*Tadius erirhinoides Pascoe 1885 [incl. T. australiae Lea 1928, T. erirhinoides lineatus Faust 1894, T. erirhinoides suspiciosus Faust 1894]Z93

*Type species of generic name indicated

References

[A02] Anderson, R. S. 2002. Curculionidae Latreille 1802. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 722–815. CRC Press.

[A89] Auld, T. D. 1989. Larval survival in the soil and adult emergence in Melanterius Erichson and Plaesiorhinus Blackburn (Coleoptera: Curculionidae) following seed feeding on Acacia and Bossiaea (Fabaceae). Journal of the Australian Entomological Society 28: 235–238.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d’Urville. Faune entomologique de l’océan Pacifique, avec l’illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature’s gems. Ivy Press: Lewes (United Kingdom).

[BG06] Bretfeld, G., & A. Greigel. 2006. Acanthoneelidus nom. n. for the genus Acanthothorax Bredfeld & Griegel 1999 from northwestern Poland (Insecta, Collembola, Neelidae). Senckenbergiana Biologica 86 (1): 46.

[CW92a] Cassis, G., & T. A. Weir. 1992a. Aphodiinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 81–105. AGPS Press: Canberra.

[CW92b] Cassis, G., & T. A. Weir. 1992b. Scarabaeinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 106–173. AGPS Press: Canberra.

[CW92c] Cassis, G., & T. A. Weir. 1992c. Dynastinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 383–425. AGPS Press: Canberra.

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