Hoplia coerulea, copyright Frank Vassen.

Belongs within: Scarabaeidae.
Contains: Rhizotrogus, Automolius, Aplopsis, Maechidius, Systellopini, Liparetrus, Colpochila, Euchirini, Phyllotocus, Xylonichini, Pachydemini, Melolonthini, Rutelinae, Cetoniinae, Diphucephala, Sericini, Heteronyx, Neoheteronyx, Gnaphalopoda, Sericesthis, Scitala.

The Melolonthinae, chafers, are a cosmopolitan group of scarabaeid beetles with soil-dwelling larvae that feed on roots and hummus. Though larvae may live for up to two years, the adult life period is relatively brief and lasts only a few days or weeks. Adults of some species feed on the leaves of trees; others do not feed as adults at all (Houston & Weir 1992). As currently recognised, the melolonthines are probably not monophyletic with other phytophagous scarabaeid subfamilies such as Dynastinae, Rutelinae and Cetoniinae nested within the group.

Members of the Melolonthinae are divided between several tribes though many have only been characterised in regard to particular geographic regions. In many of these tribes, the posterior tibia has two apical spurs that are placed above and below the tibia so that the tarsus passes between the spurs when moved from side to side. However, both spurs are placed ventrally in Plectris, Pachytricha, Diphucephalini and Melolonthini. Members of the Australian genus Phyllotocidium are at least partially metallic in coloration and have a three-segmented antennal club. The Phyllotocini have deeply projecting fore coxae, large hind coxae, a slender process with a setose ending usually present on the prosternum, and strongly asymmetrical claws in the male. The Automoliini and Maechidiini have the tergite and ventrite of the penultimate abdominal segment at least partially fused, so the lateral suture is either absent or incomplete (Britton 1957).

Within the Holarctic realm, the Hopliini are distinguished by having a single claw on each tarsus and the mid and hind tibiae lack apical spurs. The Chasmatopterini have the sutures between the abdominal ventrites obliterated. The Nearctic Oncerini have all abdominal spiracles placed within the pleural membrane, both hind tibial spurs placed below the tarsus, and the labrum located on the apical margin of the clypeus (Ratcliffe et al. 2002).

The Melolonthinae: chafers and June bugs
Published 22 April 2020

Within the bewildering array that is beetle diversity, one of the more readily recognisable groups is the Scarabaeoidea, the assemblage that includes dung beetles (which, as it happens, are what I currently spend most of my days looking at) and related forms. Members of this group are easily distinguished from other beetles by their distinctive antennae, ending in an asymmetrical club with segments extending to one side like a set of fingers. Several families, many of them further subdivided into subfamilies, are currently recognised within the scarabaeoids. One of the most commonly encountered scarabaeoid subgroups is the subfamily Melolonthinae, commonly known as the chafers.

Green scarab beetles Diphucephala sp., a common genus of day-flying melolonthines here in Australia, copyright Boobook48.

Somewhere in the region of eleven thousand species around the world have been assigned to this grouping; as always, doubtless many more could be recognised by those who take the time. Melolonthinae is generally recognised as a subfamily of the family Scarabaeidae, sharing with other scarabaeids features such an antennal club in which the segments are relatively narrow and can be smoothly pressed against each other, and an exposed pygidium (the last dorsal plate on the abdomen, forming what you might think of as the ‘butt plate’). Some authors have recognised melolonthines as a distinct family but this is the less commonly utilised option. Melolonthines belong to a group of mostly plant-feeding subfamilies in which the row of abdominal spiracles bends downwards towards the rear so at least the last pair remains visible when the elytra are closed. Within this cluster, melolonthines tend to be characterised more by lacking the features of the other subfamilies than by distinctive features of their own (more on that in a moment) but general features include mandibles that are not visible when looking down on the top of the head, fore coxae that do not protrude much ventrally, equal claws on each leg (at least on the mid and hind legs) and only one visible spiracle when the elytra are closed. The labrum (the piece at the front of the mouthparts that might be thought of as the insect’s top lip) is usually hardened and may be more or less fused with the clypeus (the lower- or foremost section [depending how you look at it] of the front of the head capsule). Many melolonthines are noticeably hairy and/or dull in comparison with other scarabaeoids but others may be shiny and/or metallic in coloration.

Sugarcane white grub beetle Lepidiota stigma, copyright Bernard Dupont.

For the most part, melolonthines are plant-feeders at both larval and adult stages of the life cycle (Lawrence & Britton 1991). The greater part of the active life cycle is taken up by the larval stage which may last for many months (Britton 1957). Larvae mostly live underground, feeding on plant roots and humus. A number of species have made themselves known as significant pests in this manner because of the damage they may inflict on pastures or agricultural crops (the grass grub Costelytra zealandica comes immediately to mind as a good example of this in my native New Zealand). Pupation also occurs underground in subterranean cells and mature adults may remain dormant in these cells for some months waiting for conditions to be just right for emergence. Once they do emerge from the ground, however, the adult life span is quite brief, only lasting a few weeks or even days. Because of this brief emergence, and because their habit of waiting for specific environmental cues means that large numbers may appear seemingly all at once, many species have been awarded vernacular names that reflect their seasonality such as June bug (in the Northern Hemisphere) or Christmas beetle (in the Southern). Some species will feed on foliage as adults, some may visit flowers for pollen and nectar, other particularly short-lived species will not feed as adults at all. The majority of adult melolonthines are active at dusk or night, spending the days sheltered in secluded locations, but a number of flower-feeding species are active by day (Britton 1957).

The infamous grass grub Costelytra zealandica, illustrated by Desmond Helmore.

The classification of melolonthines can charitably be described as an absolute mess. As noted above, we can confidently say that they belong to a clade with other subfamilies of plant-feeding scarabaeids (the Cetoniinae, Rutelinae and Dynastinae) but the features setting them apart from these other subfamilies are likely to be primitive for the group. As such, it comes as little surprise that phylogenetic studies have failed to establish the Melolonthinae as monophyletic (e.g. Eberle et al. 2018; Woolley 2016). However, it seems that no-one thinks that an adequately expansive study that would allow them to be appropriately divvied up has yet been done. Matters are not helped by the absence of a well-established internal classification for melolonthines. Various distinct subgroups can be recognised and between twenty or thirty tribes have been recognised around the world. But the relationships between these tribes remain uncertain, as does the tribal position of many genera. Much of the revisionary work that has been done has been conducted at a regional level only. Thus, for instance, the tribal classification of Australian melolonthines established by Britton (1957) applies only to Australian species and the tribal distinctions Britton recognised may end up falling apart if one attempted to apply them to species from elsewhere. Not that the authors should be criticised for this situation: after all, when one is dealing with over 11,000 species, things rapidly tend to become unmanageable.

Systematics of Melolonthinae

Characters (from Houston & Weir 1992): Head and pronotum of males unarmed; mandibles completely concealed from above; abdominal spiracles diverging, with one pair exposed beneath edge of elytra; fore coxae transverse, mesothoracic epimera not visible from above; mid coxae transverse or only slightly oblique; tarsal claws equal. Larvae with apical antennal segment about as wide as penultimate segment; mandibles withouyt transverse granular ridges forming stridulatory areas; galea and lacinia either partly fused proximally or fitting tightly together; anal cleft usually Y-shaped or angulate.

Melolonthinae [Chasmatopterinae, Melolonthidae, Melolonthides, Sericinae]
|--Phyllotocini [Phylloticides, Phyllotocina]A06
| |--PhyllotocusA06
| `--Anthotocus Britton 1957A06, HW92
| |--*A. luridus (Macleay 1864)HW92 (see below for synonymy)
| |--A. antennalis (Lea 1919) [=Phyllotocus antennalis]HW92
| |--A. cribriceps (Lea 1919) [=Phyllotocus cribriceps]HW92
| |--A. fugitivus (Lea 1920) [=Phyllotocus fugitivus]HW92
| `--A. perissus Britton 1957HW92
|--Hoplia Illiger 1803RJS02 [HopliiniA06]
| |--H. aureola [incl. H. aureola var. sibirica]C01
| |--H. aurotinctaF89
| |--H. campestrisF89
| |--H. coerulea (Drury 1773)B14
| |--H. duodecimpunctataF89
| |--H. equina LeConte 1880A06
| |--H. farinosa [=Melolontha farinosa]G20
| |--H. formosaR26
| |--H. graminicola (Fabricius 1792)A06
| |--H. minuscula Fairmaire 1889F89
| |--H. nebulosa Fairmaire 1889F89
| |--H. ochreata Fairmaire 18989F89
| |--H. parvulaC01
| |--H. philanthusR26
| |--H. rufopicta Fairmaire 1889F89
| |--H. sulphurea Chevrolat 1846E12
| `--H. validipes Fairmaire 1889F89
| `--+--PachydeminiA06
| |--ChasmatopteriniA06
| | |--Chnaunanthus Burmeister 1844 [incl. Acratus Horn 1867 non Lac. 1866, Pseudacratus Dalle Torre 1912]RJS02
| | |--Chaunocolus Saylor 1937RJS02
| | | `--C. cornutusRJS02
| | `--ChasmatopterusA06
| | |--C. annamariae Branco 2004F05
| | `--C. hirtus (Illiger 1803)A06
| `--+--MelolonthiniA06
| `--+--+--RutelinaeMF15
| | `--CetoniinaeA06
| `--Macrodactylini [Dichelonycini]HW92
| |--Macrodactylus Dejean 1821 [incl. Stenothorax Harris 1827]RJS02
| |--Gymnopyge Linell 1895RJS02
| |--Coenonycha Horn 1876RJS02
| |--Ceraspis LePeletier & Serville 1828 [incl. Faula Blanchard 1850]RJS02
| |--Dichelonyx Harris 1827 [incl. Dichelonycha Kirby 1837]RJS02
| |--Isonychus Mannerheim 1829MF15, RJS02
| `--Plectris Saint-Fargeau & Serville 1828HW92
| |--*P. tomentosa Saint-Fargeau & Serville 1828HW92
| `--P. aliena Chapin 1934HW92
`--+--Diphucephalini [Diphucephalides]HW92
| |--DiphucephalaA06
| `--Cunderdinia Lea 1916HW92
| |--*C. variabilis Lea 1916HW92
| `--C. setistriata Lea 1930HW92
`--+--+--Sericoides [Sericoidini]A06
| `--+--+--Athlia rustica Erichson 1835A06
| | `--+--SericiniA06
| | `--Ablaberini [Camentini]A06
| | |--Ablabera hirsuta Blanchard 1850A06
| | |--CyrtocamentaA06
| | | |--C. pygidialis Frey 1968A06
| | | `--C. pygmaea Brenske 1897A06
| | `--+--Empecamenta buettikeri Ahrens 2000A06
| | `--CamentaA06
| | |--C. elongata Frey 1960A06
| | `--C. westermanni Harold 1878A06
| `--HeteronyciniB00
| |--HeteronyxA06
| |--NeoheteronyxHW92
| |--Callabonica Blackburn 1895HW92
| | `--*C. propria Blackburn 1895HW92
| |--Cubidens Britton in Houston & Weir 1992 [=Paraheteronyx Britton 1988 non Moser 1924]HW92
| | `--*C. unguiculatus (Burmeister 1855) [=Heteronyx unguiculata, *Paraheteronyx unguiculata]HW92
| |--Eurychelus Blanchard 1850HW92
| | `--*E. marmoratus Blanchard 1850HW92
| |--Nepytis Erichson 1842HW92
| | `--*N. russula Erichson 1842HW92
| |--Proborhinus Britton 1988HW92
| | `--*P. cornutus (Blackburn 1910) [=Heteronyx cornutus]HW92
| |--Odontotonyx Macleay 1871HW92
| | |--*O. brunneipennis Macleay 1871HW92
| | `--O. ruficeps Lea 1919HW92
| |--Webbella Britton 1988HW92
| | |--*W. firma (Blackburn 1909) [=Heteronyx firmus]HW92
| | `--W. labralis (Blackburn 1908) (see below for synonymy)B00
| |--Acheilo Britton 1988HW92
| | |--*A. capitalis (Blackburn 1910) [=Heteronyx capitalis]HW92
| | |--A. clypeatus Britton 1988HW92
| | `--A. pedarius (Blackburn 1910) [=Heteronyx pedarius]B00
| `--Pseudoheteronyx Blackburn 1892HW92
| |--*P. baldiensis (Blackburn 1892) [=Heteronyx baldiensis]HW92
| |--P. basicollis Lea 1919HW92
| |--P. creber Blackburn 1908 [incl. P. seticollis Lea 1919]HW92
| |--P. helaeoides Blackburn 1892HW92
| |--P. laticollis Blackburn 1908HW92
| `--P. puncticollis Lea 1919HW92
|--Colpochelyne Britton 1987HW92
| `--*C. dux Britton 1987HW92
|--Idanastes Britton 1987HW92
| `--*I. abditus Britton 1987HW92
|--Phorine Britton 1987HW92
| `--*P. anomala (Blackburn 1907) [=Byrrhomorpha anomala]HW92
|--Synchilus Britton 1955HW92
| `--*S. gisleni Britton 1955HW92
|--Xyrodes Britton 1987HW92
| `--*X. calorata (Blackburn 1907) [=Frenchella calorata]HW92
|--Protelura Britton 1987HW92
| |--*P. nana Britton 1987HW92
| `--P. guttata Britton 1987HW92
|--Hadrops Britton 1987HW92
| |--*H. halei Britton 1987HW92
| `--H. flavus (Lea 1919) [=Engyops flavus]HW92
|--Colpochilodes Blackburn 1898HW92
| |--*C. raucipennis Blackburn 1898HW92
| `--C. peregrina Britton 1987HW92
|--Xyrine Britton 1987HW92
| |--*X. carnei Britton 1987HW92
| `--X. inusitatus Britton 1987HW92
|--Byrrhomorpha Blackburn 1892HW92
| |--*B. verres Blackburn 1892HW92
| |--B. basicollis Lea 1919HW92
| `--B. ponderosa Blackburn 1892 [incl. B. rudis Lea 1919]HW92
|--Ophropyx Britton 1987HW92
| |--*O. hispida (Blackburn 1898) [=Frenchella hispida]HW92
| |--O. approximans (Blackburn 1898) [=Frenchella approximans]HW92
| `--O. ciliata (Boisduval 1835)HW92 (see below for synonymy)
|--Xyroa Britton 1987HW92
| |--*X. polita Britton 1987HW92
| |--X. alta Britton 1987HW92
| |--X. amoena Britton 1987HW92
| `--X. nitida Britton 1987HW92
|--Homolotropus Macleay 1871 [=Homalotropus (l. c.)]HW92
| |--*H. luridipennis Macleay 1871HW92
| |--H. metallicus Britton 1970HW92
| |--H. sagus Britton 1970HW92
| |--H. sericeus Britton 1970HW92
| `--H. taylori Britton 1970HW92
`--Telura Erichson 1842A06, HW92
|--*T. vitticollis Erichson 1842HW92
|--T. alta Britton 1987HW92
|--T. imparilis Britton 1987HW92
|--T. monticola Britton 1987HW92
`--T. petiolata Britton 1987HW92

Melolonthinae incertae sedis:
|--P. festivaT27
|--P. laetaB35
`--P. refulgens Dupont in Boisduval 1835B35
Ancistrosoma klugii Curtis 1835B14
Chlorochiton suturalis [=Stethaspis suturalis]L19
Comophorina Strand 1928 (see below for synonymy)HW92
`--*C. testaceipennis (Blanchard 1850) [=*Comophorus testaceipennis, *Comophus testaceipennis]HW92
Phyllotocidium Blackburn 1898 [Phyllotocidiini]HW92
|--*P. macleayi (Blackburn 1892) [=Cheiragra macleayi]HW92
|--P. bimaculiflavum Lea 1917HW92
|--P. pictum Lea 1916HW92
`--P. viridis Britton 1957HW92
Pachytricha Hope 1841 [=Pachytrichia (l. c.); Pachytrichini]HW92
|--*P. castanea Hope 1841 [incl. P. munda Sharp 1874, P. pallens Sharp 1874]HW92
|--P. demarzi Frey 1966HW92
|--P. minor Sharp 1874HW92
|--P. robusta Sharp 1874HW92
`--P. tecta Sharp 1874HW92
Automoliini [Automolini]HW92
|--Bryantella Britton 1957HW92
| `--*B. castanea Britton 1957HW92
|--Maechidinus Lea 1919HW92
| |--*M. latericollis Lea 1919HW92
| |--M. marginalis Lea 1919HW92
| `--M. sculptilis Britton 1957HW92
`--Phyllochlaenia Blanchard 1846 (see below for synonymy)HW92
|--*P. rufescens Blanchard 1846HW92 (see below for synonymy)
|--‘Caulobius’ modestus Blanch. 1850M86
`--P. villosus (Guillou 1844) (see below for synonymy)HW92
Maechidiini [Maechidiides]HW92
|--Microcoenus Britton 1957HW92
| `--*M. nanus Britton 1957HW92
|--Harpechys Britton 1957HW92
| `--*H. chilo Britton 1957HW92
|--Microthopus Burmeister 1855HW92
| `--*M. castanopterus Burmeister 1855 [incl. Macleayia hybrida Blackburn 1888]HW92
|--Termitophilus Britton 1957HW92
| `--*T. spadix Britton 1957HW92
`--Epholcis Waterhouse 1875HW92
|--*E. divergens Waterhouse 1875 [incl. Maechidius albertisi Fairmaire 1877]HW92
|--E. bilobiceps (Fairmaire 1877) [=Maechidius bilobiceps]HW92
|--E. gracilis (Waterhouse 1875) [=Maechidius gracilis]HW92
|--E. longior Blackburn 1898HW92
`--E. uniformis Britton 1957HW92
Liparetrini [Colpochilini]HW92
| i. s.: ‘Melolontha’ astrolabei Boisduval 1835 [=Haplonycha astrolabei]HW92
|--Dysphanochila Blackburn 1898HW92
| `--*D. pilosipennis Blackburn 1898HW92
|--Parasciton Britton 1990HW92
| `--*P. inermis Britton 1990HW92
|--Allara Britton 1955HW92
| `--*A. insularis Britton 1955HW92
|--Anacanthodes Britton 1990 [=Anacanthopus Blackburn 1898 non Montandon 1894]HW92
| `--*A. inermis (Blackburn 1898) [=*Anacanthopus inermis]HW92
|--Macleayella Britton 1990 [=Macleayia Blackburn 1888 non Haswell 1880]HW92
| `--*M. singularis (Blackburn 1888) [=*Macleayia singularis]HW92
|--Aneucomides Blackburn 1898HW92
| `--*A. coloratus Blackburn 1898 [incl. Haplonycha mauricei Blackburn 1906]HW92
|--Aphanesia Britton 1990HW92
| `--*A. greyi Britton 1990HW92
|--Astibicola Britton 1990HW92
| `--*A. bicolor Britton 1990HW92
|--Cheilo Britton 1990HW92
| `--*C. liparetroides Britton 1990HW92
|--Dikellites Britton 1990HW92
| `--*D. abditus Britton 1990HW92
|--Engyopsina Britton 1990 [=Engyops Blackburn 1898 non Bauer & Bergenstamm 1889]HW92
| `--*E. spectans (Blackburn 1898) [=Engyops spectans]HW92
|--Hadropechys Britton 1990HW92
| `--*H. victoriae (Blackburn 1897) [=Pachygastra victoriae]HW92
|--Ictigaster Britton 1986HW92
| `--*I. ruficollis (Lea 1917) [=Haplonycha ruficollis]HW92
|--Leonotus Britton 1986HW92
| `--*L. pilosicollis (Lea 1930) [=Haplonycha pilosicollis]HW92
|--Pachygastra Germar 1848HW92
| `--*P. tasmanica Germar 1848 [incl. Prochelyna rubella Schaufuss 1882]HW92
|--Paronyx Britton 1990HW92
| `--*P. setifera Britton 1990HW92
|--Petinopus Blackburn 1898HW92
| `--*P. aegrotus Blackburn 1898HW92
|--Scitaloides Britton 1990HW92
| `--*S. malanda Britton 1990HW92
|--Stenochelyne Britton 1990HW92
| `--*S. noctis Britton 1990HW92
|--Xyridea Britton 1990HW92
| `--*X. hirticollis (Blackburn 1912) [=Aneucomides hirticollis]HW92
|--Nosphisthis Blackburn 1898HW92
| |--*N. parvicornis Blackburn 1898HW92
| `--N. perkinsi Blackburn 1907HW92
|--Biphyllocera White 1841 [=Diphyllocera Agassiz 1847]HW92
| |--*B. kirbyana White 1841HW92 [=Diphyllocera kirbyanaM86]
| `--B. fabriciana White 1841HW92
|--Glossocheilifer Blackburn 1898HW92
| |--*G. labialis Blackburn 1898 [incl. G. addendus Blackburn 1907]HW92
| `--G. bidentatus Lea 1919HW92
|--Teluroides Britton 1990HW92
| |--*T. suturalis (Lea 1920) [=Telura suturalis]HW92
| `--T. clypealis (Lea 1919) [=Telura clypealis]HW92
|--Sciton Blackburn 1892HW92
| |--*S. ruber Blackburn 1892HW92
| |--S. flavocastaneus Lea 1917HW92
| |--S. paullus Blackburn 1898HW92
| `--S. variicollis Blackburn 1907HW92
|--Neso Blackburn 1898HW92
| |--*N. usta Blackburn 1898HW92
| |--N. castaneus (Lea 1920) [=Engyops castaneus]HW92
| |--N. ducalis Blackburn 1907HW92
| |--N. flavipennis (Macleay 1887) [=Platydesmus flavipennis; incl. N. planicollis Blackburn 1898]HW92
| |--N. minuta (Lea 1926)HW92 [=Haplonycha minutaHW92, Pteroplatydesmus minutusB86]
| `--N. yorkensis Blackburn 1898HW92
|--Frenchella Blackburn 1892HW92
| |--*F. lubrica Blackburn 1892HW92
| |--F. cribriceps Lea 1919HW92
| |--F. fimbriata Lea 1919HW92
| |--F. gagatina Lea 1919HW92
| |--F. hirticollis Blackburn 1898HW92
| |--F. iridescens (Blanchard 1850) [=Haplonycha iridescens]HW92
| `--F. russula Britton 1959HW92
`--Colobostoma Blanchard 1850 (see below for synonymy)HW92
|--*C. rufipennis (Boisduval 1835) (see below for synonymy)HW92
|--C. castaneus (Lea 1919) [=Platydesmus castaneus]HW92
|--C. hirsuta (Frey 1966) [=Frenchella hirsuta]HW92
|--C. inamoenus (Blackburn 1907) [=Platydesmus inamoenus]HW92
|--C. inusitatus (Blackburn 1907) [=Platydesmus inusitatus]HW92
|--C. major (Blackburn 1907) [=Platydesmus major]HW92
|--C. obscuricornis (Blanchard 1850) [=Haplonycha obscuricornis]HW92
`--C. punctulaticeps (Blackburn 1908) [=Platydesmus punctulaticeps]HW92
Leucophilis [Leucopholini]TB21
|--L. irrorata Chevrolat 1841TB21
`--L. roridaB28
Oncerini [Oncerinae]RJS02
|--Oncerus LeConte 1856RJS02
| `--O. floralis Leconte 1856S38
`--Nefoncerus Saylor 1938S38
`--*N. convergens (Horn 1894) [=Oncerus convergens]S38
Podolasiini [Lasiopodes]RJS02
|--Podolasia Harold 1869 [=Lasiopus LeConte 1856 (preoc.)]RJS02
`--Podostena Howden 1997RJS02
Diplotaxis Kirby 1837 [incl. Alobus LeConte 1856, Diazus LeConte 1860, Orsonyx LeConte 1856; Diplotaxini]RJS02

*Anthotocus luridus (Macleay 1864)HW92 [=Cheiragra luridaHW92, Phyllotocus luridaB57; incl. P. truncatidens Lea 1920HW92]

Colobostoma Blanchard 1850 [incl. Platydesmus Macleay 1887 non Lucas 1843, Pteroplatydesmus Dalla Torre 1912]HW92

*Colobostoma rufipennis (Boisduval 1835) [=Sericesthis rufipennis; incl. Frenchella sparsiceps Blackburn 1898, *Platydesmus sulcipennis Macleay 1887, *Pteroplatydesmus sulcipennis]HW92

Comophorina Strand 1928 [=Comophorus Blanchard 1850 non Agassiz 1846, Comophus Britton 1978; Comophini, Comophorini, Comophorinini]HW92

Ophropyx ciliata (Boisduval 1835)HW92 [=Melolontha ciliataHW92, Aplonycha ciliataB86, Haplonycha ciliataB86, Heteronyx ciliataB86; incl. Frenchella aspericollis Blackburn 1898HW92, Ha. rugosa Burmeister 1855HW92]

Phyllochlaenia Blanchard 1846 [=Philochlaenia (l. c.); incl. Caulobius Guillou 1844 non Duponchel 1838, Deuterocaulobius Dalla Torre 1912]HW92

*Phyllochlaenia rufescens Blanchard 1846HW92 [=Caulobius rufescensM86, Deuterocaulobius rufescensB57, Heteronyx rufescensB57, Philochlaenia rufescensB57; incl. Caulobius advena Blackburn 1898HW92]

Phyllochlaenia villosus (Guillou 1844)HW92 [=*Caulobius villosusHW92, *Deuterocaulobius villosusHW92; incl. Hymenoplia villigera Blanchard 1846HW92, Caulobius villigerM86, Omaloplia villigeraB57]

Webbella labralis (Blackburn 1908) [=Heteronyx labralis; incl. H. anomalus Blackburn 1908, H. leai Blackburn 1909]B00

*Type species of generic name indicated


[A06] Ahrens, D. 2006. The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae). Systematic Entomology 31: 113–144.

[B28] Betrem, J. G. 1928. Monographie der Indo-Australischen Scoliiden mit zoogeographischen Betrachtungen. H. Veenman & Zonen: Wageningen.

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