Aphodiinae (Insecta: Coleoptera: Scarabaeidae)

64 Pages·2001·1.1 MB·

by  Stebnicka, Z Teresa

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Fauna of New Zealand
Ko te Aitanga Pepeke o Aotearoa
Number / Nama 42
Aphodiinae
(Insecta: Coleoptera: Scarabaeidae)
Z. T. Stebnicka
Institute of Systematics and Evolution of Animals
Polish Academy of Sciences, Cracow 31–016, Poland
[email protected]
Manaaki
Whenua
P R E S S
Lincoln, Canterbury, New Zealand
2001
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Cataloguing in publication
STEBNICKA, Z. T. (Zdzis»awaTeresa), 1932–
Aphodiinae (Insecta: Coleoptera: Scarabaeidae) / Z. T. Stebnicka
– Lincoln, Canterbury, N.Z. : Manaaki Whenua Press, 2001.
(Fauna of New Zealand, ISSN 0111–5383 ; no. 42).
ISBN 0-478-09341-1
I. Title II. Series
UDC 595.764.1(931)
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printing at Landcare Research, Mt Albert Research Centre, Private Bag 92170, Auckland, New Zealand
M~ori text by H. Jacob, Huatau Consultants, Levin.
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Front cover: Acrossidius tasmaniae (Hope) (Illustrator: D. W. Helmore).
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Fauna of New Zealand 42 5
POPULAR SUMMARY HE WHAKARAPOPOTOTANGA
Class Insecta
Order Coleoptera
Family Scarabaeidae
Subfamily Aphodiinae
Illustration / Whakaahua: Phycocus graniceps Broun (Il-
lustrator / Kaiwhakaahua: A. C. Harris).
Dung beetles Ng~~~~~ p§§§§§tara tãããããtae
Representatives of the Aphodiinae are found throughout
Kitea ai ng~ mea o te wh~nau iti Aphodiinae i ng~ koko
the world but it is generally assumed that the warm part
katoa o te ao, heoi ko te whakapae, ko ng~ w~hi mahana
of the Old World is their centre of distribution. The New
o te ‘Ao Tawhito’ tÇ r~tou tino k~inga. O ng~ iwi me ng~
World and Australia, on the other hand, possess only very
puninga ka kitea i reira, he ruarua noa iho ka kitea anÇ i
few members of those tribes and genera that occur in the
te ‘Ao Hou’ me Ahitereiria. Heoi, ko te nuinga o ng~
Old World — instead, endemic genera and species of other
puninga me ng~ iwi ka kitea i ‘nei takiw~, k~ore e kitea i
tribes of Aphodiinae are numerous.
w~hi k‘.
Approximately 3100 species of aphodiines are known,
Kei te ~hua 3100 ng~ momo aphodiine e mÇhiotia
and many more may be expected, especially from the
ana, otir~, ar~ noa atu pea ng~ momo k~ore anÇ i ~ta
Neotropical and Oriental regions. The beetles are vari-
tautuhia. E tino h~ngai ana t‘nei whakapae ki ng~ takiw~
ously shaped and sculptured, and range from 0.8 mm to
o te Pã o Amerika me Amerika ki te Tonga, tae atu ki
16.0 mm in length. They are most commonly called “dung
}hia. He rerek‘ te hanga o t‘n~, o t‘n~, ~, ko te rahi, kei
beetles”, though in fact only a part of this subfamily is
waenga i te 0.8 mm me te 16.0 mm. Ko te ‘p§tara tãtae’
usually collected in dung. Aphodiinae are coprophagous
tÇ r~tou ingoa k~rangaranga, engari ko t‘tahi w~hanga
or saprophagous, feeding and breeding in the soil on vari-
noa iho o t‘nei wh~nau iti ka kitea e noho ana i te tãtae.
ous kinds of excrement and/or in vegetable debris. Sev-
He kai tãtae, he kai popo ng~ aphodiinae, ~, ka kitea e
eral species found in Europe, Asia, America, and Austra-
kai ana, e whakaputa uri ana i roto i ‘nei mea i te oneone.
lia, and two species (Acrossidius tasmaniae (Hope) and
Kua rangahaua ng~ momo maha tonu e noho ana i âropi,
Ataenius picinus Harold) in New Zealand have been stud-
i }hia, i Amerika, me Ahitereiria, me ‘tahi momo e rua o
ied as minor pests of cultivated plants. Some species, e.g.,
Aotearoa (ko te Acrossidius tasmaniae (Hope) me te
Phycocus graniceps Broun and Tesarius sulcipennis (Lea),
Ataenius picinus Harold), he kaikai m~ra nÇ r~tou. Ar~
are strongly associated with coastal sand dunes. Many
anÇ ‘tahi momo p‘r~ i a Phycocus graniceps Broun me te
species in other countries are associated with ants and
Tesarius sulcipennis (Lea) e kitea nuitia ana ki ng~ t~huna
termites and some others live in the burrows of small
o te tahamoana. He maha ng~ momo i t~w~hi e noho ana
mammals. A few species are known to be kleptoparasitic,
i waenga i a ng~i pÇpokorua me ng~i pÇpokotea, ko ‘tahi
breeding in the brood balls of nest-building dung beetles
anÇ e noho ana ki ng~ rua o ‘tahi kararehe wh~ngote
(Scarabaeidae: Scarabaeinae). Adults of Aphodiinae are
moroiti. Ko ‘tahi momo ruarua nei, ka wh~nako i ng~
attracted to light and are often found in various kinds of
kÇhanga o ng~ p§tara tãtae hanga kÇhanga (ar~ ng~
excrement, in decaying vegetation, under logs and moss,
Scarabaeidae: Scarabaeinae), hei w~hi whakaputa uri mÇ
in rotten wood, and under loose bark of dead trees. The
r~tou. Whaiwhai ai ng~ aphodiinae pakeke i te m~rama.
immatures are known for a relatively small number of
Ka kitea nuitia hoki i ng~ momo tãtae maha, i ng~ popo,
species.
(continued overleaf) (haere tonu)
6 Stebnicka (2001): Aphodiinae (Insecta: Coleoptera: Scarabaeidae)
Twenty species of Aphodiinae are now known from i raro i ng~ poro r~kau me ng~ pãkohu, i ng~ r~kau pirau,
New Zealand. Eleven of these have been introduced, prob- i raro r~nei i te hiako o ng~ r~kau kua mate noa. E mÇhiotia
ably through human commerce — six are from Australia, ana te hanga kÇhungahunga o ‘tahi momo ~hua ruarua
two from America, one from Africa, and the last two are nei.
cosmopolitan species of European derivation. E rua tekau ng~ momo Aphodiinae o Aotearoa e
The remaining nine species of the genera Phycocus mÇhiotia ana in~ianei. Tekau m~ tahi o ‘nei, kua rere mai
Broun (introduced to Tasmania) and Saprosites i t~w~hi, n~ ng~ mahi tauhokohoko pea i heri mai. E ono
Redtenbacher are indigenous to New Zealand. One spe- i tau mai i Ahitereiria, e rua mai i Amerika, kotahi mai i
cies is found only on the South Island, and seven are found }wherika, e rua atu anÇ nÇ âropi taketake ake, engari
only on the North Island including one species collected kua puta atu ki ng~ hau e wh~.
also on the Kermadec Islands, and one on the Chathams
T‘r~ anÇ ng~ momo e iwa o te puninga Phycocus
Islands. One species is indigenous to the Three Kings
Broun (kua tae atu anÇ ki Tahim~nia) me Saprosites
Islands, one to the Chatham Islands, and one to the
Redtenbacher, he momo tãturu nÇ Aotearoa. Kotahi te
Kermadec Islands (introduced to the North Island). The
momo ka kitea i Te Waipounamu anake, e rima ka kitea i
non-endemic species, of adventive origin are frequently
Te Ika a M~ui anake, kotahi ka kitea i Te Ika a M~ui me
abundant and conspicuous in towns, orchards, pastures,
ng~ moutere Kermadec, kotahi anÇ i Te Ika a M~ui me
and other modified environments. The faunal relation-
R‘kohu. Kotahi te momo nÇ Manawa-t~whi tãturu, kotahi
ships of the native New Zealand Aphodiinae are clearly
nÇ R‘kohu tãturu, kotahi nÇ ng~ Kermadec tãturu (kua
closest to those of Australia.
tau mai anÇ ki Te Ika a M~ui). Ko ng~ momo ehara nÇ
konei taketake ake, engari kei konei e noho ana, ka kitea
nuitia i ng~ taone, i ng~ m~ra huar~kau, i ng~ p~t§t§, me
‘r~ atu taiao n~ te ringa tangata i raweke. Ko ng~ hono i
waenga i ng~ Aphodiinae o Aotearoa taketake ake, e ~hua
rite ana ki ‘r~ o Ahitereiria.
I wh~nau mai te kaituhi, a Zdzis»awa Teresa Stebnicka
(ko Eichler tana ingoa wh~nau), i PÇrana. Ko ng~ mahi
toi me ng~ pãtaiao m~ori ~na kaupapa matua i te whare
w~nanga, ~ ka whakawhiwhia ia ki tana T~kutatanga
Pãtaiao i te W~nanga Pãtaiao o PÇrana, i Cracow. Kei te
Pãtahi WhakarÇpã, Kunenga Kararehe, i raro i te maru o
te W~nanga Pãtaiao o PÇrana, a ia e mahi ana. Ko t~na
kaupapa rangahau, ko te whakarÇpã, te whakatairite hanga
rauropi, te koiora-matawhenua, me te kunenga o ‘tahi
rÇpã p§tara, otir~, me te aro wh~iti ki ng~ tini Aphodiinae,
huri i te ao. Neke atu i te 80 ng~ kÇrero n~na anÇ i tuhi e
Contributor Zdzis»»»»»awa Teresa Stebnicka, family name
titiro ana ki te pãnaha whakarÇpã, te w~hi noho me te
Eichler, was born in Poland, educated in fine arts and
~hua o t‘n~, o t‘n~ momo, te taupuhi kaiao, tae atu ki te
natural sciences, and was awarded a Dr. Sc. degree at the
koiora-matawhenua o ng~ aphodiine, me ‘tahi atu huinga
Polish Academy of Sciences in Cracow. She is employed
Coleoptera maha. [Whakaahua: T~kuta Stebnicka i te Whare
at the Institute of Systematics and Evolution of Animals
Taiao o K~nata, Ottawa, Pipiri 2000; n~ François Génier.]
of the Polish Academy of Sciences where her research is
on the taxonomy, comparative morphology, biogeogra-
phy, and phylogeny of various groups of beetles, and in Translation by H. Jacob
particular the Aphodiinae of the world. She has authored Huatau Consultants, Levin
over 80 original papers and books covering systematics,
faunal, ecological, and biogeographical treatments of
aphodiines and many other groups of Coleoptera. [Photo-
graph: Dr Stebnicka at the Canadian Museum of Nature, Ottawa,
June 2000; by François Génier].
Fauna of New Zealand 42 7
ABSTRACT
The Aphodiinae of New Zealand are comprehensively revised for the first time
and their relationships are discussed. Twenty species in 4 tribes and 9 genera are
recognised. Two new species are described (Saprosites kingsensis, Saprosites
watti), and 2 new synonymies are proposed (Ataenius macilentus Blackburn,
1904 = A. brouni (Sharp, 1876); Saprosites candens (Broun, 1880) = S. distans
(Sharp, 1876)). Eleven of the 20 species are introduced, 6 of them originating in
Australia, 2 in America, and 1 in Africa; the other 2 are cosmopolitan anthropo-
genic species of European origin. Members of the genus Saprosites Redtenbacher
are endemic to New Zealand, except for 1 Australian species. The species of
Phycocus and Tesarius are indigenous to both New Zealand and Tasmania. The
morphology and diagnostic characters of aphodiines are reviewed and illustrated
in detail. A key is given for the tribes, genera, and species known from New
Zealand. The biological associations, known seasonality, and geographic distri-
bution of the species are summarised, with maps indicating locality records. The
distributions of the indigenous species and their systematic affinities suggest a
Gondwanan origin.
Keywords. Coleoptera, Scarabaeidae, Aphodiinae, New Zealand, taxonomy,
classification, key, new species, new synonymy, distribution, ecology, biology,
dispersal power, species endemism, fauna.
Stebnicka, Z. T. 2001. Aphodiinae (Insecta: Coleoptera: Scarabaeidae). Fauna
of New Zealand 42, 64 pp.
Received: 18 March 1999. Accepted: 24 January 2001.
CHECKLIST OF TAXA
Genus Tesarius Rakovi…, 1981.................................. 22
Subfamily APHODIINAE........................................ 17 sulcipennis (Lea, 1904)....................................... 22
Tribe Aphodiini......................................................... 17 Tribe Proctophanini.................................................. 23
Genus Aphodius Illiger, 1798.................................... 17 Genus Proctophanes Harold, 1861........................... 23
granarius (Linnaeus, 1767) ................................ 18 sculptus (Hope, 1846)......................................... 23
adelaide Hope, 1846 minor (Blackburn, 1897) [interception record].. 24
lividus (Olivier, 1789)......................................... 19 Genus Australaphodius Balthasar, 1942.................. 24
cincticulus Hope, 1847 frenchi (Blackburn, 1892)................................... 25
spilopterus Germar, 1848 melbournicus (Balthasar, 1942)
pseudolividus Balthasar, 1941 Tribe Eupariini ......................................................... 25
lividus pseudolividus Endrödi & Rakovi…, 1981 Genus Parataenius Balthasar, 1961.......................... 25
Genus Acrossidius Schmidt, 1913............................ 19 simulator (Harold, 1868).................................... 26
tasmaniae (Hope, 1847)...................................... 20 schwarzi (Linell, 1896)
howitti (Hope, 1847) granuliceps Petrovitz, 1971
longitarsus (Redtenbacher, 1867) Genus Ataenius Harold, 1867................................... 26
andersoni (Blackburn, 1904) brouni (Sharp, 1876)........................................... 27
pallidihirtus (Balthasar, 1941) macilentus Blackburn, 1904 new synonymy
Tribe Psammodiini.................................................... 21 picinus Harold, 1867........................................... 28
Genus Phycocus Broun, 1886................................... 21 duplopunctatus Lea, 1923
graniceps Broun, 1886........................................ 21 boucomonti Paulian, 1937
lobatus Broun, 1893 rugosus (Richards, 1959)
8 Stebnicka (2001): Aphodiinae (Insecta: Coleoptera: Scarabaeidae)
Genus Saprosites Redtenbacher, 1858...................... 29 calities with their area codes, and in preparation of the
mendax (Blackburn, 1892) ................................. 30 maps. Des Helmore prepared many of the habitus illus-
punctatus Richards, 1959 trations and Birgit Rhode the SEM’s. Rowan Emberson
exsculptus (White, 1846).................................... 31 reviewed the manuscript.
pascoei (Sharp, 1876) The present contribution was supported in part by
suspectus (Sharp, 1876) Grant Nr 6 P04C 055 15 received from the Committee of
distans (Sharp, 1876).......................................... 32 the Scientific Research (KBN) in Poland.
candens (Broun, 1880) new synonymy
raoulensis (Broun, 1910).................................... 33
INTRODUCTION
communis (Broun, 1880).....................................33
kingsensis sp. nov................................................ 34 The Aphodiinae is a large and diverse group of species
fortipes (Broun, 1881)......................................... 35 inhabiting all zoogeographical regions. The fauna of
watti sp. nov......................................................... 36 aphodiines is generally well known for most of the world
sulcatissimus (Broun, 1911)............................... 37 with the exception of Central and South America. These
areas are the object of current studies of the author. The
faunas of Australia, New Guinea, and neighbouring ar-
CONTENTS
chipelagos have been recently treated in a series of pa-
Acknowledgments........................................................ 8 pers by Stebnicka & Howden (1994, 1995, 1996, 1997)
Introduction.................................................................. 8 and Stebnicka (1998b). Before the present contribution
Systematics................................................................... 9 was undertaken, very little was known of the faunistic
Morphology.................................................................. 9 composition of Aphodiinae in New Zealand. Early treat-
Diagnostic characters for tribes in N.Z................ 11 ments of New Zealand aphodiines from 1846 to 1910
Biology ....................................................................... 12 consist of isolated species descriptions by Broun (6 spe-
Biogeography.............................................................. 13 cies), Sharp (4 species), White (1 species), and Lea (1
Collecting and preparation methods.......................... 15 species). The first native New Zealand species to be dis-
Conventions................................................................ 15 covered was Saprosites exsculptus (as Oxyomus
Key ....................................................................... 16 exsculptus) which was collected at “Port Nicholson”
Descriptions................................................................ 17 (Wellington) and described by White (1846). Other than
References.................................................................. 38 the listing of the species in a catalogue (Schmidt 1910)
Illustrations................................................................. 43 and their misleading diagnoses in a monograph by Schmidt
Distribution maps....................................................... 53 (1922), the only subsequent papers dealing with the tax-
Taxonomic index........................................................ 58 onomy of New Zealand species were the review of the
genus Saprosites by Richards (1959) and some synonymi-
cal notes by Watt (1984). The status of nearly all species
ACKNOWLEDGMENTS
of Saprosites has been particularly enigmatic, since their
The Biodiversity & Conservation Group of Manaaki types were not readily accessible to researchers as they
Whenua initiated this study when they contacted the au- were in European collections (mostly in BMNH), and
thor in 1996 concerning a possible contribution to the specialists on Aphodiinae never saw them.
Fauna of New Zealand on New Zealand Aphodiinae. This The present survey is based mostly on a large holding
request led over the intervening years to the present revi- of material in the New Zealand Arthropod Collection,
sion. During the period of this study I have enjoyed the Auckland, and on the collections of various institutions
co-operation in arranging loans of specimens, including mentioned in the section on Conventions. Approximately
types, of a succession of individuals at several institu- 1800 specimens and the type-series of all New Zealand
tions. I here thank them, and they are listed with their species were examined. Twenty species are now known
respective institutions in the section on Abbreviations. from New Zealand, nine of which are indigenous. Eleven
In particular I would like to acknowledge the late are definite introductions from other countries; six from
Tymone Duval, Trevor Crosby, and Richard Leschen of Australia — Acrossidius tasmaniae (Hope), Tesarius
the New Zealand Arthropod Collection and Rowan sulcipennis (Lea), Proctophanes sculptus ((Hope), P. mi-
Emberson and John Marris of the Lincoln University for nor ((Blackburn), Ataenius brouni (Sharp), Saprosites
their unfailing encouragement and co-operation. Grace mendax (Blackburn); two from the Americas —
Hall assisted greatly through association of specimen lo- Parataenius simulator (Harold) and Ataenius picinus
Fauna of New Zealand 42 9
Harold; one from Africa — Australaphodius frenchi one time or another been treated as subfamilies, how-
(Blackburn); and two cosmopolitan species of ultimate ever, they are not separated by a decided gap (if any), and
European origin — Aphodius lividus (Olivier) and A. none of the taxa covered here can seriously be elevated to
granarius (Linnaeus). Two species from the remaining subfamily rank. As in other groups of Scarabaeoidea, there
nine are here described as new, and the genera Tesarius has been considerable parallel evolution within the sub-
and Parataenius are recognised from New Zealand for family, and almost all characters claimed to be diagnostic
the first time. In general, the New Zealand fauna of at tribal or generic level have evolved independetly sev-
aphodiines is similar to that of Australia but strongly im- eral times. In some cases, groups which are easily sepa-
poverished in the number of genera and species. The bio- rated in the Old World, e.g., Psammodiini–Eupariini–
geographic affinities of the New Zealand Aphodiinae Aphodiini, become almost indistinguishable in South
clearly indicate an ancient, Gondwanan origin in com- America or Australia. Often seemingly distinct genera in
mon with the Australian fauna. temperate regions break down in the tropics and poten-
tially strong defining characters are diluted and must be
qualified. The following characterisation of Aphodiinae
on a world basis and diagnostic characters for tribes found
SYSTEMATICS
in New Zealand clearly indicate that all taxa above spe-
The higher classification of the subfamily Aphodiinae cies level have to be defined by a mosaic of characters, a
(Lawrence & Newton 1995) has varied considerably over “polythetic classification” — no character in isolation be-
the last 80 years, and is still not completely settled. The ing diagnostic for all members.
basic systematic arrangement for the world species pro-
posed by Schmidt (1922), based on 1137 species known
at that time, is that on which new additions have been
MORPHOLOGY
based. The latest world catalogue (Dellacasa 1988, 1989,
1991, 1996) lists about 3100 species, but additional taxa Characterisations of Aphodiinae (Aphodiidae of authors)
have been described since. The tribe Aphodiini forms the hitherto given by many authors are far from being ex-
core of the subfamily, and is based on the huge genus haustive and concern mainly superficial, often second-
Aphodius Illiger, with over 30 associated genera. The ary, characters from casual examples of various groups of
genus Aphodius is actually split into about 135 subgenera, species. The following, general description is based on
with a number of additional species that cannot be prop- all known tribes, of which only four are represented in
erly assigned. Consequently, the existing keys for tem- New Zealand. The principal characters of the subfamily
perate Aphodius have become unworkable, and identifi- are first listed here in a synoptic form (excluding sexual
cation of species very difficult. My belief is that the use dimorphism), and then the tribal characters are discussed
of species groups permits organisation within large gen- at greater length.
era (i.e., Ataenius: Stebnicka & Howden 1997) without General characters used in Aphodiinae for ident-
the attendant nomenclatural problems associated with ification are illustrated in Fig. 3–6. A number of species
subgeneric names. It is obvious in these circumstances have unusual or unique characters which allow them to
that a compromise must often be made between the prac- be identified without needing to use keys and descriptions.
tical aims of classification, its phylogenetic basis, and a
system of naming.
The higher categories, such as tribes, genera, and spe- PRINCIPAL CHARACTERS OF APHODIINAE
cies groups are poorly understood and nearly every au-
Body. Shape and sculpture diverse; length 0.8 mm to 16.0
thor has a different arrangement. Part of the problem is
mm from clypeal apex to elytral apex.
due to the tremendous numbers of species, some of which
Head. Clypeus dilated to cover mouthparts or shortened,
fill the gaps between any arrangement of higher catego-
exposing mouthparts or clypeus of intermediate type.
ries so far devised. Examples of confusion in relation-
Antenna short, of 9, rarely 8, antennomeres, club cir-
ships and classification are numerous, resulting from dif-
cular, ovoid, or elongate, of 3 antennomeres.
ferent levels of knowledge of the various species groups
Eye very large, moderate in size, vestigial or absent.
by authors studying only limited faunal areas. Recently a
trend may be observed of a category inflation. The break- Mouthparts. Maxillary endites (galea and lacinia) sepa-
ing up of polyphyletic groups does not necessarily led to rate or approximate, consisting of soft, flexible, densely
a more practical system, if we do not know where the setose lobes; third maxillary palpomere cylindrical, fusi-
fragments belong. Various tribes of Aphodiinae have at form or triangular, its length varied.
10 Stebnicka (2001): Aphodiinae (Insecta: Coleoptera: Scarabaeidae)
Either: scissorial lobes of mandibles sclerotised, den- Abdomen. Five or six sternites visible with intersegmen-
tate, or truncate; labrum partially sclerotised and more or tal membranes or sternites coalesced; pygidium covered
less exposed in front of clypeus, e.g., Aegialiini (indi- by elytra or partially exposed, e.g., Proctophanini,
cates “hard saprophagy” as a component of feeding) Odontolochini, and Eupariini.
Or: scissorial lobes of mandibles dorso-ventrally flat- Mesocoxae. Approximate (space between mesocoxae
tened, flexible plate-like; labrum membranous, fully cov- smaller than width of mesofemur), or moderately sepa-
ered by clypeus, e.g., Aphodiini (indicates “soft rate (space between mesocoxae equal to width of
saprophagy or coprophagy” as a component of feeding). mesofemur), or widely separate (space between
Labrum. Anterior margin evenly rounded, truncate, mesocoxae nearly twice as wide as mesofemur); coxal
bilobed or trilobed; inner surface — epipharynx, consists cavities parallel or oblique, elongate, rarely round, con-
of various elements and sensory organs (Stebnicka 1985, vergent in shape with those of Zopheridae (Tenebrionoidea
Stebnicka & Howden 1995, 1996). [Labro-epipharyngeal and many other beetles), e.g., Neotropical tribe
structures are of some use in species separation, e.g., Lomanoxiini (Stebnicka 1999b).
Aphodiini, and especially indicative at a tribal level].
Metacoxae. Always contiguous; trochanters short, rarely
Pronotum. Rectangular or subquadrate in shape; basal elongate, e.g., Phycocus Broun.
and lateral edges margined or not, and/or crenate,
Legs. Middle and hind legs short (femora not or barely
explanate or sulcate or otherwise modified; disc evenly
visible from above), or moderate in length (femora ex-
convex or sulcate, fossulate and/or carinate, or with hump.
tending a little beyond body outline), or long (femora pro-
Scutellum. From very large to minute; shape triangular, jecting for about 1/3 their length).
pentagonal, semi-oval, or cordate. Profemur short, enlarged, flattened dorso-ventrally;
Elytra. Striate or not; usually with 10 striae and 10 inter- meso- and metafemur parallel-sided or fusiform, some-
vals including sutural ones, or number of striae reduced; times posterior edge dentate.
rarely elytra non-striate, convergent in sculpture like those Protibia short or moderate in length, usually with ter-
of Tenebrionidae, e.g., African Eremazus Mulsant; minal spur; outer side with 1-5 teeth or serrate, e.g., Asian
epipleura gradually narrowed to apices, abruptly narrowed Setylaides Stebnicka.
or absent posteriorly, or widely inflexed, e.g., Neotropical Meso- and metatibia cylindrical, or flattened dorso-
Euparixia Brown, Euparixoides Hinton, and Lomanoxia ventrally or parallel-sided; transverse ridges clearly de-
Martinez, or with edge upturned, e.g., Neotropical Selviria veloped, e.g., Aphodiini, Proctophanini, and some
Stebnicka (1999a), and Nearctic Hornietus (Stebnicka Aegialiini, Psammodiini, and Eupariini, or vestigial, e.g.,
2000b); humeral umbone strongly or moderately devel- some Aphodiini, Psammodiini, Aegialiini, and Eupariini,
oped (indicates wings present even if reduced), or hu- or absent, e.g., Didactyliini, Odontolochini, Eupariini, and
meral area flattened and elytra egg-shaped (indicates Psammodiini, or tibia otherwise sculptured, e.g.,
wings absent), e.g., Aphodiini, Aegialiini, Psammodiini, Eupariini; apex of meso- and metatibia with two spurs,
and Eupariini; in some species-groups elytra with rarely with one spur; apical spurs of metatibia slender,
preapical umbone, e.g., Australian Podotenus Schmidt. spatulate, or foliaceous, placed on each side of tarsal in-
Wings. Fully developed and functional, or non-functional, sertion, e.g., Aphodiini and Aegialiini, or placed close
brachypterous, or absent. together below tarsal insertion, e.g., Didactyliini,
Odontolochini, Proctophanini, Psammodiini, and
Prosternum. Usually with median tubercle, or with tri-
Eupariini, or intermediate character state occurs, e.g.,
angular or hastate process.
some Aphodiini and Eupariini.
Mesosternum. Evenly convex or deplanate and/or cari-
External sexual dimorphic characters. Strongly devel-
nate, or with variously shaped callosities, e.g., Asian-
oped to invisible or absent.
Australian Airapus Stebnicka & Howden, and Neotropical
Auperia Chevrolat and Euparixoides Hinton. Male genitalia. Phallobase of aedeagus usually with dor-
sal hump, junction of phallobase and parameres usually
Metasternum. Approximately equal to the length of
membranous, e.g., Aphodiini, or phallobase of aedeagus
mesosternum (indicates wings present even if reduced),
evenly rounded, phallobase and parameres sometimes
or significantly shorter than mesosternum (frequently
coalesced, e.g., Eupariini and Psammodiini; apical por-
wings absent but not always); surface usually with me-
tion of internal sac membranous, frequently furnished with
dian longitudinal line or furrow, sometimes with pits;
sclerites and/or spicules.
posterior plates frequently with metasternal triangle, e.g.,
Eupariini and Psammodiini. Female genitalia. Genital plate “stylus” (the divided 10th