Paratoxicum iridescens, from Matthews & Bouchard (2008).

Belongs within: Tenebrionoidea.
Contains: Adesmia, Lagriinae, Phrenapatinae, Cossyphodini, Sepidiini, Pimeliini, Coniontini, Cnemeplatiini, Vacronini, Anepsiini, Asidini, Epitragini, Eurymetopini, Nycteliini, Tentyria, Alleculinae, Diaperinae, Stenochiinae, EleodesHelops, Toxicini, Blaps, Amarygmini, Bolitophagini, Ulomini, Opatrini, Palorini, Pedinini, Heleini, Triboliini, Titaenini, Tenebrio.

The Tenebrionidae are a large and varied family of beetles, most of which are scavengers on dead plant and fungal material, and are one of the most successful families of beetles in arid environments. The majority of species are heavily sclerotised and often dark in coloration. Many tenebrionids are able to discharge defensive chemicals from glands in the anal region, though these glands are absent in some significant subgroups such as the Pimeliinae. In those groups lacking defensive glands, the sternites of the abdomen hinge along the midline but those tenebrionids with defensive glands have sternites hinging laterally to allow the abdomen to expand before expelling the defensive fluid (Matthews & Bouchard 2008).

The Tenebrionidae may be divided between three major lineages, the Lagriiformes, Pimeliiformes and Tenebrioniformes, based primarily on features of the male and female reproductive systems (Matthews & Bouchard 2008). Members of the Lagriiformes branch include the Neotropical genus Nilio, false ladybird beetles, adults of which have a circular outline and a setose, nearly hemispherical dorsal surface (Bouchard 2014). The Pimeliinae are the most diverse subgroup of the tenebrionids in arid environments in most parts of the world except Australia (Matthews & Bouchard 2008). The Zolodininae are a small Australasian group that differ from other tenebrionids in the presence of externally open procoxal cavities (Matthews & Bouchard 2008).

Many members of the Tenebrioniformes possess complex, stellate antennal sensoria. The Tenebrioninae tend to have fairly small defensive gland reservoirs with broad openings, and the bursa copulatrix in the female genital system tends to be lost and be replaced by a spermatheca that may be coiled in a tight ball (Matthews & Bouchard 2008).

Darklings, tok toks and pie-dishes
Published 7 January 2018
False wireworm beetle Gonocephalum sp., copyright EBKauai.

It has been noted to the point of cliché that the Creator has an inordinate fondness of beetles. Even within the massive range of beetle diversity, though, certain families stand out as particularly diverse. One such family is the Tenebrionidae, with over twenty thousand known species worldwide. The family is sometimes referred to as the darkling beetles but no one vernacular name is really sufficient for this group. Not only are tenebrionids taxonomically diverse, they are morphologically diverse, varying from long-legged and elongate to hemi-spherical and robust, from smooth and shining to ornate and hairy, from dull-coloured and retiring to bright and striking. Habits vary from detritivorous to xylophagous (feeding on decaying wood) to herbivorous to mycetophagous, with even a few predators. Larvae of some species are of economic significance as pests: the false wireworms feed on the roots of crops or lawns, while mealworms and flour beetles attack stored products (mealworms are, of course, also used as pet food and occasionally even as human food). Several species live as inquilines of social insects such as ants or termites. The highest diversity of tenebrionids is in relatively arid regions; some species, such as the tok tok beetles of southern Africa and the pie-dish beetles of Australia, are familiar sights in such habitats.

Pie-dish beetle Helea sp., copyright Australian Museum.

With such high diversity, it is not easy to define this group without encountering exceptions, but generally tenebrionids have the antennae eleven-segmented and inserted below lateral expansions of the genae. The procoxal cavities are usually closed externally, and the legs of most species have a 5-5-4 tarsal formula. The first three sternites of the abdomen are fused (Kergoat et al. 2014). Several subfamilies are recognised, but they are commonly grouped into three clusters known as the lagrioid, pimelioid and tenebrionoid branches of the family (Matthews & Bouchard 2008). Many members of the lagrioid and tenebrionoid branches possess well-developed defensive glands in the abdomen. The rear sternites of the abdomen in these species are hinged on the sides rather than along the midline as in more primitive forms, allowing the abdomen to expand as the gland reservoirs fill with a repugnant fluid that can be expelled when required. Many larger tenebrionids have a tendency to walk with their rear ends tilted upwards, ready to unleash at a moment’s notice.

Allecula rhenana, copyright Stanislav Krejčík.

Members of the pimelioid branch, including the subfamilies Pimeliinae and (possibly) Zolodininae, lack abdominal defensive glands. In many parts of the world, pimelioids are the dominant tenebrionids in dry habitats. The lagrioid branch includes the single subfamily Lagriinae, defined by features of the genitalia. Matthews & Bouchard (2008) also listed the small subfamily Phrenapatinae in this branch but a molecular phylogenetic analysis of the family by Kergoat et al. (2014) placed this latter subfamily in the tenebrionoid branch. The tenebrionoid branch also includes the Tenebrioninae, Diaperinae, Alleculinae and Stenochiinae, though monophyly of the Tenebrioninae and Diaperinae is uncertain (Kergoat et al. 2014). Diaperines include a number of shiny, sometimes strikingly coloured species; members of the tribe Leiochrinini look more like ladybeetles of the Coccinellidae than typical tenebrionids. The Tenebrioninae include such notable members as the false wireworms of the tribe Opatrini, the mealworms of the Tenebrionini and the flour beetles of the Triboliini. Finally, the Alleculinae are a distinctive group of often relatively soft-bodied tenebrionids readily distinguished from other members of the family by their pectinate claws; in some older classifications, alleculines were treated as a separate family of their own.

Systematics of Tenebrionidae

Characters (from Lawrence & Britton 1991): Usually black or brown and subglabrous, with concealed antennal insertions and usually stout, moniliform or incrassate antennae. Eyes usually emarginate; lateral pronotal carinae usually simple and complete; prosternal process usually moderately broad, strongly curved dorsally behind coxae, neither strongly nor abruptly expanded apically; junctions of first three ventrites different from those separating last two, the basal ventrites being strongly connate and the apical ones more or less movable, often with exposed membrane. Externally open fore coxal cavities only occur in Zolodininae, internally open ones in some Tenebrioninae. Tarsal formula usually 5-5-4, sometimes 4-4-4. Most species with paired glandular reservoirs, associated with chemical defence sustem, located usually at base of fifth ventrite. Larvae almost always elongate (ovate in Leiochrini) and subcylindrical to slightly flattened. May be very lightly sclerotised, moderately sclerotised or yellow in colour, with or without darker head and urogomphi, or uniformly heavily sclerotised and darkly pigmented; vestiture consists of scattered fine hairs except in Lagriini whose larvae are densely setose.

<==Tenebrionidae (see below for synonymy)
| |--LagriinaeMW15
| |--PhrenapatinaeMB08
| `--Nilio Latreille 1802L02 [=NilionL02; Nilionidae, NilioninaeMB08]
| |--*N. villosus [=Coccinella villosa]L02
| `--N. lanatus Germar 1824B14
`--+--+--Zolodininae [Zolodinini]MW15
| | |--Zolodinus zelandicusMB08, LB91
| | `--Tanylypa Pascoe 1869MB08
| | `--*T. morio Pascoe 1869MB08
| `--Pimeliinae [Pimeliiformes, Tentyriinae]MW15
| | i. s.: OnymacrisB14
| | |--O. bicolor (Haag-Butenberg 1875)B14
| | |--O. laevigatusB14
| | `--O. unguicularisB14
| | BothrolesC68
| |--CossyphodiniB14
| |--SepidiiniB14
| |--CeratanisiniMB08
| |--PimeliiniMB08
| |--LachnogyiniMB08
| |--ConiontiniAT02
| |--CnemeplatiiniMB08
| |--VacroniniMB08
| |--AnepsiiniAT02
| |--AsidiniAT02
| |--Cnemodinus Cockerell 1906 [incl. Cnemodus Horn 1870 non Herrich-Schaeffer 1850; Cnemodini]AT02
| |--EpitraginiAT02
| |--EurymetopiniAT02
| |--Salax Guérin-Méneville 1834 [Trilobocarini]MB08
| | `--*S. lacordairei Guérin-Méneville 1834MB08
| |--Nyctoporis Eschscholtz 1831 [incl. Emeax Pascoe 1866, Enneacoides Fairmaire 1881; Nyctoporini]AT02
| | `--N. cristata Eschscholtz 1831 [incl. Emeax sculpturatus Pascoe 1866]MB08
| |--Branchus LeConte 1862 [Branchini]AT02
| | |--B. floridanusAT02
| | `--B. obscurusAT02
| |--CryptochiliniB14
| | |--VansoniumB14
| | `--Calognathus chevrolatiB14
| | |--C. c. chevrolatiB14
| | `--C. c. eberlanzi Koch 1950B14
| |--NycteliiniB14
| |--StenosiniAT02
| | |--Araeoschizus LeConte 1851 [Araeoschizina]AT02
| | | `--A. andrewsiAT02
| | `--Typhlusechus Linell 1897 [Typhlusechina]AT02
| | `--T. singularisAT02
| |--Tentyriini [Gnathosini]MB08
| | |--TentyriaK03
| | `--Stenosida Solier 1835MB08
| | |--‘Tagenia’ striatopunctata Wiedemann 1821 [incl. *Stenosida tenuicollis Solier 1835]MB08
| | |--S. australis Medvedev 1995MB08
| | `--S. darevskiiMB08
| `--CryptoglossiniAT02
| |--Schizillus Horn 1874AT02
| |--Cryptoglossa Solier 1836 (see below for synonymy)AT02
| | `--C. muricataAT02
| `--Asbolus LeConte 1851AT02
| `--A. verrucosusAT02
|--Trachelostenus [Trachelostenidae]MW15
`--Tenebrioninae [Opatrinae]MW15
| i. s.: Eleodini [Amphidorini]AT02
| |--EleodesB14
| |--Embaphion Say 1824AT02
| |--Neobaphion Blaisdell 1925AT02
| |--Lariversius Blaisdell 1947AT02
| | `--L. tibialis Blaisdell 1947AT02
| |--Trogloderus LeConte 1879AT02
| | `--T. costatus LeConte 1879AT02
| `--Eleodimorpha Blaisdell 1909AT02
| `--E. bolcan Blaisdell 1909AT02
| Rhyzodina mniszechii Chevrolat 1873B14
| EulabiniAT02
| |--Apsena LeConte 1862AT02
| |--Eulabis Eschscholtz 1829AT02
| | `--E. bicarinata Eschscholtz 1829AT02
| `--Epantius LeConte 1862AT02
| `--E. obscurus LeConte 1851AT02
| CerenopiniAT02
| |--Argoporis Horn 1870 [incl. Threnus Motschulsky 1870]AT02
| `--Cerenopus LeConte 1851AT02
| `--C. concolor LeConte 1851AT02
| HelopiniAT02
| |--HelopsAT02
| |--Tarpela Bates 1870 [incl. Lamperos Allard 1876]AT02
| `--Nautes Pascoe 1866AT02
| `--N. viridimicans Horn 1878AT02
| CentronopiniAT02
| |--Metaclisa Jacquelin du Val 1861MB08, AT02 [incl. Amarantha Faust 1875AT02]
| |--Centronopus Solier 1848 (see below for synonymy)AT02
| `--Scotobaenus LeConte 1859AT02
| Apocrypha Eschscholtz 1831 [incl. Compsomorphus Solier 1851; Apocryphini]AT02
| `--A. setosaAT02
|--Cheirodes Gené 1839MB08 (see below for synonymy)
| |--*C. sardous Gené 1839 [incl. Anemia caulobioides Carter 1920]MB08
| `--C. californica (Horn 1870)AT02
|--Platynotini [Platynotina]MB08
| | i. s.: ‘Tenebrio’ dispar Herbst 1797 [incl. Menearchus impressosulcatus Carter 1920]MB08
| |--AnomalipusB14
| | |--A. elephas Fåhraeus 1870B14
| | `--A. expansicollisG91
| `--Opatrinus Dejean 1821 (see below for synonymy)AT02
| |--‘Hopatrinus’ corvinus Muls 1852G89
| |--O. gemellatus Ol. 1795 [incl. O. clathratus]FS90
| `--‘Hopatrinus’ niloticus Muls 1852G89
|--Acropteron Perty 1830FS90 [AcropteroniniMB08]
| |--A. chabrieri Fleutiaux & Sallé 1890FS90
| `--A. transversicolleFS90
| |--Diaclina Jacquelin du Val 1861MB08
| | |--*D. chrysomelina (Carter 1908) [=Tenebrio chrysomelinus]MB08
| | `--D. nitida (Carter 1908) [=Heterocheira nitida; incl. D. immaculata Gebien 1920]MB08
| `--Alphitobius Stephens 1829MB08 (see below for synonymy)
| |--A. laevigatus (Fabricius 1781) (see below for synonymy)MB08
| |--A. diaperinus (Panzer 1797) [=Tenebrio diaperinus]MB08
| `--A. piceus Ol. 1795 [incl. A. convexus, A. fagi, A. manillensis, A. orizae]FS90
|--Rhinandrus LeConte 1866 [incl. Exerestus Bates 1870, Proderops Fairmaire 1873]AT02
| `--R. sublaevisAT02
|--Bius Mulsant 1854AT02
| `--B. estriatus LeConte 1851AT02
|--Idiobates Casey 1890AT02
| `--I. castaneus (Knoch 1801)AT02
|--Neatus LeConte 1862AT02
| `--N. tenebrioides (Beauvois 1805)AT02
|--Zophobas Blanchard 1845 [incl. Macrozophobas Pic 1913]AT02
| |--Z. atratusAT02
| `--Z. morio Fab. 1777 [incl. Z. elongatus, Z. infuscatus, Z. nigritus]FS90
`--Paratoxicum Champion 1894 [incl. Schizophthalmotribolium Kaszab 1940]MB08
|--*P. iridescens Champion 1894MB08
|--P. australiae (Kaszab 1940) [=*Schizophthalmotribolium australiae]MB08
`--P. nigricans Oke 1932MB08

Tenebrionidae incertae sedis:
Zeadelium gratiosumED04
Parastizopus armaticepsGE05
Pseudognaptorina Kaszab 1977K77
`--*P. nepalica Kaszab 1977K77
Freudeia Kaszab 1961K77
|--F. granulipleuris Kaszab 1977K77
|--F. martensi Kaszab 1977K77
|--F. nepalica Kaszab 1961K77
`--F. punctipleuris Kaszab 1977K77
Blaptyscelis Koch 1961 [=Blaptyscellis Pierre 1961]K77
|--B. alpinus (Kaszab 1965)K77
|--B. costulatus (Fairmaire 1901)K77
|--B. dreuxi Pierre 1961K77
|--B. nepalicus (Kaszab 1973) [=Tagenoides nepalicus]K77
|--B. subcarinatus (Blair 1927)K77
`--B. zurstrasseni Kaszab 1977K77
Platolenes nepalensis Kaszab 1977K77
|--H. auriculatus Marseul 1876K77
`--H. pustuliferF89
Agnaptoria Reitter 1887K77
|--A. rubripes Reitter 1887K77
|--A. seidlitzi Reitter 1893K77
`--A. subcarinata Blair 1927K77
Platynoscelis (Leipopleura) politicollis Kaszab 1940K77
Hexarhopalus entomogonoides (Allard 1896) [=Ptilonix entomogonoides]K77
Zophosis Latreille 1802L02
|--*Z. testudinaria [=Erodius testudinarius]L02
|--Z. abbreviata Sol. 1834G89
|--Z. acuticosta Fairm. 1885G89
|--Z. plana [=Erodius planus]G89
|--Z. punctata Brullé 1832G89
|--Z. pygmaea Sol. 1834G89
|--Z. quadricostata Sol. 1834G89
|--Z. sabaea Baudi 1881G89
`--Z. sulcata Deyr. 1867G89
|--A. impressifrons Baudi 1881G89
`--A. rotundatus Sol. 1834G89
|--E. gibbusL02
`--E. puncticollis Sol. 1834G89
Rhytidonota praelonga (Reiche 1850) [=Axumia praelonga]G89
|--M. laevicollis Sol. 1835G89
|--M. puncticollis Sol. 1835G89
`--M. punctipennis Sol. 1835G89
Mesostenopa arabica Gestro 1881G89
Micipsa philistina Reiche 1857G89
|--O. hegeterica (Reiche 1857) [=Melanerus hegetericus]G89
`--O. pygmaea (Reiche 1857) [=Melanerus pygmaeus]G89
Sceleodis castaneus [=Cratopus castaneus]G89
Phaeotribon pulchellusG89
Himatismus villosus Haag 1870G89
Pogonobasis ornata Sol. 1837G89
Peristeptus gestroi Haag 1875G89
Stenosis costulata Baudi 1881G89
Eutagenia arabs (Baudi 1881) [=Stenosis arabs]G89
|--T. crinita (Klug 1830) [=Pimelia crinita]G89
`--T. murina Baudi 1881G89
Gedeon abyssinicusG89
|--S. orientale (Fabr. 1775) [=Opatrum orientale]G89
`--S. sulcatum Baudi 1876G89
|--H. famelicum Oliv. 1811G89
|--H. hispidum Brullé 1838G89
|--H. lugensG89
|--H. micansG89
|--H. murinum Baudi 1876G89
|--H. rusticum Oliv. 1811G89
`--H. sericeum Baudi 1876G89
Halonomus subplumbeus (Fairm. 1856) [=Platydema subplumbea]G89
Hopatroides punctulatusG89
Calcar raffrayi Fairm. 1873G89
Dilamus pictus Baudi 1881G89
|--M. gerstaeckeriG89
|--M. rugulosus Gestro 1878G89
`--M. tentyrioidesG89
|--H. elegans Baudi 1881G89
`--H. psalidiformis Ancey 1881G89
|--P. carbonariaW41
|--P. gagatina Mäklin 1864G89
|--P. marginata [=Helops marginatus]W41
`--P. rubripesW41
Boletoxenus bellicumK98
|--A. idunaC01
|--A. paradoxaC01
`--A. sulcipennis Reitt. in Csiki 1901C01
|--M. ciliataC01
|--M. heydeni Csiki 1901C01
|--M. maximaC01
`--M. mongolica Csiki 1901C01
Melaxumia angulosaC01
Scytosoma arcibasisC01
Microdera interruptaC01
Dordanea kraatziC01
|--T. kraatziC01
|--T. laeviusculaC01
|--T. reitteri Csiki 1901C01
|--T. seidlitziC01
`--T. zichyi Csiki 1901C01
Platyscelis rugifronsC01
|--D. barbarus Lucas 1847E12
|--D. extensusC01
|--D. rotundicollis Lucas 1847E12
|--D. subvariolosus Lucas 1847E12
`--D. tristisR26
|--S. carinatumC01
`--S. tuberculiferumC01
Ammophorus [Scotobiini]MB08
`--A. bifoveatusPP72
|--A. (Eurychora) ciliataG20
|--A. (Akis) collarisG20
|--A. punctataR26
`--A. reflexaL02
Tagenia Latreille 1802L02
|--*T. filiformis [=Akis filiformis]L02
`--T. algirica Lucas 1846E12
Scaurus striatusL02
Hegeter Latreille 1802L02
|--*H. elongatus [=Blaps elongata]L02
`--H. striatusG20
Orthocerus Latreille 1802L02
|--O. hirticornisR26
`--O. muticus [=Sarrotrium muticum]L02
‘Silvanus’ costatus Steinheil 1869 (n. d.)H73
|--B. laticollis Boisduval 1835 (n. d.)MB08
`--B. politus Boisduval 1835 (n. d.)MB08 [=Meneristes politusM86]
Chrysobalus fulgidipennis Boisduval 1835 (n. d.)MB08 [=Cyphaleus fulgidipennisM86]
Pseudostrongylium viridipenne Kraatz 1880 (n. d.)MB08
Tetraphyllus sumptuosus Hope 1843 (n. d.)MB08
|--B. egenus Pascoe 1866M86
|--B. macleayi Pascoe 1866M86
`--B. saccharatus Pascoe 1870M86
Ulodica hispida Pascoe 1869M86
|--E. crassicornis Pascoe 1860M86
`--E. lunatus Pascoe 1860M86
Ennebaeus ovalis Waterh. 1878M86
|--N. brevicornis Westwood 1842W42
|--N. confusus Westwood 1842W42
|--N. crenata Boisd. 1835M86
|--N. hypocrita Westwood 1842 [=Iphthinus hypocrita (n. n.); incl. I. guineensis (n. n.)]W42
|--N. moerens Westwood 1842W42
|--N. punctatus [=Helops punctatus]W42
`--N. rotundicollis Westwood 1842W42
Acrothymus caenosus Pascoe 1866M86
Misolampus hofmannseggiiG20
Hesiodobates antiquus Kaszab & Schawaller 1984P92
|--S. galapagoensisPP72
|--S. laevigatumPP72
`--S. rugosumPP72
Trientoma Sol. 1835FS90
|--T. guadeloupensis Fleutiaux & Sallé 1890FS90
`--T. salleiFS90
Diastolinus Muls & Rey 1859FS90
`--D. perforatus Sahlb. 1823FS90
Hesiodus Champ. 1885FS90
|--H. caraibus Fleutiaux & Sallé 1890FS90
`--H. longitarsisFS90
Allegoria Cast. 1840FS90
|--A. castelnaui Fleutiaux & Sallé 1890FS90
`--A. dilatataFS90
Antimachus Gistl 1829FS90
|--A. divergensFS90
`--A. roudeni Petit in Fleutiaux & Sallé 1890FS90
Cyrtosoma Perty 1830FS90
`--C. lherminieri Chv. 1838FS90
Elenophorus collarisR26

Alphitobius Stephens 1829MB08 [incl. Cryptops Solier 1851 non Leach 1815AT02, Heterophagus Redtenbacher 1845AT02, Microphyes Macleay 1872MB08]

Alphitobius laevigatus (Fabricius 1781) [=Opatrum laevigatum; incl. Helops picipes Panzer 1794, *Alphitobius picipes, *Microphyes rufipes Macleay 1872]MB08

Centronopus Solier 1848 [incl. Menechides Motschulsky 1872, Pachyurgus LeConte 1866, Pyres Champion 1885, Scotobates Horn 1875]AT02

Cheirodes Gené 1839MB08 [incl. Ammidaenemia Reitter 1904AT02, Anaemia Horn 1870AT02, Anemia Laporte 1840AT02, Anemiadena Ardoin 1971AT02, Eremonomus Wollaston 1861AT02, Histiaea Fairmaire 1892AT02, Pseudanemia Wollaston 1864 AT02, Spinianemia Ardoin 1971AT02, Trichanemia Ardoin 1971AT02; MelaniminiMB08]

Cryptoglossa Solier 1836 [incl. Amblycyphrus Motschulsky 1870, Centrioptera Mannerheim 1843, Oochila LeConte 1862]AT02

Opatrinus Dejean 1821 [incl. Hopatrinus Gemminger & Harold 1869, Zidalus Mulsant & Rey 1852, Zodinus Mulsant & Rey 1852]AT02

Tenebrionidae [Blapsidae, Cossyphores, Diaperidae, Helopidae, Petriidae, Pimeliariae, Pimeliidae, Rhysopaussidae, Tenebrionites, Tentyriidae, Ulominae]

*Type species of generic name indicated


[AT02] Aalbu, R. L., C. A. Triplehorn, J. M. Campbell, K. W. Brown, R. E. Somerby & D. B. Thomas. 2002. Tenebrionidae 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. 463–509. CRC Press.

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[ED04] Ewers, R. M., & R. K. Didham. 2004. A first assessment of New Zealand beetles restricted to large forest areas. Weta 27: 33–40.

[F89] Fairmaire, L. 1889. Coléoptères de l’intérieur de la Chine. 5e partie. Annales de la Société Entomologique de France, 6e série 9: 5–84.

[FS90] Fleutiaux, E., & A. Sallé. 1890. Liste des coléoptères de la Guadeloupe et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France, 6e série 9: 351–484.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 5–72.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[G91] Groves, C. P. 1991. A Theory of Human and Primate Evolution, revised ed. Clarendon Press: Oxford.

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