Exemplar of the Cymatodera aegra complex, copyright Mike Quinn.

Belongs within: Cleroidea.
Contains: Lemidia, Stigmatium, Eleale, Hydnocerinae, Enopliinae, Epiphloeinae, Clerinae, Necrobia.

The Cleridae, checkered beetles, are a family of elongate, hairy beetles, some if which are brightly coloured. Most clerids are predatory on other insects both as adults and larvae. Members of the family have five-segmented tarsi but the fourth tarsomere is concealed within the third in the Korynetinae, Tarsosteninae, Epiphloeinae and Enopliinae. The antennal club is loose and longer than the remainder of the flagellum in Epiphloeinae and Enopliinae but compact and not so long in Korynetinae and Tarsosteninae. Korynetinae have a broad-oval habitus and entire carina along the lateral margin of the pronotum whereas Tarsosteninae have a narrow-rectangular habitus and the lateral pronotal carina is not entire.

The Tillinae are distinguished by having the fore coxal cavities closed behind whereas they are open in other subfamilies. The Californian species Bostrichoclerus bicornis has a prominent pair of horns on the frons in front of the eyes. Another North American genus Monophylla is distinguished by having the last antennal segment flattened and much increased in length (Opitz 2002).

Today’s Tillinae
Published 9 May 2021
Tillus elongatus, copyright Gilles San Martin.

The above individual is a representative of a species of the subfamily Tillinae of the beetle family Cleridae. Clerids are a widespread group of moderate-sized beetles, larger individuals being about a centimetre in length, but most species tend to attract little attention from humans. They are mostly predators in confined spaces (Gunter et al. 2013): larvae hunt down wood-boring insects in their burrows, or the young of bees and wasps in their nests, whereas adults hunt for other insects under bark. Adults are more or less elongate in shape and commonly have an even covering of setae and a prominently punctate dorsum. The legs have five-segmented tarsi, each tarsus often with multiple segments lobed. Clerids are commonly referred to as ‘checkered beetles’ in reference to the contrasting colour patterns of many species but, as you can clearly see, not all clerids have such checkered patterns.

The subfamilial classification of clerids has shifted around a lot in the past but the Tillinae have been one of the more consistently recognised subfamilies. The feature most consistently separating tillines from other clerids is that the fore coxal cavities are both externally and internally closed: that is, the external rim and internal collar around the fore coxae are both complete rather than being interrupted posteriorly. Other distinctive features of the tillines are that all five tarsal segments are well developed and distinct, the pronotum is campanulate (bell-shaped) or bisinuate, and the eyes usually have coarse ommatidia (Burke & Zolnerowich 2017).

Cylidrus megacephalus, copyright Udo Schmidt.

The taxonomy of tillines is (as always) in great need of study. Around 550 or more species have been assigned to the subfamily worldwide, with the highest diversity in the Afrotropical and Oriental regions. However, many species can be quite variable in appearance and it is suspected that many previously described species may turn out to be synonyms. The situation is not helped by many species being rarely collected. For instance, Bostrichoclerus bicornis is a remarkable species distinguished by the presence of a pair of prominent, apically bifurcate horns arising alongside the antennal insertions. To date, this species is known from just two specimens, collected at separate locations in Baja California and southern California (Burke & Zolnerowich 2017).

A molecular phylogenetic analysis of the Cleridae by Gunter et al. (2013) suggested that the Tillinae represent the sister group of all other clerids. While unexpected from a morphological perspective, this result does tally with the long recognition of the tillines as a distinctive group. They may prove to have interesting things to tell us about the evolution of the clerids as a whole.

Systematics of Cleridae

Characters (from Lawrence & Britton 1991): Elongate, parallel-sided, clothed with erect, or erect and decumbent, hairs, often either metallic in colour or patterned with red or yellow. Head usually more or less deflexed; eyes usually slightly to strongly emarginate; antennae short; labial palps often enlarged and securiform; pronotum usually without lateral carinae; prosternum well developed in front of coxae; tarsi with one or more segments lobed or with membranous appendages; first tarsal segment sometimes reduced and concealed within tibial apex. Larvae elongate, cylindrical to somewhat flattened, lightly sclerotised except for head, one or more tergal plates on thorax and T9 but cuticle often pigmented with yellow, pink, blue or brown. Head well sclerotised dorsally and ventrally; gula long and narrow; mandible with hyaline process at base; epicranial stem usually absent, median endocarina usually present.

<==Cleridae [Clerii, Corynetidae, Korynetidae]
|--Tillinae [Tillides]RS15
| | i. s.: ‘Clerus’ sanguinalis Westw. 1852 [=Lissaulicus sanguinalis; incl. Macrotelus sanguineus]G01
| |--Aphelochroa carneipennisG01
| |--Bostrichoclerus Van Dyke 1938O02
| | `--*B. bicornis Van Dyke 1938O02
| |--Araeodontia Barr 1952O02
| | `--*A. penninsularis Schaeffer 1904O02
| |--Cymatoderella Barr 1962O02
| | `--*C. collaris (Spinola 1844) [=Tillus collaris]O02
| |--Callotillus Wolcott 1911O02
| | |--*C. eburneocinctus Wolcott 1911O02
| | `--C. elegansO02
| |--CylidrusLB91
| | |--C. basalis Macleay 1872M86
| | |--C. centralis Pascoe 1860M86
| | `--C. nigrinus White 1849M86
| |--Graptoclerus Gorham 1901G01
| | |--*G. signatus Gorham 1901G01
| | |--G. equestris [=Clerus equestris]G01
| | `--G. quadripunctatus Gorham 1901G01
| |--Prosymnus Laporte de Castelnau 1836 [incl. Ryparus Spin. 1844]G01
| | |--P. cribripennisG01
| | |--‘Ryparus’ tomentosusG01
| | `--P. villosus Gorham 1901G01
| |--OpetiopalpusG01
| | |--O. caffrarius Gorham 1901G01
| | |--O. collarisG00
| | `--O. rubricollis Klug 1837–1840G00
| |--Monophylla Spinola 1841 [incl. Elasmocerus LeConte 1849, Macrotelus Klug 1842]O02
| | |--*M. megatoma Spinola 1841O02
| | |--M. californicaO02
| | `--M. terminataO02 [=Macrotelus terminatusG00]
| |--Bogcia Barr 1978O02
| | |--*B. disjuncta Barr 1978O02
| | |--B. oaxacaeO02
| | `--B. obliquefasciataO02
| |--Lecontella Wolcott & Chapin 1918O02
| | |--*L. cancellata (LeConte 1854) [=Cymatodera cancellata]O02
| | |--L. brunneaO02
| | `--L. gnaraO02
| |--TillusM86
| | |--T. dux Westw. 1852M86
| | |--T. elongatusL02
| | |--T. formicariusL02
| | |--T. hilarisM86
| | `--T. unifasciatusR26
| `--Cymatodera Grey 1832O02
| |--*C. hopei Grey 1832O02
| |--C. aegraO02
| |--C. belfrageiB14
| |--C. bicolorO02
| |--C. californicaMW15
| |--C. horniO02
| |--C. oblitaO02
| |--C. ovipennisO02
| `--C. tricolor Skinner 1905B14
| |--L. rufipes (Klug 1842) [=Enoplium rufipes]B14
| `--L. zonataB14
|--Lebasiella Spinola 1844O02
| |--*L. erythrodera Spinola 1844O02
| `--L. pallipesO02
|--Loedelia Lucas 1920 [incl. Necrobioides Gahan 1919 non Fairmaire 1882]O02
| |--*L. mexicana (Gahan 1910) [=Necrobiodes mexicanus]O02
| `--L. maculicollisO02
|--C. abdominalisG00
|--C. analis Klug 1842 [incl. C. pallicornis]G00
|--C. concolor Gorham 1900G00
|--C. rufipes [=Anobium rufipes; incl. C. australis]B35
`--C. unicolor Chev. 1878M86

Cleridae incertae sedis:
Phloiocopus tricolorG89
Tarsobaenus letourneauaeDL99
Phlogistus eximiusB70
|--E. maculatumMC94
`--E. modestumMC94
|--A. brevipennis Pascoe 1860M86
`--A. ctenostomoides Waterh. 1836M86
|--S. albocinctus Pascoe 1860M86
|--S. idoneus Newm. 1842M86
`--S. splendidus Newm. 1840 [incl. S. reichei]M86
|--Paratillus Gorham 1876O02
| |--*P. carus (Newman 1840) [=Clerus carus]O02
| `--P. basalis Gorham 1878M86
`--Tarsostenus Spinola 1844O02
|--*T. univittatus (Rossi 1792) [=Clerus univittatus]O02
|--T. mastersi Macleay 1872M86
|--T. pulcher Macleay 1872M86
`--T. zonatusM86
|--O. olivaceus Westw. 1852M86
`--O. prasinus Westw. 1852M86
|--N. cribricollisM86
|--N. mastersi Macleay 1872M86
|--N. porcata [incl. N. heros Sturm 1843]M86
`--N. titana Thoms. 1860M86
Orthrius cylindricus Gorham 1876M86
Cleromorpha novenguttatus (Westw. 1852) [=Clerus novemguttatus]M86
|--O. australisM86
|--O. cruentatus Chev. 1874M86
`--O. gracilis Gorham 1876M86
Eburiphora patriciaM86
|--T. abbreviatus White 1849M86
|--T. ruficollis Macleay 1872M86
`--T. telephroides Pascoe 1860M86
|--P. anthicoides Newm. 1842M86
|--P. bicinctus Newm. 1842M86
|--P. fatuus [incl. P. ochropus, P. passerinii]M86
`--P. pallipes Macleay 1872M86
Tanocleria Hong 2002H02
`--*T. tenuis Hong 2002H02
Lissaulicus laevisG01
|--P. compressicornis (Klug 1842) [=Tillus compressicornis; incl. T. uniformis, Macrotelus uniformis]G00
`--P. pretiosus Gorh. 1883G00
|--P. flavonotatusG00
`--P. undulatus Gorham 1900G00
Dolichopsis Gorh. 1877G00
|--D. auronitens Gorh. 1900G01
`--D. haplocnemodesG00

*Type species of generic name indicated


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