| Cleridae | |
|---|---|
 | |
| Some checkered beetles described in the mid-19th century | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Coleoptera |
| Suborder: | Polyphaga |
| Infraorder: | Cucujiformia |
| Superfamily: | Cleroidea |
| Family: | Cleridae Latreille, 1802 |
| Subfamilies | |
ClerinaeLatreille, 1802 | |
Cleridae are a family ofbeetles of the superfamilyCleroidea. They are commonly known ascheckered beetles. The family Cleridae has a worldwide distribution, and a variety ofhabitats and feeding preferences.
Cleridae have manyniches and feeding habits. Most genera arepredaceous and feed on other beetles and larvae; however other genera arescavengers orpollen feeders. Clerids have elongated bodies with bristly hairs, are usually bright colored, and have variableantennae. Checkered beetles range in length between 3 and 24 millimetres (0.12 and 0.94 in). Cleridae can be identified based on their 5–5–5tarsal formula, division ofsternites, and the absence of a special type of vesicle. Female Cleridae lay between28–42eggs at a time predominately under thebark of trees. Larvae are predaceous and feed vigorously before pupation and subsequently emergence as adults.
Clerids have a minor significance inforensic entomology. Some species are occasionally found on carrion in the later dry stages ofdecay. Also, some species are pests (stored product entomology) and are found infesting various food products. Research efforts related to Cleridae have focused primarily on using certain species as biological controls. This is a very effective technique for controllingbark beetles due to the voracious appetite of many clerid species.
Generally, checkered beetles are elongated andoval in shape and range from 3–24 millimetres (0.12–0.94 in) in length.[1] Their entire bodies are covered with bristly hairs and many display an ornate body color pattern.[1] These often brightly color patterns can be red, yellow, orange, or blue.[2] The antennae are clubbed at the tip for most species, but others can be "clubbed, saw-tooth, or thread-like."[1][2] The pronotum region is nearly cylindrical and characteristically narrower than theelytra (special hardened front wings), while the head is as wide or wider than thepronotum.[2] Their elytra have tiny pits or depressions, and never expose more than twotergites (dorsal plates).[1]
Clerid beetles fall under the suborderPolyphaga. Key characteristics of Polyphaga are that the hindcoxa (base of the leg), do not divide the first and second abdominal/ventral plates which are known assternites. Also, thenotopleural suture (found under the pronotal shield) is not present.[1] To further identify Clerid beetles, a few additional characteristics need to be examined.
Clerid beetles have unique legs that help to distinguish them from other families. Theirtarsal formula is 5–5–5, meaning that on each of the front, middle and hind legs there are 5 tarsomeres (individual subsegments of the feet/tarsi).[1] One or more of these subsegments on each leg is typically lobed, and the 4th tarsomere is normally difficult to distinguish. Furthermore, an important feature that eliminates many other families of beetles is that clerids' front coxae (base of the leg) expose the second segment of the legs known as the trochanter.[1]
The second defining characteristic of the family Cleridae is that clerids never have eversible vesicles (small usually hidden balloon-like structures thought to be scent glands) on theirabdomen andpronotum. This characteristic distinguishes them from a similar familyMelyridae which sometimes has these glands.[1] This trait is very important in correctly differentiating checkered beetles from Melyridae.
Cleridae can be found in the Americas, Africa, Europe, the Middle East and even in Australia. There are approximately 3,500 species in the world and about 500 species in North America.[3] Due to this wide distribution there are many differenthabitats in which the checkered beetles can be found.
Many of the species are known as "flower visitors", that prey on otherflower visitinginsects and also feed onpollen. These species are found in moist, sunny environments where flowering plants are found in abundance.[4]
Another habitat commonly inhabited by clerid beetles istrees. These "tree living species" are found in forests across the world with various climates and an array of easily preyed upon insects. They seek protection under the bark and hunt for other insects above and below thebark.[4] The primary source of prey for these bark living hunters is bark beetles.
The third type of clerid beetles is the "nest robbing species" which live in shrubbery and in trees. Unlike the tree living species, these species do not actuallyburrow into the bark. Nest robbing species typically hunt termite, bee, and wasp larvae, and one particular species has been noted to prey primarily on grasshopper egg masses.[4] Not all nest robbing species actively hunt live prey, some species for example prefer to feed only on dead honey bee larvae and adults.[4]
The Cleridae contains many species of predaceous beetles that feed on other beetles and beetle larvae in their natural habitat.[3] The most common prey item for checkered beetles are bark beetles and wood boring beetles.[5]
In general, the bulk of adult Cleridae feed mainly on other adult beetles while the larvae stage feed on other beetle larvae. Some checkered beetles are known to have an extremely voracious appetite with some larvae able to consume "several times their own body weight" in a day.[6]
Although most species of checkered beetles are predaceous in nature, some are scavengers and others have been found feeding on flower pollen.[7] Because of the checkered beetles predaceous nature and insatiable appetite, they are often key players in thebiological control of other insects. The checkered beetles have also developed a unique adaptation to aid in their quest for prey. The beetles usepheromones to help them locate, kill, and consume their prey.[8]
The diversity of checkered beetle's feeding habits is quite evident when different species are examined.Necrobiaspp. are attracted to dry carrion and other decomposing animal matter such as bones and skin as well as various meat products.[9]Thanasimus spp. are found inwoodland areas where bark beetle species constitute their main source ofprey.[9] The primary source of prey forPhyllobaenus spp. arewood borers, immatureweevils, andhymenoptera larvae.[9] One of the more diversegenera isTrichodes, in which larvae feed on the pollen of flowering plants and adults prey upon grasshoppers and wasps.[9]
The general life cycle of clerids has been known to last anywhere from 35 days to more than 3 years, and is strongly dependent on the life cycle of their prey.[10] While the life cycle can vary in length between genus and species, temperature is also a major determinant in the length of time spent in each stage of development. The warmer the temperature is, the quicker the life cycle, and the cooler the temperature is the slower the life cycle. If temperatures dip below a threshold temperature for an extended period of time, clerids and most other insects will have growth and developmental progress arrested. Like all beetles, Cleridae follow aholometabolous life cycle: the egg hatches into alarva, which grows and feeds, changing its skin to form apupa, and the pupa shedding its skin to emerge as an adult. The larvae of the majority of the known species of Cleridae feed upon the eggs and young ofwood-boring beetles, while the adults feed on the adult bark beetles.[11]
Copulation takes place while the female feeds, because females need a large amount of food for egg development.[12] The female lays her eggs 36–72 hours after copulation. The eggs are laid in between pieces of bark on wood-borer-infested trees or under stones in the soil.[11] She may lay 28–42 eggs at a time.[12] For the longer-living species such asThanasimus, this occurs in late summer or early fall to give thelarvae enough time for proper growth before having tooverwinter.[13]
When larvae hatch from their eggs, they are either red or yellow.[11] Their bodies have a slender and flat appearance with short legs due to their minimal movement. The larvae are covered in hair and have two horn-type projections on thedorsal area of the last body segment.[11] Immediately after birth, they start searching for food close to where they hatched. They feed on wood-borer insects on trees, or feed on their species' substrate or prey of choice.[11] Feeding is the main purpose of the larval stage to prepare for pupation. Once their larval stage is complete the tree dwelling species make their way to the bottom of the tree to pupate.[13] Thepupal stage can last from 6 weeks to one year depending on the need to overwinter, and how short the overall life cycle is for a particular species. A majority of clerid species pupate in earthen cells which are made from soil and certain enzymes secreted from their mouths.[11] The rest remain in pupal cells. Adult beetles emerge from pupation and spend a variable time of their life maturing, and eventuallyoviposit. Sexually mature adults orimagos ofThanasiumus overwinter inside the wood-borer-infested trees and oviposit during thespring.[12]
Necrobia rufipes, commonly known as the red-legged ham beetle, is of particular importance instored product entomology. It infestsdried orsmoked meats, especially those products that are stored unwrapped for long periods of time. Adults feed on the surface of the products, while the larvae damage the meat by boring down usually in the fatty parts.[14]Necrobia rufipes has been recorded feeding upon a large variety of items ranging from hides and dried figs to Egyptian mummies.[14] In addition, products such aswool andsilk can become infested, but not destroyed.[14]
Since clerids are predaceous in nature, they have been found feeding on fly larvae as well as the skin and bones ofcarrion.[9] Most clerids are not useful inforensics because of their food choice, but some species such asNecrobia rufipes can be useful.Necrobia rufipes is attracted towards carrion in the later stages of decomposition, so its arrival on carrion can help provide an estimate for thepost-mortem interval or PMI. Although the checkered beetle is not the most significant insect on carrion, the beetle's predaceous nature and its ability to reproduce in carrion that is exposed to the environment provides some forensic importance.[15]
There is ongoing research with some clerid species. Forensic research is limited because of their late arrival on carrion, but members such asThanasimus undatulus have been researched as a possible role inintegrated pest management or IPM.Thanasimus undatulus is a predator of bark beetles. Some species of bark beetles such as thesouthern pine beetle and themountain pine beetle can become pests to thelumber industry because in large numbers they can cause damage and kill live trees.Thanasimus undatulus has been researched as a possiblebiological control agent for these pests. Researchers and forestry officials have used bark beetle aggregation pheromones to attract the checkered beetle to specific trees. This causes the bark beetles to be overwhelmed, extensively preyed upon by the clerid beetles, and typically eliminated.[16]There is also additional research being done pertaining to the impact of clerids on pollination in flowers.[17]
Thegenera of Cleridae are divided among severalsubfamilies, though some genera still defy easy classification. Severaltaxonomic schemes exist, recognizing for example a group aroundNeorthopleura as distinct subfamilyNeorthopleurinae, or splitting off the Thaneroclerinae as distinct family, or circumscribing the Korynetinaesensu stricto orsensu lato. The following list of tribes and selected genera is thus preliminary. Some notable species are also listed. The oldest members of the family areProtoclerus andWangweiella the lateMiddle Jurassic (Callovian)Daohugou bed in Inner Mongolia, China.[18]
| Enopliinae (sometimes in Korynetinae)
Epiphloeinae (sometimes in Korynetinae)
Hydnocerinae (including Phyllobaeninae)
Tarsosteninae (sometimes in Korynetinae)
Thaneroclerinae (tentatively placed here)
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