Telephone-pole beetle

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Telephone-pole beetle
Temporal range: Miocene–Present
Micromalthus debilis.jpg
"Ghost adult" stage
Micromalthus debilis larva.png
Larval stage
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Family: Micromalthidae
Genus: Micromalthus
LeConte, 1878
Species:
M. debilis
Binomial name
Micromalthus debilis
LeConte, 1878
Other species
  • †Micromalthus eocenicusKirejtshuk, Nel & Collomb, 2010
  • †Micromalthus priabonicusPerkovsky, 2016
Synonyms [1]

Micromalthus anansiPerkovsky, 2008

The telephone-pole beetle (Micromalthus debilis) is a beetle native to the eastern United States and the only known living representative of the otherwise extinct family Micromalthidae. Larvae of the beetle live in decaying wood and can be pests to wooden structures, lending them their common name, the 'telephone-pole beetle.'

Contents

The larvae of Micromalthus debilis start as tiny white creatures with well-developed legs, resembling carabid larvae. Larvae bore into moist, decaying chestnut and oak logs, creating galleries as they consume wood fibers. Adult beetles are dark brown to blackish with vestigial reproductive organs. Mating behavior includes sex-role reversal, with females exhibiting more aggression and competition for mates. Micromalthus's evolutionary history dates back millions of years, with fossil records found in various ambers. Their larvae infest timber, weakening structures and attracting fungi, as seen in South African gold mines in the 1930s.

The telephone-pole beetle used to have reproducing adults, but has evolved to become obligately paedogenetic. The rare adult is sometimes referred to as a 'ghost adult' due to its vestigial existence. They have an unusual reproductive system involving asexually reproducing (parthenogenetic) female larvae. Being one of few parthenogenetic haplodiploid species, the telephone-pole beetle is an interesting subject of coleopteran reproductive behavior and physiology.

Distribution

Reports of the species are infrequent and it is unknown whether they are rare, or common and unrecognized. A recent study by Bertone et al. (2016) [2] found telephone-pole beetles in a survey of the indoor arthropod fauna in 50 houses located in and around Raleigh, North Carolina. [3] A recent survey found that the species had spread to every continent except Australia. With finds in South Africa, Hong Kong, Belize, [4] Cuba, Brazil, Japan, Hawaii, Italy and Austria, the dispersal is likely connected to the timber trade. [5]

Taxonomy

Classification of M. debilis was historically controversial and unsettled. The species, first reported by John Lawrence LeConte in 1878, was long considered one of the Polyphaga, and placed in the Lymexylidae or Telegeusidae, or as a family within the Cantharoidea. However, characteristics of larvae, wings, and male genitalia show that it is in the suborder Archostemata, where it has been placed since 1999. [6]

Description

Larvae

Female-producing (thelytokous) female larvae

The thelytokous female larva resembles a carabid larva when first hatched. The first instar larva is the smallest in length, measuring about 1.26mm. The body is white, distinct lateral bulges are present, and the legs are well-developed. [7] The second instar and subsequent instars have similar morphologies, with the head and body size and width increasing with each subsequent instar. The larvae measure between 2.7 to 3.3 millimeters in length. The head displays a light brown hue and notable sclerotization. In almost all segments, the cuticle covering the thorax and abdomen lacks pigmentation. The body is characterized by its slender, elongated form, maintaining parallel sides and cylindrical shape, featuring dorsal and ventral ampullae along with lateral bulges on numerous segments. The thorax is shorter than the combined length of abdominal segments I to III. [7] In later instars, the body is slightly flatter, broader, and shorter. [7] Legs are not present in second and older instars which is considered a trait unique to M. debilis. [7]

Male-producing (arrhenotokous) female larvae

The penultimate instar of the male-producing larva is vaguely described as having a dense white color due to much fat. The body is cylindrically shaped and marked by segmented constrictions. [8]

Male larvae

Unlike female larvae, which resemble the carabid type when first hatched, male larvae resemble that of a weevil. [9] The body is short and stubby with stump-like legs. [8]

Adults

The adult beetle is elongated and a dark brown to blackish color, with brownish-yellow legs and antennae. The head is larger than the thorax, with large eyes protruding from either side. [8] According to H.S. Barber, the adult female is responsible for

Feeding

The larvae are wood-borers that feed on moist and decaying chestnut and oak logs. Telephone-pole beetle larvae infest timber by burrowing into the wood, where they feed and develop. They tunnel through the timber, creating galleries as they consume the wood fibers.

Life cycle

Diagram of life cycle Micromalthus life cycle.webp
Diagram of life cycle

The life cycle of M. debilis is one of the most complicated life cycles among beetles because it involves viviparity, hypermetamorphosis, parthenogenesis, and paedogenesis. [8] The life cycle is also highly unusual because the larvae rarely mature into adults in both sexes. Haplodiploidy is another notable feature of this species; haploid males are hatched from eggs within the mother's body via haploid parthenogenesis, and diploid females are birthed live via diploid parthenogenesis. [10]

Females

Upon being birthed live, the larva emerges as a minute white creature with long, slender legs that resemble the carabid type. [9] This stage is primarily focused on migration, with the young larvae crawling away from the location of their birth to find new areas within their habitat. During this phase, they may consume the remains of their mother before migration. After a period of wandering, the larvae begin to burrow into wood or other suitable substrates again. They feed minimally during this time and undergo their first molt, transforming into a legless form that resembles the larva of cerambycid beetles. This stage is characterized by the development of an inconspicuous anal armature. Additional molts occur in this form, likely to accommodate head growth and overall body development. The larvae continue to bore through wood, packing their galleries with dust as they progress. The color of their bodies may darken due to the accumulation of food in their alimentary tract. As the larvae near maturity, the eggs in the ovaries of what will become the paedogenetic form become visible. At this stage, the larvae reverse their position in the gallery, construct a cell, and enter an aestivation phase. During aestivation, their bodies gradually turn white as they consume all available food in their system. [9]

Depending on circumstances, the larvae may either pupate (though this is described as rare) or undergo another molt, ultimately revealing the paedogenetic form, in which they can produce young. [11] Paedogenesis is the process by larvae reproduce by giving birth to more larvae without the production of adults and is a process exclusive to females. [11] Once the paedogenetic form emerges, it typically takes around two weeks for the new generation to be born. The young larvae are born tail-first and begin the cycle anew, continuing the species' life cycle. [9]

Males

Male larvae are hatched from a single, large egg that adheres to the male-producing larva's body for 8–10 days. The larvae feed on the mother's body and will grow rapidly. [9] The male rarely pupates and transforms into the imago, and will often die as a larva. [11] While male adults only develop from uni-oviparous paedogenetic females, adult females only develop from cerambycoid paedogenetic females. [9]

Larval sex ratio

In naturally occurring paedogenetic larvae, the sex ratio is strongly biased toward females. None of the three canonical explanations for biased sex ratios, local mate competition, local resource competition, and local resource enhancement, are likely explanations for the biased sex ratio in telephone-pole beetle larvae. Local mate competition selects for female-biased sex ratios when male siblings compete to fertilize their female siblings, but this is unlikely in this species which females tend to avoid mating with siblings. [11] Local resource competition selects for biased sex ratios but typically involves competition between females for resources and thus selects for male-biased ratios. [11] Lastly, local resource enhancement can select for biased sex ratios if the offspring of one sex increases the fitness of parents. [11] However, because female offspring feed on the mother, there is more likely competition between female larvae, contradicting this explanation. [11] As such, the cause of sex ratio deviation remains unclear. [11]

Adulthood

Adult telephone-pole beetles are unable to copulate, and adult females do not have the physiological mechanisms to reproduce because they are unable to lay eggs or produce live progeny, either sexually or by parthenogenesis. [11]

Since adults do not have a role in reproduction, they are not a physiological part of the life cycle. Thus, the rarity of adult development in the natural world may be an evolutionary response to the lack of their reproductive role. In laboratory settings, development into adults can be induced by high temperatures, but this also results in high mortality because only one out of hundreds of heat-treated larvae will survive and pupate into an adult. [11]

The adult females live for about six days and males only live for around 12 hours, with a strongly biased sex ratio towards females. The adults of both sexes are sterile and are vestigial remnants of a time when the life cycle involved sexual reproduction. The loss of sexual reproduction is likely associated with its infection by Wolbachia bacteria. [11]

Literature

Limited observations and experiments on M. debilis have resulted in conflicting observational conclusions, particularly regarding the beetle's reproduction, in the existing literature. Pollock & Normack reported the existence of reproductive adult males, but this was based on the incorrect conclusions by Barber. [9] [11] However, all existing experimental literature states that adults are fertile. [11]

Mating behavior

A 2016 experimental study used heating to generate substantial numbers of adults to simulate the now non-functional adult reproductive behavior. Due to parthenogenetic reproduction in telephone-pole beetles, information on the sexual mating system of this beetle was previously lost, but recent research is working to uncover this information. Both adult female and adult male modern telephone-pole beetles are sterile, but they still exhibit mating behavior. [11]

Sex role reversal

The study revealed sex-role reversal, meaning that females face more competition for mates compared to males. This was demonstrated by greater initiative by females to mate and increased fighting between females in the presence of unrelated males. Female also display more aggressive mating behavior, as they may grasp the male genitalia with their own genitalia. [11]

Rejection behavior

Adult female beetles exhibit rejection behavior to avoid inbreeding with related males, which are adult male beetles that are located on the same natal log. Right after pupating, males expose their reproductive organs as a mate-seeking gesture. Despite tending to compete for male mates, females will ignore these males because they originate from the same log. The males would need to take a short flight to a neighboring log for female mates. Females also perform a 'kin dance' involving shaking of their abdomens and beating of their wing, which is thought to be a deterring signal to related males. Such rejection behaviors demonstrate that sex roles are not fixed because females can also display choosy behavior. The rejection behavior also supports that the female-biased sex ratio is not due to local mate competition. [11]

Mounting

Females initiated more interactions by actively mounting males, further supporting that female compete for male mates. In cases where several females pile on top of a male, a female may try to dislodge the others with her mandible. Female-female mounting can also occur, and the frequency of this does not change depending on the presence or absence of males. [11]

Evolutionary history

Genetic studies have placed Micromalthus as more closely related to Ommatidae than to Cupedidae within Archostemata. [12] A close relationship between Ommatidae and Micromalthidae is supported by several morphological characters, including those of the mandibles and male genitalia. [13] A close relationship to the enigmatic family Crowsoniellidae has been suggested by some sources. [14] The oldest record of Micromalthidae is Archaeomalthus from the Upper Permian of Russia around 252 million years old, which is morphologically similar in many respects to Micromalthus including an only weakly sclerotised body. [15] Several other fossil genera of the family are known including Cretomalthus , known from a larva found in Early Cretaceous (Barremian) Lebanese amber, [16] as well as Protomalthus from the mid-Cretaceous (Albian-Cenomanian) Burmese amber of Myanmar. [17]

Fossils of Micromalthus are known from the Miocene aged Dominican amber (adults and larvae, which were found to not be distinguishable from the living species [1] ) and Mexican amber (larvae), the late Eocene aged Rovno amber of Ukraine (Micromalthus priabonicus), [18] and the early Eocene (Ypresian) aged Oise amber of France (Micromalthus eocenicus). [19] A possible specimen of Micromalthus is known from Burmese amber, [20] but the poor preservation of the specimen makes the assignment tentative. [17]

Wood pests

Telephone-pole beetle larvae infest timber by burrowing into the wood, where they feed and develop. They tunnel through the timber, creating galleries as they consume the wood fibers. This activity weakens the structural integrity of the timber, leading to decay and potential collapse. Additionally, their presence can attract fungi, further contributing to the degradation of the timber. [21]

In the 1930s, telephone-pole beetle larvae were reported as the perpetrators of a gold mine infestation in the Witwatersrand Basin in South Africa. The primary timbers employed in the mines, including Acacia , Eucalyptus , and Pinus species, are all susceptible to infestation, particularly in conditions with ample moisture. Heavy infestations were noted in aged timbers within poorly ventilated shafts, especially in environments with temperatures ranging from 88 to 93 °F. The presence of stagnant or running water may cause even further decay, sometimes resulting in complete pulverization. [22]

Related Research Articles

<span class="mw-page-title-main">Micromalthidae</span> Family of beetles

Micromalthidae is an ancient family of small beetles belonging to the suborder Archostemata. The only known living representative is the telephone-pole beetle, Micromalthus debilis. A few extinct species have been described, the oldest being the Late Permian Archaeomalthus synoriacos. It is one of the oldest families of beetles still existing today.

<span class="mw-page-title-main">Mealworm</span> Species of darkling beetle, larval form

Mealworms are the larval form of the yellow mealworm beetle, Tenebrio molitor, a species of darkling beetle.

<span class="mw-page-title-main">Archostemata</span> Suborder of beetles

The Archostemata are the smallest suborder of beetles, consisting of 50 living species in five families and over 200 described fossil species. They are an ancient lineage with a number of primitive characteristics. Antennae may be thread-shaped (filiform) or like a string of beads (moniliform). This suborder also contains the only beetles where both sexes are paedogenic, Micromalthus debilis. Modern archostematan beetles are considered rare, but were more diverse during the Mesozoic.

<span class="mw-page-title-main">Hydrophilidae</span> Family of beetles

Hydrophilidae, also known colloquially as water scavenger beetles, is a family of beetles. Aquatic hydrophilids are notable for their long maxillary palps, which are longer than their antennae. Several of the former subfamilies of Hydrophilidae have recently been removed and elevated to family rank; Epimetopidae, Georissidae, Helophoridae, Hydrochidae, and Spercheidae. While the majority of hydrophilids are aquatic, around a third of described species are terrestrial, mostly belonging to the subfamily Sphaeridiinae.

<span class="mw-page-title-main">Lymexylidae</span> Family of wood-boring beetles

The Lymexylidae, also known as ship-timber beetles, are a family of wood-boring beetles. Lymexylidae belong to the suborder Polyphaga and are the sole member of the superfamily Lymexyloidea.

<span class="mw-page-title-main">Japanese rhinoceros beetle</span> Species of beetle

The Japanese rhinoceros beetle, also known as the Japanese rhino beetle, the Japanese horned beetle, or by its Japanese name kabutomushi, is a species of rhinoceros beetle. They are commonly found in continental Asia in countries such as China, the Korean peninsula, Japan, and Taiwan. In these areas, this species of beetle is often found in broad-leaved forests with tropical or sub-tropical climates. This beetle is well known for the prominent cephalic horn found on males. Male Japanese rhinoceros beetles will use this horn to fight other males for territory and access to female mating partners. Upon contact, males will attempt to flip each other onto their backs or off of their feeding tree. In response to selective pressures, smaller male A. dichotoma have adapted a "sneak-like behavior". These smaller beetles will attempt to avoid physical confrontation with larger males and try to mate with females.

<span class="mw-page-title-main">Ripiphoridae</span> Family of beetles

Ripiphoridae is a cosmopolitan family of some 450 described species of beetles sometimes called "wedge-shaped beetles". Ripiphoridae are unusual among beetle families in that many species are hypermetamorphic parasitoids, an attribute that they share with the Meloidae. Members of the family differ in their choice of hosts, but most attack various species of bees or wasps, while some others attack cockroaches or beetles. Many species of Ripiphoridae have abbreviated elytra, and flabellate or pectinate antennae.

<i>Lampyris noctiluca</i> Species of beetle

Lampyris noctiluca, the common glow-worm of Europe, is the type species of beetle in the genus Lampyris and the family Lampyridae.

<span class="mw-page-title-main">American carrion beetle</span> Species of beetle

The American carrion beetle is a North American beetle of the family Silphidae. It lays its eggs in, and its larvae consume, raw flesh and fungi. The larvae and adults also consume fly larvae and the larvae of other carrion beetles that compete for the same food sources as their larvae. They prefer to live in marshy and woody habitats. Necrophila americana emerge from their larval state in the early summer. The P. ashtoni cuckoo bumble bee displays close mimicry with the American carrion beetle. They are important in forensic studies because of their tendency to thrive on large carcasses.

<i>Crowsoniella</i> Family of beetles

Crowsoniella is a genus of beetles in the order Archostemata. It contains only a single species, Crowsoniella relicta, and is the only member of the monotypic family Crowsoniellidae. It is known only from three male specimens collected in 1973 in the Lepini mountains of central Italy by Roberto Pace. In a degraded pasture, the beetles were found among the roots of a large hawthorn tree, in deep calcareous soil. No other specimens have been found since.

<span class="mw-page-title-main">Ommatidae</span> Family of beetles

The Ommatidae are a family of beetles in the suborder Archostemata. The Ommatidae are considered the extant beetle family that has most ancestral characteristics. There are only seven extant species, confined to Australia and South America. However, the geographical distribution was much wider during the Mesozoic spanning across Eurasia and Australia, suggesting that they were widespread on Pangea. So far, over 26 extinct genera containing over 170 species of these beetles have been described. Three extant genera have been assigned to this family: Omma,Tetraphalerus and Beutelius. The family is considered to be a subfamily of Cupedidae by some authors, but have been found to be more closely related to Micromalthidae in molecular phylogenies. A close relationship with Micromalthidae is supported by several morphological characters, including those of the mandibles and male genitalia. Due to their rarity, their ecology is obscure, it is likely that their larvae feed on deadwood.

<i>Gastrophysa viridula</i> Species of beetle

Gastrophysa viridula, known as the green dock beetle, green dock leaf beetle or green sorrel beetle, is a species of beetle native to Europe.

<i>Austroplatypus incompertus</i> Species of beetle

Austroplatypus incompertus, a type of ambrosia beetle, is endemic to Australia. They are found in mesic forests, and subtropical and tropical ecosystems along the east coast of Australia. There are many unique characteristics attributable to the A. incompertus, like their gallery excavation in several Eucalyptus species, their obligate eusocial behavior, their relationship with fungi, and their unique sexual dimorphism. These beetles are one of the only insects that display obligate eusocial behavior. Additionally, their sexually dimorphic traits are of interest, since body size is reversed with males having smaller torsos than female A. incompertus beetles.

<span class="mw-page-title-main">Madrone butterfly</span> Species of butterfly

Eucheira socialis, commonly known as the madrone butterfly is a lepidopteran that belongs to the family Pieridae. It was first described by John O. Westwood in 1834. Locally known as Mariposa del madroño or tzauhquiocuilin, it is endemic to the highlands of Mexico, and exclusively relies on the madrone as a host-plant. The species is of considerable interest to lepidopterists due to gregarious nest-building in the larval stages, and heavily male-biased sex ratio. It takes an entire year for this adult butterfly to develop from an egg. The eggs are laid in the month of June and the adults emerge the following May–June. The adults have a black and white pattern on their wings, and the males are generally much smaller and paler than the females. The larvae do not undergo diapause and continue to feed and grow communally in the coldest months of the year. There are two subspecies of E. socialis, named E. socialis socialis and E. socialis westwoodi.

<i>Callosobruchus chinensis</i> Common species of beetle

Callosobruchus chinensis, also known as the adzuki bean weevil, pulse beetle, Chinese bruchid or cowpea bruchid, is a common species of beetle found in the bean weevil subfamily. Although it is commonly known as the "adzuki bean weevil" it is in fact not a true weevil, belonging instead to the leaf beetle family, Chrysomelidae.

<i>Calleida viridipennis</i> Species of ground beetle

The Calleida viridipennis is a species of ground beetle belonging to the Carabidae family, and are referred to as carabid beetles. It is found in various states, including New York, New Mexico, Louisiana, and Florida. Habitat preferences include forests and swamps. C. virdipennis is a particular carabid beetle that is, on average, 10 mm long. Recognizable by a green-black metallic exoskeleton, it has a trapezoidal head shape. Its large eyes are also characteristic of the species. Below the exoskeleton reside functional wings, giving the beetle flight capacity.

<i>Xyleborinus saxesenii</i> Species of beetle

Xyleborinus saxesenii, commonly known as the fruit-tree pinhole borer, is a species of ambrosia beetle in the family Curculionidae. It is native to the Palaearctic region but has been introduced in many locations, including North America. X. saxesenii typically live in freshly dead wood, but it has also been reported to attack live trees. Such attacks on live trees may lead to economic damage.

<i>Rivacindela hudsoni</i> Species of beetle

Rivacindela hudsoni is an Australian species of the family Cicindelinae or "tiger beetle" and is the fastest running insect. The genus Rivacindela is contentiously treated as a subgenus of the broader Cicindela and are typically found in salty habitats such as dry salt lakes and salt streams. The species was discovered in South Australia and described in 1997, with an adult form of approximately 20–21mm in length and a running speed of 2.49 m/s, or 120 body lengths per second.

<i>Gnatocerus cornutus</i> Species of beetle

Gnatocerus cornutus, or the broadhorned flour beetle, is a common species of beetle in the Tenebrionidae family. It is distributed worldwide and is commonly found in warm, tropical regions, preferring grains, yeasts, and flours as its main diet. The total development time is about 47 days. G. cornutus is predominately found in mills as a storage pest and has a wide distribution.

Carabus japonicus is a ground beetle endemic to Japan. It inhabits the southwestern Japanese forests and is most frequently observed between May and September, with peak observations occurring mid-August. C. japonicus is a univoltine species, meaning it has one brood of offspring per year. Its larvae prey exclusively on earthworms. At lower altitudes, the reproductive cycle has been shown to coincide with the abundance of earthworms. This demonstrates the synchronization between the life cycles of the two species. The males tend to be slightly smaller than the females with both sexes having median body sizes of 24 mm and 26 mm, respectively. However, there is a large variation in intraspecific body size due to factors other than sex. These factors include parent size, food availability, and habitat temperature.

References

  1. 1 2 Hörnschemeyer, Thomas; Wedmann, Sonja; Poinar, George (February 2010). "How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera)". Zoological Journal of the Linnean Society. 158 (2): 300–311. doi: 10.1111/j.1096-3642.2009.00549.x . (Erratum:  doi:10.1111/j.1096-3642.2010.00702.x)
  2. ↑ Bertone, MA; Leong, M; Bayless, KM; Malow, TL; Dunn, RR; Trautwein, MD (19 January 2016). "Arthropods of the great indoors: characterizing diversity inside urban and suburban homes". PeerJ. 4: e1582. doi: 10.7717/peerj.1582 . PMC   4727974 . PMID   26819844.
  3. ↑ Milman, Oliver (19 January 2016). "Hundreds of tiny spiders, lice and more crawling through US homes, study says". The Guardian . Retrieved 19 January 2016. Matthew Bertone, an entomologist at North Carolina State University, said he was amazed at the variety of species found in what he stressed were 'clean and normal' homes in Raleigh, North Carolina. 'We were pretty surprised with what we found, such as the smallest wasp in the world, which is just 1mm long,' he said. 'I saw a lot of things in homes that I had never seen in the wild before, things we've previously tried to trap. There is a weird species of beetle, called telephone pole beetles, where the babies can produce babies. And tiny crickets called ant-loving ants because they are found near ant nests. I've never seen one of those before.'
  4. ↑ Philips, T. Keith (2001). "A Record of Micromalthus debilis (Coleoptera: Micromalthidae) from Central America and a Discussion of Its Distribution". The Florida Entomologist. 84 (1): 159–160. doi:10.2307/3496680. ISSN   0015-4040. JSTOR   3496680.
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  6. ↑ Ross H. Arnett, Jr. and Michael C. Thomas, American Beetles (CRC Press, 2001), chap. 2
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  12. ↑ McKenna, Duane D.; Shin, Seunggwan; Ahrens, Dirk; Balke, Michael; Beza-Beza, Cristian; Clarke, Dave J.; Donath, Alexander; Escalona, Hermes E.; Friedrich, Frank; Letsch, Harald; Liu, Shanlin (2019-12-03). "The evolution and genomic basis of beetle diversity". Proceedings of the National Academy of Sciences. 116 (49): 24729–24737. Bibcode:2019PNAS..11624729M. doi: 10.1073/pnas.1909655116 . ISSN   0027-8424. PMC   6900523 . PMID   31740605.
  13. ↑ Hörnschemeyer, Thomas (July 2009). "The species-level phylogeny of archostematan beetles-where do Micromalthus debilis and Crowsoniella relicta belong?". Systematic Entomology. 34 (3): 533–558. Bibcode:2009SysEn..34..533H. doi:10.1111/j.1365-3113.2009.00476.x. S2CID   84795808.
  14. ↑ Li, Yan-Da; Tihelka, Erik; Yamamoto, Shûhei; Newton, Alfred F.; Xia, Fang-Yuan; Liu, Ye; Huang, Di-Ying; Cai, Chen-Yang (2023-08-22). "Mesozoic Notocupes revealed as the sister group of Cupedidae (Coleoptera: Archostemata)". Frontiers in Ecology and Evolution. 11. doi: 10.3389/fevo.2023.1015627 . ISSN   2296-701X.
  15. ↑ Yan, Evgeny Viktorovich; Beutel, Rolf Georg; Lawrence, John Francis; Yavorskaya, Margarita Igorevna; Hörnschemeyer, Thomas; Pohl, Hans; Vassilenko, Dmitry Vladimirovich; Bashkuev, Alexey Semenovich; Ponomarenko, Alexander Georgievich (2020-09-13). "Archaeomalthus -(Coleoptera, Archostemata) a 'ghost adult' of Micromalthidae from Upper Permian deposits of Siberia?". Historical Biology. 32 (8): 1019–1027. Bibcode:2020HBio...32.1019Y. doi:10.1080/08912963.2018.1561672. ISSN   0891-2963. S2CID   91721262.
  16. ↑ Song, Zhenyu; Zhao, Xianye; Jarzembowski, Edmund A.; Xiao, Chuantao (2022-09-01). "Overview of and taphonomic bias in Archostemata (Insecta: Coleoptera) from Cretaceous amber". Palaeontographica Abteilung A. 323 (4–6): 119–137. Bibcode:2022PalAA.323..119S. doi:10.1127/pala/2022/0131. ISSN   0375-0442. S2CID   250394641.
  17. 1 2 Tihelka, Erik; Huang, Diying; Cai, Chenyang (March 2020). "A new genus and species of Micromalthidae from Burmese amber (Coleoptera: Archostemata)". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 111 (1): 39–46. Bibcode:2020EESTR.111...39T. doi:10.1017/S1755691019000185. ISSN   1755-6910. S2CID   210267382.
  18. ↑ Perkovsky, E. E. (May 2016). "A new species of Micromalthidae (Coleoptera) from the Rovno amber: 1. Adult morphology". Paleontological Journal. 50 (3): 293–296. Bibcode:2016PalJ...50..293P. doi:10.1134/S0031030116030047. ISSN   0031-0301. S2CID   89095000.
  19. ↑ Kirejtshuk, Alexander G.; Nel, André; Collomb, François-Marie (January 2010). "New Archostemata (Insecta: Coleoptera) from the French Paleocene and Early Eocene, with a note on the composition of the suborder". Annales de la Société Entomologique de France. N.S. 46 (1–2): 216–227. doi:10.1080/00379271.2010.10697661. ISSN   0037-9271. S2CID   55400656.
  20. ↑ Yamamoto, Shûhei (2021-07-03). "The extant telephone-pole beetle genus Micromalthus discovered in mid-Cretaceous amber from northern Myanmar (Coleoptera: Archostemata: Micromalthidae)". Historical Biology. 33 (7): 941–948. Bibcode:2021HBio...33..941Y. doi:10.1080/08912963.2019.1670174. ISSN   0891-2963. S2CID   208560568.
  21. ↑ Normark, Benjamin B. (2013). "Micromalthus debilis". Current Biology. 23 (10): R430–R431. Bibcode:2013CBio...23.R430N. doi: 10.1016/j.cub.2013.02.045 . ISSN   0960-9822. PMID   23701680.
  22. ↑ Pringle, J. A. (September 27, 1938). "A Contribution to the Knowledge of Micromalthus debilis LeC. (Coleoptera)". Transactions of the Royal Entomological Society of London. 87 (12): 271–286. doi:10.1111/j.1365-2311.1938.tb00088.x – via CABI Digital Library.
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