Hkamti Amber
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Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
. The amber is dated to around 100 million years ago, during the latest Albian to earliest
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
ages of the mid- Cretaceous period. The amber is of significant palaeontological interest due to the diversity of flora and fauna contained as inclusions, particularly arthropods including insects and arachnids but also birds, lizards, snakes, frogs and fragmentary dinosaur remains. The amber has been known and commercially exploited since the first century AD, and has been known to science since the mid-nineteenth century. Research on the deposit has attracted controversy due to its alleged role in funding internal conflict in Myanmar and hazardous working conditions in the mines where it is collected.


Geological context, depositional environment and age

The amber is found within the Hukawng Basin, a large Cretaceous-
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
sedimentary basin within northern Myanmar. The strata have undergone folding and faulting. The Hukawng basin is part of the larger Myanmar Central Basin, a N-S orientated synclinal basin extending to the Gulf of Martaban to the south. The basin is considered to be a part of the West Burma Block or Burma Terrane, which has a debated tectonic history, it is considered to be associated with the concepts of the Cimmeria and Sibumasu terranes. The block was part of
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
during at least the Early Paleozoic, but the timing of rifting is very uncertain, with estimates ranging from the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
to Early Cretaceous. It is also disputed whether the block had accreted onto the Asian continental margin by the time of the amber deposition. Some members of the flora and fauna have Gondwanan affinities, while others have Laurasian affinities. A recent paleomagnetic reconstruction finds that the Burma Terrane formed an island land mass in the Tethys Ocean during the Mid Cretaceous at a latitude around 5-10 degrees south of the equator. At Noije Bum, located on a ridge, amber is found within fine grained
clastic rock Clastic rocks are composed of fragments, or clasts, of pre-existing minerals and rock. A clast is a fragment of geological detritus,Essentials of Geology, 3rd Ed, Stephen Marshak, p. G-3 chunks, and smaller grains of rock broken off other rocks ...
s, typically medium to greyish green in colour, resulting from the constituent grains being black, yellow, grey and light green. The fine grained rocks are primarily fine to very fine grained sandstone, with beds of silt and
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
and laterally persistent thin (1–2 mm thick) coal horizons. Massive micritic limestone interbeds of 6-8 centimetre thickness, often containing coalified plant material also occur. This
facies In geology, a facies ( , ; same pronunciation and spelling in the plural) is a body of rock with specified characteristics, which can be any observable attribute of rocks (such as their overall appearance, composition, or condition of formatio ...
association is typically around 1 metre thick and typically thinly bedded and laminated. Associated with the fine grained facies is a set of medium facies primarily consisting of medium to fine grained sandstones also containing thin beds of siltstone, shale and
conglomerate Conglomerate or conglomeration may refer to: * Conglomerate (company) * Conglomerate (geology) * Conglomerate (mathematics) In popular culture: * The Conglomerate (American group), a production crew and musical group founded by Busta Rhymes ** Co ...
, alongside a persistent conglomerate horizon. A specimen of the
ammonite Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish) ...
''
Mortoniceras ''Mortoniceras'' is an ammonoid genus belonging to the superfamily Acanthocerataceae, named by Meek in 1876, based on ''Ammonites vespertinu'', named by Morton in 1834. ''Mortoniceras'' is the type genus of the Mortoniceratinae, one of 4 subfam ...
'' has been found in a sandstone bed 2 metres above the amber horizon, alongside indeterminate gastropods and bivalves. Lead-uranium dating of zircon crystals of volcanic clasts within the amber bearing horizons has given a maximum age of 98.79 ± 0.62  million years ago ( Ma), making the deposit earliest
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
in age. Unpublished data by Wang Bo on other layers suggests an age range of deposition of at least 5 million years. The amber does not appear to have undergone significant transport since hardening or be redeposited. The strata at the site are younging upwards,
striking Strike may refer to: People *Strike (surname) Physical confrontation or removal *Strike (attack), attack with an inanimate object or a part of the human body intended to cause harm *Airstrike, military strike by air forces on either a suspected ...
north north-east and dipping 50-70 degrees E and SE north of the ridge and striking between south south-east and south-east and dipping 35-60 degrees south-west south of the ridge, suggesting the site is on the northwest limb of a syncline plunging to the northeast. A minor fault with a conspicuous gouge zone was noted as present, though it appeared to have no significant displacement. Several other localities are known, including the colonial Khanjamaw and the more recent Inzutzut, Angbamo, and Xipiugong sites, within the vicinity of Tanai. The Hkamti site SW of the Hukawng basin has been determined to be significantly older, dating to the early Albian around ca. 110 Ma and is therefore considered distinct.


Paleoenvironment

The Burmese amber paleoforest is considered to have been a tropical rainforest, situated near the coast, where resin was subsequently transported into a shallow marine environment. The shell of a dead juvenile ''Puzosia'' (''Bhimaites'') ammonite, four marine gastropod shells (including '' Mathilda'') and littoral or supralittoral isopods entombed in a piece of amber with shell sand, along with growth of Isocrinid crinoids, corals and oysters on the surface of some amber pieces indicate marine conditions for final deposition. Additionally pholadid (piddock) bivalve borings into amber specimens along with at least one pholadid which became trapped was interpreted to show that the resin was still fresh and unhardened when it was being moved into the tidal areas. However, the phloladids in question, belonging to the extinct genus '' Palaeolignopholas,'' were later interpreted as a freshwater species, and the presence of numerous freshwater insects suggests that the initial environment of deposition was a downstream
estuarine An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environment ...
to freshwater section of a river, with the forests extending across coastal rivers, river deltas, lakes, lagoons, and coastal bays. The forest environment may have been prone to fire, similar to modern tropical peat swamps, based on the presence of fire adapted plants and burned plant remains found in the amber. The amber itself is primarily disc-shaped and flattened along the bedding plane, and is typically reddish brown, with the colour ranging from shades of yellow to red. The opacity of the amber ranges from clear to opaque. Many amber pieces have thin
calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
veins that are typically less than , but up to thick. The number and proportion of veins in a piece of amber varies significantly, in some pieces veins are virtually absent, while others are described as being "packed with veinlets" The amber is considered to be of
coniferous Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extant ...
origin, with a likely araucarian source tree, based on spectroscopic analysis and wood fragment inclusions, though a pine origin has also been suggested.


Fauna and flora

The list of taxa is extraordinarily diverse, with over 42 classes, 108
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
, 569 families, 1017
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and 1379 species described as of the end of 2019, with over 300 species described in 2019 alone, the vast majority (94%) of which are arthropods. A complete list of taxa up until the end of 2018 can be found in Ross 2018 And a supplement covering most of 2019 can be found in Ross 2019b. For the sake of brevity, a complete list of taxa is not given here, and the classification is mostly at family level. For a more complete list of taxa, see
Paleobiota of Burmese amber Burmese amber is fossil resin dating to the early Late Cretaceous Cenomanian age recovered from deposits in the Hukawng Valley of northern Myanmar. It is known for being one of the most diverse Cretaceous age amber paleobiotas, containing rich arth ...
.


Invertebrates

Well over 1000 species of invertebrates are known from the deposit, including, notably the oldest members of Palpigradi (''
Electrokoenenia ''Electrokoenenia yaksha'' is a Palpigrade that lived approximately 100 million years ago during the Cretaceous period. It is the first microwhip scorpion fossil from this period to be found and is currently the oldest known Palpigrade. The pa ...
'') and Schizomida ('' Mesozomus'') the oldest Velvet worm (''
Cretoperipatus ''Cretoperipatus burmiticus'' is an extinct species of velvet worm that is known from Cretaceous Burmese amber approximately 100 million years old. It was found in Kachin state, Myanmar. Taxonomy The species can be assigned to one of the moder ...
'') and the only known fossil members of Mesothelae and Ricinulei since the Paleozoic. '' Chimerarachne'' is a unique stem spider still possessing a tail, with similar forms only known from the Paleozoic.


Arachnids


= Araneae

= Forty-four families of
spiders Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species dive ...
are known from the Burmese amber, including: Archaeidae, † Burmadictynidae, † Burmascutidae, † Burmathelidae, Clubionidae, Corinnidae, † Cretaceothelidae, Deinopidae,
Dipluridae The family Dipluridae, known as curtain-web spiders (or confusingly with other distantly related ones as funnel-web tarantulas) are a group of spiders in the infraorder Mygalomorphae, that have two pairs of booklungs, and chelicerae (fangs) that ...
, † Eopsilodercidae, †
Fossilcalcaridae Fossilcalcaridae is an extinct Mygalomorphae spider family in the clade Avicularioidea containing the single species ''Fossilcalcar praeteritus''. The family genus and species were described in 2015 from a male fossil entombed in Cretaceous age ...
,
Hersiliidae Hersiliidae is a tropical and subtropical family of spiders first described by Tamerlan Thorell in 1869, which are commonly known as tree trunk spiders. They have two prominent spinnerets that are almost as long as their abdomen, earning them ...
, Hexathelidae, †
Lagonomegopidae Lagonomegopidae is an extinct family of spiders known from the Cretaceous period. Members of the family are distinguished by a large pair of eyes, positioned on the anterolateral flanks of the carapace, with the rest of the eyes being small. They ...
,
Leptonetidae Leptonetidae is a relatively primitive family of spiders first described by Eugène Simon in 1890. It is made up of tiny haplogyne spiders, meaning they lack the hardened external female genitalia. Their six eyes are arranged in a semicircle of fo ...
,
Liphistiidae The spider family Liphistiidae, recognized by Tamerlan Thorell in 1869, comprises 8 genera and about 100 species of medium-sized spiders from Southeast Asia, China, and Japan. They are among the most basal living spiders, belonging to the subord ...
, † Micropalpimanidae, †
Mongolarachnidae ''Mongolarachne'' is an extinct genus of spiders placed in the monogeneric family Mongolarachnidae. The genus contains only one species, ''Mongolarachne jurassica'', described in 2013, which is presently the largest fossilized spider on record. ...
, Mysmenidae,
Ochyroceratidae Ochyroceratidae is a six-eyed spider family, with 165 described species in ten genera. They are common inhabitants of caves and the tropical forest litter of South Africa, the Caribbean, Asia and South America. Considered an ecological counterpar ...
, Oecobiidae, Oonopidae, Oxyopidae,
Palpimanidae Palpimanidae, also known as palp-footed spiders, is a family of araneomorph spiders first described by Tamerlan Thorell in 1890. They are widely distributed throughout the tropical and subtropical regions of the world, the Mediterranean and one in ...
, † Parvithelidae, Pholcidae, † Pholcochyroceridae, † Plumorsolidae, † Praearaneidae, † Praeterleptonetidae,
Psechridae Psechridae is a family (biology), family of Araneomorphae, araneomorph spiders with about 70 species in two genera. These are among the biggest cribellate spiders with body lengths up to and funnel webs more than in diameter. The family belon ...
,
Psilodercidae ''Psilodercidae'' is a family of spiders first described as a subfamily of Ochyroceratidae by Machado in 1951 and raised to family rank by J. Wunderlich in 2008. These spiders can be distinguished by the "segestriid positioning" of their six eyes, ...
, Salticidae,
Segestriidae Tube-dwelling spiders (Segestriidae) are a family of araneomorph spiders first described by Eugène Simon in 1893. It consists of five genera, two large and widespread, '' Segestria'' and ''Ariadna'', and three smaller genera, '' Citharoceps'', ...
,
Telemidae Telemidae, also known as long-legged cave spiders, is a family of small haplogyne spiders. Most are cave dwelling spiders with six eyes, though some do not have any eyes at all. There are about 104 described species in sixteen genera. Physical fe ...
, Tetrablemmidae,
Tetragnathidae Long-jawed orb weavers or long jawed spiders (Tetragnathidae) are a family of araneomorph spiders first described by Anton Menge in 1866. They have elongated bodies, legs, and chelicerae, and build small orb webs with an open hub with few, wid ...
,
Theridiosomatidae The ray spiders (Theridiosomatidae) are a family of spiders first described by Eugène Simon in 1881. They are most recognizable for their construction of cone-shaped webs. The family contains several genera which actively hunt for prey by using ...
, Theridiidae, Thomisidae, Uloboridae and † Vetiaroridae.


= Acariformes

= Twenty families of acariformes are known from the Burmese amber, including:
Anystidae Anystidae is a family of mites, based on the genus '' Anystis''. The family has a cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appro ...
, Archaeorchestidae,
Bdellidae Bdellidae is a family of snout mites in the order Trombidiformes. There are about 11 genera and at least 260 described species in Bdellidae. In terms of size, they are medium to large-sized predatory mites. They are known to inhabit soil, leaves ...
,
Caeculidae Caeculidae, also known as rake-legged mites, is a family of mites in the order Trombidiformes, the only family of the superfamily Caeculoidea. There are about 9 genera and about 100 described species in Caeculidae which occur world-wide. The olde ...
,
Cheyletidae Cheyletidae is a family of mites in the order Trombidiformes The Trombidiformes are a large, diverse order of mites. Taxonomy In 1998, Trombidiformes was divided into the Sphaerolichida and the Prostigmata. The group has few synapomorphies ...
, Enantioppiidae,
Eremaeidae Eremaeidae is a family of mites belonging to the order Sarcoptiformes. Genera: * ''Asperemaeus'' Behan-Pelletier, 1982 * ''Carinabella'' Hammer, 1977 * ''Eremaeus'' Koch, 1835 * ''Eueremaeus'' Mihelcic, 1963 * ''Rhynchobella'' Hammer, 1961 * ''T ...
, Erythraeidae,
Eupodidae Eupodidae is a family in the order Trombidiformes. There are at least two genera and three described species in Eupodidae. Genera * '' Claveupodes'' * ''Eupodes ''Eupodes'' is a genus of mite Mites are small arachnids (eight-legged arthr ...
,
Gymnodamaeidae Gymnodamaeidae is a family of oribatids in the order Oribatida. There are about 8 genera and at least 60 described species in Gymnodamaeidae. Genera * ''Adrodamaeus'' Paschoal, 1984 * ''Arthrodamaeus'' Grandjean, 1954 * ''Austrodamaeus'' Balogh & ...
,
Malaconothridae Malaconothridae is a family of mites belonging to the order Sarcoptiformes The Sarcoptiformes are an order of Acari comprising over 15,000 described species in around 230 families. Previously it was divided into two suborders, Oribatida and As ...
,
Microtrombidiidae Microtrombidiidae is a family of micro velvet mites in the order Trombidiformes The Trombidiformes are a large, diverse order of mites. Taxonomy In 1998, Trombidiformes was divided into the Sphaerolichida and the Prostigmata. The group has f ...
,
Neoliodidae Neoliodidae is a family of oribatids in the order Oribatida. There are at least 4 genera and 50 described species in Neoliodidae. Genera * ''Neoliodes'' Berlese, 1888 * ''Platyliodes'' Berlese, 1916 * ''Poroliodes'' Grandjean, 1934 * ''Teleiolio ...
,
Oribatellidae Oribatellidae is a family of mites belonging to the order Sarcoptiformes. Genera Genera: * ''Berniniella'' Özdikmen, 2008 * ''Cavernella'' Bernini, 1975 * ''Cuspidozetes'' Hammer, 1962 * ''Fberninia'' Özdikmen, 2008 * ''Fenestrobates'' Balog ...
, Oribotritiidae, Resinacaridae, Smarididae, Trombellidae, Trombidiidae and
Tuckerellidae The peacock mites of the genus ''Tuckerella'' ( the only genus of the mite family Tuckerellidae) are a significant herbivorous pest in the tropics, for example on citrus fruit. Other species dwell in grasses, possibly as root feeders. The peaco ...
.


= Opiliones

= Nine families of opiliones are known from the Burmese amber, including: Beloniscidae,
Epedanidae Epedanidae is a family of the harvestman infraorder Grassatores with about 200 described species. They are the sister group of the Gonyleptoidea. (2007): Epedanidae. Sørensen, 1886. In: Pinto-da-Rocha ''et al.'' 2007: 188. The Epedanidae are e ...
, † Halithersidae, † Mesokanidae,† Monooculricinuleidae, ? Pyramidopidae Sclerosomatidae,
Stylocellidae The Stylocellidae are a family of harvestmen with about 30 described species, all of which occur from India to New Guinea. Members of this family are from one to seven millimeters long. While ''Stylocellus'' species have eyes, these are absent in ...
and Tithaeidae.


= Pseudoscorpiones

= Twelve families of pseudoscorpions are known from the Burmese amber, including:
Atemnidae Atemnidae is a family of pseudoscorpion Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseudoscorpionida or Chelonethida. Pseudoscorpi ...
,
Cheiridiidae Cheiridiidae is a family of pseudoscorpions belonging to the order Pseudoscorpiones Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseu ...
,
Cheliferidae Cheliferidae is a family of pseudoscorpions in the order Pseudoscorpiones Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseudoscorpion ...
, Chernetidae,
Chthoniidae Chthoniidae is a family of pseudoscorpions within the superfamily Chthonioidea. The family contains more than 600 species in about 30 genera. Fossil species are known from Baltic, Dominican, and Burmese amber.Biology Catalog Chthoniidae now inc ...
,
Feaellidae The Feaellidae are a family of pseudoscorpions with the single genus ''Feaella''. Species Feaella Ellingsen, 1906 * subgenus ''Feaella'' Ellingsen, 1906 ** '' Feaella mirabilis'' Ellingsen, 1906 — western Africa ** '' Feaella mombasica'' Beie ...
,
Garypinidae Garypinidae is a family of pseudoscorpion Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseudoscorpionida or Chelonethida. Pseudosco ...
, Hyidae
Ideoroncidae Ideoroncidae is a family of pseudoscorpions belonging to the order Pseudoscorpiones. Members of the family are known from Asia, Africa, western North America and South America. Genera: * '' Afroroncus'' Mahnert, 1981 * '' Albiorix'' Chamberlin, 1 ...
, Neobisiidae, Pseudocheiridiidae and
Withiidae Withiidae is a family of pseudoscorpions belonging to the order Pseudoscorpiones Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseudos ...
.


= Scorpiones

= Seven families of scorpions are known from the Burmese amber, including: Buthidae,
Chaerilidae Chaerilidae is a family of scorpions. It contains two genera, the extant ''Chaerilus'' (Simon, 1877) and the extinct ''Electrochaerilus'' (Santiago-Blay, Fet, Soleglad & Anderson, 2004). References External links

* Scorpion familie ...
, † Chaerilobuthidae, † Palaeoburmesebuthidae, † Palaeoeuscorpiidae, † Palaeotrilineatidae and † Sucinolourencoidae.


= Parasitiformes

= Seven families of parasitiformes are known from the Burmese amber, including: Argasidae, † Deinocrotonidae, † Khimairidae, Ixodidae, Opilioacaridae,
Polyaspididae Polyaspididae is a family of mites in the order Mesostigmata Mesostigmata is an order of mites belonging to the Parasitiformes. They are by far the largest group of Parasitiformes, with over 8,000 species in 130 families. Mesostigmata includes ...
and
Sejidae Sejidae is a family of mites in the order Mesostigmata. The oldest known record of the group is an indeterminate deutonymph from the mid Cretaceous (Albian-Cenomanian) aged Burmese amber of Myanmar. Species '' Archaeopodella'' Athias-Henriot ...
.


= Schizomida

= One family of schizomida is known from the Burmese amber: Hubbardiidae


= Palpigradi

= One genus of palpigradi is known: ''
Electrokoenenia ''Electrokoenenia yaksha'' is a Palpigrade that lived approximately 100 million years ago during the Cretaceous period. It is the first microwhip scorpion fossil from this period to be found and is currently the oldest known Palpigrade. The pa ...
'', which belongs to Eukoeneniidae


= Amblypygi

= Two genera of Amblypygi are known: '' Kronocharon'' and '' Burmacharon'' which do not belong to any extant family.


= Solfugae

= One genus of
camel spider Solifugae is an order of animals in the class Arachnida known variously as camel spiders, wind scorpions, sun spiders, or solifuges. The order includes more than 1,000 described species in about 147 genera. Despite the common names, they are ne ...
is known: '' Cushingia'', which does not belong to any extant family.


= Thelyphonida

= Two genera of whip scorpion are known: '' Mesothelyphonus'', which belongs to
Thelyphonidae Thelyphonidae is the sole family in the Thelyphonida order of arachnids. Description These arachnids can reach a length of . The body consists of a cephalothorax coated with chitin and of an abdomen divided into 12 segments. Fossil record Th ...
and '' Burmathelyphonia,'' which does not belong to any extant family.


= Ricinulei

= Three genera of Ricinulei are known: '' Hirsutisoma'', ?'' Poliochera'' (an otherwise Carboniferous taxon) and '' Primoricinuleus,'' none of which belong to extant families


Myriapoda

Sixteen families of Myriapods are known, including:
Anthroleucosomatidae Anthroleucosomatidae is a family of millipedes in the order Chordeumatida. These millipedes range from 3.5 mm to 28 mm in length. Adult millipedes in this family have 26, 28, 30, or 31 segments (counting the collum as the first segment and the tel ...
, † Electrocambalidae,
Tingupidae Tingupidae is a family of millipedes in the order Chordeumatida. Adult millipedes in this family have 28 or 30 segments (counting the collum as the first segment and the telson The telson () is the posterior-most division of the body of an ar ...
,
Glomeridellidae Glomeridellidae is a family of millipedes belonging to the order Glomerida Glomerida is an order of pill-millipedes found primarily in the Northern Hemisphere. They superficially resemble pill-bugs or woodlice, and can enroll into a protecti ...
, Andrognathidae,
Paradoxosomatidae Paradoxosomatidae, the only family in the suborder Paradoxosomatidea (also known as Strongylosomatidea), is a family of flat-backed millipedes in the order Polydesmida. Containing nearly 200 genera and 975 species , it is one of the largest fami ...
,
Polydesmidae Polydesmidae is a family of millipedes in the order Polydesmida. These millipedes range from 4 mm to 30 mm in length. This family includes species notable for featuring sexual dimorphism in segment number: Adult females in the genus ''Perapolydes ...
, Polyxenidae,
Synxenidae Synxenidae is a family of bristly millipedes (Polyxenida). Three genera and around 10 species are known. Adult synxenids possess 15 or 17 pairs of legs, with the last two pair modified for small jumps. Adults in most species in this family have 1 ...
,
Polyzoniidae Polyzoniidae is a family of millipedes in the order Polyzoniida __NOTOC__ Polyzoniida is an order of millipedes in the Class_(biology)#Hierarchy_of_ranks_below_and_above_the_level_of_class, subterclass Colobognatha containing three families and ...
, Siphoniulidae,
Siphonophoridae Siphonophoridae is a family of millipede in the order Siphonophorida. There are about 12 genera and more than 110 described species in Siphonophoridae. Genera These 12 genera belong to the family Siphonophoridae: * ''Bactrois'' Cook, 1896 * ''B ...
,
Siphonorhinidae Siphonorhinidae is a family of millipede in the order Siphonophorida. There are at least 4 genera and about 12 described species in Siphonorhinidae. Genera These four genera belong to the family Siphonorhinidae: * '' Illacme'' Cook & Loomis, 19 ...
,
Zephroniidae Zephroniidae (sometimes misspelled "Zephronidae") is a family of giant pill millipedes in the taxonomic order Sphaerotheriida. They occur in southeast Asia from the Himalayas and China south and east to Sulawesi and to Australia, and also inhab ...
,
Cambalidae Cambalidae is a family of millipedes in the order Spirostreptida. There are at least 20 genera and 80 described species in Cambalidae. Genera References Further reading

* * * * Spirostreptida Millipede families {{myria ...
, Scolopendrellidae and †
Burmanopetalidae ''Burmanopetalum'' is an extinct genus of millipede containing the single species ''Burmanopetalum inexpectatum'' from the Cretaceous of Myanmar. It is a member of the order Callipodida and is the only member of the suborder Burmanopetalidea and ...
.


Entognatha

Eight families of Entognathans are known, including:
Campodeidae The Campodeidae are a family of hexapods belonging to the order Diplura. These pale, eyeless hexapods, the largest of which grow to around 12 mm in length, can be recognised by the two long, many-segmented cerci at the end of the abdomen. ...
, Japygidae, Isotomidae, †
Praentomobryidae :''Entomobryoidea is also the old name of Entomobryomorpha, when these were placed in the "Arthropleona" The Entomobryoidea are a superfamily of springtails (Collembola), tiny hexapods related to insects. In the modern sense, this group is placed ...
,
Tomoceridae The Entomobryomorpha are one of the three main groups (order) of springtails (Collembola), tiny hexapods related to insects. This group was formerly treated as a superfamily, the Entomobryoidea. They can be best distinguished from the other sp ...
, Neanuridae,
Odontellidae Odontellidae is a family of springtails in the order Poduromorpha. There are about 9 genera and more than 50 described species in Odontellidae. Genera These nine genera belong to the family Odontellidae: * '' Austrodontella'' Ellis & Bellinger, ...
and
Sminthuridae Sminthuridae is a family of springtails of the order Symphypleona. Sminthurids are commonly referred to as globular springtails. Description Like other Symphypleona, Sminthuridae are globular in shape and have a furcula that allows them to ju ...
.


Insects


= Archaeognatha

= Two families of archaeognathans are known from the Burmese amber: Machilidae and
Meinertellidae The Meinertellidae are a small family of basal insects belonging to the order Archaeognatha. They are sometimes known as rock bristletails. These insects can be distinguished from members of the other Archaeognatha family, Machilidae, by the l ...


= Zygentoma

= One family of Zygentoman is known: Lepismatidae


= Ephemeroptera

= Seven families of
mayfly Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) are aquatic insects belonging to the ord ...
are known: † Australiphemeridae, Baetidae, Ephemeridae,
Heptageniidae The Heptageniidae (synonym: Ecdyonuridae) are a family of mayflies with over 500 described species mainly distributed in the Holarctic, Oriental, and Afrotropical regions, and also present in the Central American Tropics and extreme northern Sou ...
, † Hexagenitidae, Isonychiidae,
Prosopistomatidae Prosopistomatidae is a family of mayflies. There is one extant genus, '' Prosopistoma,'' with several dozen species found across Afro-Eurasia and Oceania. They are noted for their unusual beetle-shaped larvae, which live beneath rocks and stones ...
.


= Odonata

= Twenty families of odonatan are known from the Burmese amber, including: Aeshnidae, † Araripegomphidae, † Burmacoenagrionidae, † Burmaeshnidae, † Burmagomphidae, † Burmaphlebiidae, Calopterygidae, Coenagrionidae, Dysagrionidae, Gomphaeschnidae, Gomphidae, Hemiphlebiidae,
Libellulidae The skimmers or perchers and their relatives form the Libellulidae, the largest dragonfly family in the world. It is sometimes considered to contain the Corduliidae as the subfamily Corduliinae and the Macromiidae as the subfamily Macromiinae. Ev ...
,
Lindeniidae Lindeniidae is a family of dragonflies occurring in Australia. The family Lindeniidae is not recognised in the World Odonata List at the Slater Museum of Natural History, but rather its species are considered to be part of the Gomphidae family. ...
, Megapodagrionidae, † Mesomegaloprepidae, † Paracoryphagrionidae, Perilestidae, Platycnemididae, Platystictidae.


= Hymenopterans

= Over fifty families of hymenopterans have been described beginning with the papers of
Cockerell Cockerell is a surname, and may refer to: People *Allan Cockerell (1891–1975), New Zealand soldier and politician * Catherine Cockerell Cobb (1903–1995), British jeweler, silversmith, daughter of Douglas *Charles Robert Cockerell (1788–1863), ...
who described a group of
Bethylidae The Bethylidae are a family of aculeate wasps in the superfamily Chrysidoidea. As a family, their biology ranges between parasitoid wasps and hunting wasps. Overview Like most of the Chrysidoidea, the Bethylidae are stinging Hymenoptera and m ...
and
Aulacidae The Aulacidae are a small, cosmopolitan family of wasps, with two extant genera containing some 200 known species. They are primarily endoparasitoids of wood wasps (Xiphydriidae) and xylophagous beetles (Cerambycidae and Buprestidae). They are c ...
species between 1917 and 1920. The monotypic family Melittosphecidae is only known from the Burmese amber species ''
Melittosphex burmensis ''Melittosphex burmensis'' is an aculeate wasp that was formerly considered one of the two oldest-known species of bees. The species was described from an inclusion in Burmese amber in the year 2006 by George Poinar Jr., a zoologist at Oregon ...
'' and eight species belonging to ''
Aptenoperissus ''Aptenoperissus'' is a genus of extinct wasp with eight described species, placed into the monotypic family Aptenoperissidae. The type species ''Aptenoperissus burmanicus'' resembles a mix between a grasshopper, an ant, and a wasp. It was descri ...
'' of the monotypic family Aptenoperissidae are also known. Originally described as an Aneuretinae ant ''
Burmomyrma rossi ''Burmomyrma'' is an extinct genus of aculeate hymenopteran, suggested to be an ant. The genus contains a single described species, ''Burmomyrma rossi''. ''Burmomyrma'' is known from a single Middle Cretaceous fossil which was found in Asia. ...
'' was moved to the extinct Chrysidoidea family Falsiformicidae. A number of Formicidae species known, belonging to ''
Baikuris ''Baikuris'' is an extinct genus of ant in the Formicidae subfamily Sphecomyrminae, and is currently placed in the tribe Sphecomyrmini. The genus contains four described species: the type species ''Baikuris mandibularis'', along with ''Baikur ...
'' (indet) ''
Camelomecia janovitzi ''Camelomecia'' is an extinct genus of stem-group ants not placed into any Formicidae subfamily. Fossils of the single known species, ''Camelomecia janovitzi'', are known from the Middle Cretaceous of Asia. The genus is one of several ants des ...
, Ceratomyrmex ellenbergeri,'' 11 species of ''
Gerontoformica ''Gerontoformica'' is an extinct genus of stem-group ants. The genus contains thirteen described species known from Late Cretaceous fossils found in Asia and Europe. The species were described between 2004 and 2016, with a number of the specie ...
,'' 3 species of '' Haidomyrmex'', '' Linguamyrmex vladi'', 2 species of '' Zigrasimecia'', '' Dhagnathos autokrator'', '' Chonidris insolita'', '' Aquilomyrmex huangi'', '' Protoceratomyrmex revelatus'' and ''
Linguamyrmex brevicornis ''Linguamyrmex'' is an extinct genus of ant in the formicid subfamily Haidomyrmecinae, and is one of only nine genera placed in the subfamily Haidomyrmecinae. The genus contains three described species, ''Linguamyrmex brevicornis'', ''Lingua ...
''. Other families include
Ampulicidae The Ampulicidae, or cockroach wasps, are a small (about 170 species), primarily tropical family of sphecoid wasps, all of which use various cockroaches as prey for their larvae. They tend to have elongated jaws, pronounced neck-like constrictions ...
, Braconidae, Cephidae,
Ceraphronidae The Ceraphronidae are a small hymenopteran family with 14 genera and some 360 known species, though a great many species are still undescribed. It is a poorly known group as a whole, though most are believed to be parasitoids (especially of f ...
, Chalcididae, Chrysididae, Crabronidae, Diapriidae, Dryinidae,
Embolemidae Embolemidae is a family of small solitary parasitoid wasps with around 70 species in 2 genera distributed around the world.van Achterberg, Cornelis & Kats, R.. (2000). Revision of the Palaearctic Embolemidae (Hymenoptera). Zoöl. Med. 74 (2000), ...
, Evaniidae,
Gasteruptiidae The Gasteruptiidae are one of the more distinctive families among the apocritan wasps, with surprisingly little variation in appearance for a group that contains around 500 species in two subfamilies ( Gasteruptiinae and Hyptiogastrinae) and wi ...
,
Geoscelionidae Geoscelionidae is a family of wasps in the superfamily Platygastroidea. It contains three extant species in two genera, native to South America and Africa, and several other genera known from fossils. It was originally erected as the tribe Geoscel ...
,
Heloridae Heloridae is a family of wasps in the order Hymenoptera known primarily from fossils, and only one extant genus, '' Helorus'', with 12 species found worldwide. Members of ''Helorus'' are parasitic on green lacewings. Extant genera *'' Helorus'' ( ...
, Ichneumonidae,
Megalyridae Megalyroidea is a small hymenopteran superfamily of wasps that includes a single family, Megalyridae, with eight extant genera (plus around a dozen extinct ones) and 49 described species. Modern megalyrids are found primarily in the southern hem ...
,
Megaspilidae The Megaspilidae are a small hymenopteran family with 13 genera in two subfamilies, and some 450 known species, with a great many species still undescribed. It is a poorly known group as a whole, though most are believed to be parasitoids (espe ...
, Mymaridae, Mymarommatidae,
Pelecinidae Pelecinidae is a family of parasitic wasps in the superfamily Proctotrupoidea. It contains only one living genus, '' Pelecinus'', with three species known from the Americas. The earliest fossil species are known from the Jurassic, and the group ...
, Platygastridae,
Rhopalosomatidae Rhopalosomatidae is a family of Hymenoptera containing about 68 extant species in four genera that are found worldwide. Three fossil genera are known. The adults resemble ants and may be confused with them. They are yellowish with red or brown m ...
.
Rotoitidae The Rotoitidae are a very small family of rare, relictual parasitic wasps in the superfamily Chalcidoidea, known primarily from fossils (14 extinct species in two genera, '' Baeomorpha'' and '' Taimyromorpha'').Huber JT, Shih C, Dong R (2019) A n ...
,
Sapygidae The Sapygidae are a family of solitary kleptoparasitic aculeate wasps. They are generally black wasps, similar in appearance to some Tiphiidae or Thynnidae, with white or yellow markings developed to various degrees. The female oviposits her egg ...
,
Scelionidae The hymenopteran family Scelionidae is a very large cosmopolitan group (over 3000 described species in some 176 genera) of exclusively parasitoid wasps, mostly small (0.5–10 mm), often black, often highly sculptured, with (typically) elbowe ...
,
Sclerogibbidae The Sclerogibbidae are a small family of aculeate wasps in the superfamily Chrysidoidea. Overview Sclerogibbidae are ectoparasitoids of Embioptera. The female wasp oviposits an egg on the abdomen of a host. Once the larva emerges, it attaches i ...
,
Scolebythidae The Scolebythidae are a small family of aculeate wasps in the superfamily Chrysidoidea. These chrysidoid wasps are found in Africa, Australia, the Neotropics, north China, Thailand and Fiji. They are parasites on larvae of Cerambycidae and Ptini ...
,
Sepulcidae Sepulcidae is an extinct family of stem sawflies in the order Hymenoptera. The family is known primarily from late Mesozoic fossils found in 1968 in Transbaikalia. The insects were distant relatives of modern sawflies and are part of the living ...
, Sierolomorphidae, Siricidae,
Sparasionidae Sparasionidae is a family of wasps in the superfamily Platygastroidea. Known species are parasitioids of the eggs of orthopterans. Taxonomy * '' Archaeoteleia'' Masner Burmese amber, Late Cretaceous ( Cenomanian) Baltic amber, Eocene The ...
,
Sphecidae The Sphecidae are a cosmopolitan family of wasps of the suborder Apocrita that includes sand wasps, mud daubers, and other thread-waisted wasps. The name Sphecidae was formerly given to a much larger grouping of wasps. This was found to be p ...
,
Stephanidae The Stephanidae, sometimes called crown wasps, are a family of parasitoid wasps. They are the only living members of the superfamily Stephanoidea. Stephanidae has at least 345 living species in 11 genera. The family is considered cosmopolitan in ...
, Tiphiidae, Vespidae, Xiphydriidae, † Angarosphecidae, †
Aptenoperissidae ''Aptenoperissus'' is a genus of extinct wasp with eight described species, placed into the monotypic family Aptenoperissidae. The type species ''Aptenoperissus burmanicus'' resembles a mix between a grasshopper, an ant, and a wasp. It was descr ...
, † Bryopompilidae, † BurmorussidaeBurmusculidae, † Chrysobythidae, † Dipterommatidae, †
Diversinitidae Diversinitidae are an extinct family of Chalcid wasps. Three genera are known, all from the early Cenomanian aged Burmese amber. They are distinguished by the presence of multiporous plate sensilla on the first flagellomere Antennae ( antenna ...
, † Falsiformicidae, † Gallorommatidae, † Ohlhoffiidae, †
Panguidae Panguidae is an extinct family of aculeate wasps, known exclusively from the mid-Cretaceous (latest Albian-earliest Cenomanian) Burmese amber of Myanmar. The family was initially named for the genus ''Pangu'', with the genus '' Prosphex,'' orig ...
, †
Plumalexiidae ''Plumalexius'' is a genus of wasps in the extinct monotypic family Plumalexiidae, containing two species: the type species ''Plumalexius rasnitsyni'', known from the Late Cretaceous White Oaks Pit in Sayreville, New Jersey, and ''Plumalexius ohm ...
, †
Maimetshidae Maimetshidae is an extinct family of wasps, known from the Cretaceous period.M. S. Engel. 2016A new genus and species of maimetshid wasps in Lebanese Early Cretaceous amber (Hymenoptera: Maimetshidae).''Novitates Paleoentomologicae'' 18:1-14 While ...
, † Myanmarinidae, †
Othniodellithidae Praeaulacidae is an extinct family of Mesozoic parasitic wasps in the suborder Evanioidea. It among the earliest known families of the group and is characterised by more complete wing venation in comparison to other members of the suborder. It ha ...
, †
Peleserphidae Peleserphidae is an extinct family of wasps belonging to the superfamily Proctotrupoidea. It is currently known from four species in three genera. One ('' Arkadiserphus'') from the Middle-Late Jurassic Karabastau Formation of Kazakhstan, and the ...
, †
Praeaulacidae Praeaulacidae is an extinct family of Mesozoic parasitic wasps in the suborder Evanioidea. It among the earliest known families of the group and is characterised by more complete wing venation in comparison to other members of the suborder. It ha ...
, †
Proterosceliopsidae ''Proterosceliopsis'' is an extinct genus of platygastroid parasitic wasp, known from the Mid-Cretaceous of Eurasia. The genus was first described in 2014 from the Albian amber of the Escucha Formation. In 2019 additional species were described ...
, †
Serphitidae Serphitidae is a family of microscopic parasitic wasps known from the Cretaceous period. Taxonomy This family was described in 1937 by the American entomologist Charles Thomas Brues to classify a fossil insect caught in an amber piece from Cana ...
, †
Spathiopterygidae Spathiopterygidae is an extinct family of small parasitic wasps, known from the Cretaceous (Barremian to Santonian) of Laurasia and Northern Gondwana. They are suggested to be members of Diaprioidea, in part due to their similarly reduced wing v ...
, † Syspastoxyelidae and several ''incertae sedis'' taxa.


= Dipterans

= Over forty families of dipterans are known from the Burmese amber, including: Acroceridae, Anisopodidae,
Apsilocephalidae Apsilocephalidae is a family of flies in the superfamily Asiloidea. It was historically treated as a subfamily within Therevidae, but placed in a separate family in 1991, and subsequently recognized as more distantly related. The family contains ...
,
Apystomyiidae Apystomyiidae is a small family (biology), family of fly, flies containing the living genus ''Apystomyia'' and the extinct genera ''Apystomimus'' and ''Hilarimorphites''. The single living Apystomyiidae species, ''Apystomyia elinguis'', is native ...
, Asilidae,
Atelestidae Atelestidae is a family of flies in the superfamily Empidoidea. The four genera were placed in a separate family in 1983; they were formerly either in Platypezidae (which are not even particularly closely related) or considered ''incertae sedis'' ...
,
Blephariceridae The Blephariceridae, commonly known as net-winged midges, are a nematoceran family (biology), family in the order Fly, Diptera. The adults resemble tipulidae, crane flies except with a projecting anal angle in the wings, and different head shape, ...
, Bombyliidae, Cecidomyiidae, Ceratopogonidae, Chaoboridae, Chironomidae, Corethrellidae, Culicidae, Diadocidiidae, Dolichopodidae, Empididae,
Hybotidae Hybotidae, the typical dance flies, are a family of true flies. They belong to the superfamily Empidoidea and were formerly included in the Empididae as a subfamily. Some, such as '' Tachydromia'', are predators that run around on the bark of t ...
,
Ironomyiidae Ironomyiidae is a small family of flies in the order Diptera. Historically, they had been included in the family Platypezidae, and includes three extant species within the single extant genus ''Ironomyia'' endemic to Australia and a number of exti ...
, Keroplatidae, Limoniidae, Lygistorrhinidae, Mycetophilidae, Mythicomyiidae, Nemestrinidae, Phoridae, Pipunculidae, Platypezidae, Psychodidae, Ptychopteridae,
Rachiceridae The Brachyceran infraorder Xylophagomorpha is a small group that consists solely of the family Xylophagidae, which presently contains subfamilies that were sometimes considered to be two small related families (Coenomyiidae and Rachiceridae). ...
, Rhagionidae, Scatopsidae, Sciaridae,
Stratiomyidae The soldier flies (Stratiomyidae, sometimes misspelled as Stratiomyiidae, from Greek - soldier; - fly) are a family of flies (historically placed in the now-obsolete group Orthorrhapha). The family contains over 2,700 species in over 380 extan ...
, Tabanidae, Tanyderidae, Tipulidae, Valeseguyidae, Xylomyidae, †
Archizelmiridae Archizelmiridae is an extinct family of flies, known from the Jurassic and Cretaceous periods. It belongs to the Sciaroidea, and has suggested to have a close relationship with Sciaridae. Genera * '' Archimelzira'' Grimaldi et al. 2003 New J ...
, † Chimeromyiidae, †
Eremochaetidae Eremochaetidae is an extinct family of brachyceran flies known from the Jurassic and Cretaceous periods of Asia. It is part of the extinct superfamily Archisargoidea. The morphology of the ovipositor of the only 3 dimensionally preserved genus '' ...
, † Eucaudomyiidae, † Mysteromyiidae, †
Rhagionemestriidae Rhagionemestriidae is an extinct family of brachyceran flies known from the Jurassic and Cretaceous periods. It was first named as a subfamily of the Nemestrinidae by Ussatchov (1968), and was raised to full family status by Nagatomi and Yang (19 ...
, †
Tethepomyiidae Tethepomyiidae is an extinct family of small brachyceran flies known from the Cretaceous period of Laurasia. It is part of the extinct superfamily Archisargoidea. The family is characterised by "very large eyes, reduced mouthparts, a highly reduc ...
, † Zhangsolvidae and several ''incertae sedis'' taxa.


= Coleopterans

= Over ninety families of coleopterans are known from the Burmese amber, including:
Aderidae The Aderidae, sometimes called ant-like leaf beetles, are a family of beetles that bear some resemblance to ants. The family consists of about 1,000 species in about 40 genera, of which most are tropical, although overall distribution is worldwid ...
, Anthicidae, Anthribidae, † Apotomouridae, Belidae, Boganiidae, Bostrichidae, Brachypsectridae,
Buprestidae Buprestidae is a family of beetles known as jewel beetles or metallic wood-boring beetles because of their glossy iridescent colors. Larvae of this family are known as flatheaded borers. The family is among the largest of the beetles, with some ...
, Cantharidae, Carabidae, Caridae, Cerambycidae, Cerophytidae,
Cerylonidae Cerylonidae are small to tiny (), smooth, shiny, hairless beetles, only lightly punctured. There are about 450 species worldwide in 50 or so genera, mostly tropical and subtropical. They are most common under the bark of dead trees, but can also ...
,
Chrysomelidae The insects of the beetle family Chrysomelidae are commonly known as leaf beetles, and include over 37,000 (and probably at least 50,000) species in more than 2,500 genera, making up one of the largest and most commonly encountered of all beetle ...
, Ciidae, Clambidae,
Cleridae Cleridae are a family of beetles of the superfamily Cleroidea. They are commonly known as checkered beetles. The family Cleridae has a worldwide distribution, and a variety of habitats and feeding preferences. Cleridae have many niches and feed ...
,
Cucujidae The Cucujidae, "flat bark beetles," are a family of distinctively flat beetles found worldwide (except Africa and Antarctica) under the bark of dead trees. The family has received considerable taxonomic attention in recent years and now consists ...
, Cupedidae,
Curculionidae The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families, with 6,800 genera and 83,000 species described worldwide. They are the sister group to the family Brentidae. T ...
, Cyclaxyridae,
Dascillidae Dascillidae is a family of beetles within the clade Elateriformia. There are about 100 extant species in 11 genera, which are found worldwide. Dascillidae together with Rhipiceridae form the super family Dascilloidea. Taxonomy The family was na ...
, Dermestidae, Dytiscidae, Elateridae, Elmidae,
Endomychidae Endomychidae, or handsome fungus beetles, is a family of beetles with representatives found in all biogeographic realms. There are around 120 genera and 1300 species. The family was established based on the type genus '' Endomychus'', a genus ere ...
, Eucinetidae, Eucnemidae, Geotrupidae, Glaresidae, Gyrinidae, Heteroceridae, Histeridae, Hybosoridae, Hydraenidae, Hydrophilidae,
Jacobsoniidae Jacobsoniidae are a family of tiny beetles belonging to Staphylinoidea. The larvae and adults live under bark, in plant litter, fungi, bat guano and rotten wood. There are around 28 described species in three genera: Description Members of this ...
, Kateretidae,
Laemophloeidae Laemophloeidae, "lined flat bark beetles," is a family in the superfamily Cucujoidea characterized by predominantly dorso-ventrally compressed bodies, head and pronotal discs bordered by ridges or grooves, and inverted male genitalia. Size range ...
, Lampyridae,
Latridiidae Latridiidae (sometimes spelled "Lathridiidae") is a family of tiny, little-known beetles commonly called minute brown scavenger beetles or fungus beetles. The number of described species currently stands at around 1050 in 29 genera but the number ...
,
Leiodidae Leiodidae is a family of beetles with around 3800 described species found worldwide. Members of this family are commonly called round fungus beetles due to the globular shape of many species, although some are more elongated in shape. They are g ...
,
Lepiceridae ''Lepicerus'' is a genus of myxophagan beetles containing three described species in the family Lepiceridae; it is the only extant genus in the family, with another genus, '' Lepiceratus'' only known from fossils.Jałoszyński, Paweł; Luo, Xiao- ...
, Lucanidae,
Lycidae The Lycidae are a family in the beetle order Coleoptera, members of which are commonly called net-winged beetles. These beetles are cosmopolitan, being found in Nearctic, Palearctic, Neotropical, Afrotropical, Oriental, and Australian ecoregions ...
,
Lymexylidae The Lymexylidae (historically often spelled Lymexylonidae), 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. Habitat a ...
, Melandryidae, Meloidae, Melyridae, † Mesophyletidae,
Micromalthidae The telephone-pole beetle (''Micromalthus debilis'') is a beetle native to the eastern United States, and the only living representative of the otherwise extinct family Micromalthidae. They have an unusual lifecycle involving asexually reproducin ...
,
Monotomidae Monotomidae is a family of beetles in the superfamily Cucujoidea. The family is found worldwide, with approximately 240 species in 33 genera. The ecological habits of the family are diverse, with different members of the group being found under t ...
,
Mordellidae The Mordellidae are a family of beetles commonly known as tumbling flower beetles for the typical irregular movements they make when escaping predators, or as pintail beetles due to their abdominal tip which aids them in performing these tumbling ...
, †
Mysteriomorphidae ''Mysteriomorphus'' is an extinct genus of beetle, and the only member of the family Mysteriomorphidae. It is known from one described species, ''Mysteriomorphus pelevini'' found in Burmese amber, dating to the early part of the Cenomanian stage ...
, Nemonychidae, Nitidulidae, Oedemeridae,
Ommatidae 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. How ...
, Passalidae, † Parandrexidae, † Passalopalpidae,
Passandridae Passandridae, the "parasitic flat bark beetles," are a family of beetles notable for being one of the very few beetle families with larvae that are, as far as known, exclusively ectoparasitic on the immature stages of other beetles and Hymenoptera ...
,
Phloeostichidae Phloeostichidae is a family of beetles in the superfamily Cucujoidea. They are typically found under the bark of dead trees. Larvae have been found to consume plant tissue and some fungi, while the adults appear to be exclusively fungivores. The ...
, Prostomidae, Psephenidae, Ptiliidae, Ptinidae, Ptilodactylidae, Ripiphoridae, Salpingidae, Scarabaeidae, Scirtidae,
Scraptiidae The family Scraptiidae is a small group of tenebrionoid beetles sometimes called false flower beetles. There are about 400 species in 30 genera with a world-wide distribution. The adults are found on flowers, sometimes in large numbers, but are a ...
, Silphidae, Silvanidae,
Smicripidae Smicripidae is a family of beetles, in the superfamily Cucujoidea. The common name for this family is palmetto beetles. The family only has one extant genus, '' Smicrips,'' with six extant species native to tropical and subtropical regions of th ...
, Sphaeriusidae,
Staphylinidae The rove beetles are a family (Staphylinidae) of beetles, primarily distinguished by their short elytra (wing covers) that typically leave more than half of their abdominal segments exposed. With roughly 63,000 species in thousands of genera, the ...
, Tenebrionidae, Tetratomidae, Thanerocleridae, Throscidae,
Trogidae Trogidae, sometimes called hide beetles, is a family of beetles with a distinctive warty or bumpy appearance. Found worldwide, the family includes about 300 species contained in four or five genera. Trogids range in length from 2 to 20 mm. Thei ...
,
Trogossitidae Trogossitidae, also known as bark-gnawing beetles, are a small family in the superfamily Cleroidea. Many taxa formerly within this family have been removed (as of 2019) to other families, such as Lophocateridae, Peltidae, Protopeltidae, Rentoniid ...
and
Zopheridae The Zopheridae family of beetles has grown considerably in recent years as the members of two other families have been included within its circumscription; these former families are the Monommatidae and the Colydiidae, which are now both incl ...
.


= Neuroptera

= Over twenty families of
neuroptera The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in th ...
ns are known from the Burmese amber, including: † Araripeneuridae,
Ascalaphidae Ascalaphidae is a family of insects in the order Neuroptera, commonly called owlflies; there are some 450 extant species. They are fast-flying crepuscular or diurnal predators of other flying insects, and have large bulging eyes and strongly ...
, †
Babinskaiidae Babinskaiidae is an extinct family of neuropterans known from the Cretaceous period. They are part of the superfamily Myrmeleontoidea. Their distinguishing characters include: "long filiform antennae, narrowly elongated wings, with features suc ...
, Berothidae, Chrysopidae,
Coniopterygidae The dustywings, Coniopterygidae, are a family of Pterygota (winged insects) of the net-winged insect order ( Neuroptera). About 460 living species are known.Engel & Grimaldi (2007) These tiny insects can usually be determined to genus with a han ...
, † Corydasialidae, Dilaridae, †
Dipteromantispidae Dipteromantispidae is an extinct family of neuropterans known from the Cretaceous period. Unlike other neuropterans, the family possesses only a single set of fully developed forewings, with the hindwings reduced to haltere like structures. They ...
,
Hemerobiidae Hemerobiidae is a family of Neuropteran insects commonly known as brown lacewings, comprising about 500 species in 28 genera. Most are yellow to dark brown, but some species are green. They are small; most have forewings 4–10 mm long (som ...
, Ithonidae, Kalligrammatidae, Mantispidae, †
Mesochrysopidae Mesochrysopidae is an extinct family of lacewings known from the Jurassic and Cretaceous periods. They are considered to be closely related to green lacewings of the family Chrysopidae. The family are also alternatively considered a paraphyletic ...
,
Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family Myrmeleontidae. They are known for the predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. In North America, the l ...
, Nemopteridae, Nevrorthidae,
Nymphidae Nymphidae, sometimes called split-footed lacewings, are a family of winged insects of the order Neuroptera. There are 35 extant species native to Australia and New Guinea. Nymphidae stand somewhat apart from other living Myrmeleontoidea. The ...
, Osmylidae, Psychopsidae, Rachiberothidae, Sisyridae and several ''incertae sedis'' taxa.


= Hemiptera

= Over sixty families of
hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
ns are known from the Burmese amber, including: Achilidae, †
Albicoccidae ''Albicoccus'' is an extinct genus of scale insect in the extinct monotypic family Albicoccidae, containing a single species, ''Albicoccus dimai''. The genus is solely known from the Albian - Cenomanian Burmese amber deposits. History and class ...
,
Aleyrodidae Whiteflies are Hemipterans that typically feed on the undersides of plant leaves. They comprise the family Aleyrodidae, the only family in the superfamily Aleyrodoidea. More than 1550 species have been described. Description and taxonomy The A ...
, Aphrophoridae,
Aradidae Aradidae is a family of flat bugs, and a member of true bugs (Heteroptera). The common name for these insects refers to their dorsoventrally flattened bodies. With few exceptions, these cryptic insects are of no economic importance. This famil ...
, †
Berstidae ''Bersta'' is an extinct genus of hemipteran in the monotypic family Berstidae. It is known from two species found in the Cenomanian aged Burmese amber of Myanmar. The external morphology suggests that the genus were beetle mimics. Etymology ...
, †
Burmacoccidae ''Burmacoccus'' is an extinct genus of scale insect in the extinct monotypic family Burmacoccidae, containing a single species, ''Burmacoccus danyi''. The genus is solely known from the Albian – Cenomanian Burmese amber deposits. History a ...
, † Burmitaphidae,
Callaphididae Callaphididae is a family of true bugs belonging to the order Hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, ...
, Cercopidae, Cicadellidae, Cicadidae, Cimicidae, Cixiidae,
Coccidae The Coccidae are a family of scale insects belonging to the superfamily Coccoidea. They are commonly known as soft scales, wax scales or tortoise scales. The females are flat with elongated oval bodies and a smooth integument which may be cover ...
, Coreidae, Cydnidae,
Dictyopharidae Dictyopharidae is a family of planthoppers, related to the Fulgoridae. The family comprises nearly 760 species in more than 150 genera which are grouped into two subfamilies, Dictyopharinae and Orgeriinae. Description Like all other fulgoroi ...
, † Dinglidae
Dipsocoridae Dipsocoridae are a family of heteropteran bugs known as jumping ground bugs. There are about 30 widely distributed species which are placed in three genera. Fossils from Eocene amber have also been placed in the family. The antennae have four se ...
, † Dorytocidae, Enicocephalidae, Fulgoridae,
Gelastocoridae The Gelastocoridae (toad bugs) is a family of about 100 species of insects in the suborder Heteroptera. These fall into two genera, about 15 species of '' Gelastocoris'' from the New World and 85 of '' Nerthra'' from the Old World. They are rem ...
, Gerridae, † Hodgsonicoccidae, Hydrometridae, Issidae, † Jubisentidae, †
Juraphididae Juraphididae is an extinct family of aphids in the order Hemiptera. There are at least three genera and about five described species in Juraphididae. The most recent discovery in Juraphididae family was the Prolavexillapis munditia in 2018 and Is ...
, † Katlasidae, Kinnaridae, † Kozariidae, † Lalacidae,
Leptopodidae Leptopodidae is a family of spiny-legged bugs in the order Hemiptera. There are about 15 genera and more than 40 described species in Leptopodidae. Genera These 15 genera belong to the family Leptopodidae: * '' Erianotoides'' J.Polhemus & D.Po ...
, † Liadopsyllidae, † Macrodrilidae,
Margarodidae The Margarodidae (illegitimately as Margodidae) or ground pearls are a family of scale insects within the superfamily Coccoidea. Members of the family include the Polish cochineal and Armenian cochineal (genus ''Porphyrophora'') and the original ...
,
Matsucoccidae Matsucoccidae is a family of scales and mealybugs in the order Hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers ...
, †
Mimarachnidae Mimarachnidae is an extinct family of planthoppers known from the Cretaceous period. Their name is derived from spots on the wings of the first described genera, ''Mimarachne'' and ''Saltissus'', being suggestive of spider mimicry, but these char ...
, † Minlagerrontidae, Miridae,
Monophlebidae Monophlebidae is a family of scale insects commonly known as the giant scales or monophlebids. They occur in most parts of the world but more genera are found in the tropics than elsewhere.Naucoridae Naucoridae is a small family of insects commonly known as the creeping water bugs and saucer bugs. They are similar in appearance and behavior to Belostomatidae (giant water bugs), but considerably smaller, at long. Naucoridae are found around the ...
, Nabidae, †
Neazoniidae Neazoniidae is an extinct family of planthoppers. It was named by Szwedo in 2007, and contains 2 genera. Classification * † '' Akmazeina'' Szwedo, 2009 * † '' Neazonia'' Szwedo, 2007 References {{Taxonbar, from=Q21351626 Fulgoromorph ...
,
Ochteridae The Ochteridae comprise a small family of insects. Eight genera with about 80 species have been described. They occur worldwide along the shore of various types of water (for example, streams and ponds) and the greatest diversity is in tropical ...
,
Ortheziidae Ortheziidae is a family of scale insects commonly known as the ensign scales or ortheziids. They occur in most parts of the world but the majority of the species are found in the Neotropical and Nearctic regions while there are not many species i ...
,† Palaeoleptidae, † Parvaverrucosidae, †
Perforissidae Perforissidae is an extinct family of planthoppers. They are considered to belong to the group of " Cixiidae-like" planthoppers. Species are known from the Early to Late Cretaceous of Eurasia, North America and South America. The family was name ...
, † Protopsyllidiidae, † Procercopidae,
Pseudococcidae Mealybugs are insects in the family Pseudococcidae, unarmored scale insects found in moist, warm habitats. Many species are considered pests as they feed on plant juices of greenhouse plants, house plants and subtropical trees and also act as a ...
, Reduvidae,
Schizopteridae Schizopteridae is the largest family in the infraorder Dipsocoromorpha and comprises 56 genera and approximately 255 species.Emsley, M.G., 1969. The Schizopteridae (Hemiptera: Heteroptera) with the descriptions of new species from Trinidad. Memo ...
, † Sinoalidae, † Tajmyraphididae, Tettigarctidae,
Tingidae The Tingidae are a family of very small () insects in the order Hemiptera that are commonly referred to as lace bugs. This group is distributed worldwide with about 2,000 described species. They are called lace bugs because the pronotum and f ...
,
Tropiduchidae Tropiduchidae is a family of planthoppers in the order Hemiptera. There are at least 160 genera and 600 described species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of ...
, Velocipedidae, Veliidae, † Weitschatidae,
Xylococcidae Xylococcidae is a family of scales and mealybugs in the order Hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, ...
, †
Yetkhatidae Yetkhatidae is an extinct family of planthoppers. It was named by Song, Xu, Liang, Szwedo & Bourgoin in 2019, and contains 2 genera. Genera * † '' Parwaina'' Song, Szwedo & Bourgoin, 2019 * † '' Yetkhata'' Song, Szwedo & Bourgoin, 2019 Refe ...
, † Yuripopovinidae and several ''incertae sedis'' taxa such as ''
Mesophthirus ''Mesophthirus'' is an extinct genus of insect known from Burmese Amber, Burmese amber from Myanmar during the mid-Cretaceous period, about 100 million years ago. Its Monotypic taxon, sole species, ''Mesophthirus engeli'', is known from multiple ...
'', formerly believed to be an
ectoparasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
but since determined to be a basal
scale insect Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than the ...
.


= Dictyoptera

= Twenty one families of dictyopterans are known from the Burmese amber, including: Blaberidae, †
Blattulidae Blattulidae is an extinct family of cockroaches known from the Triassic to the Late Cretaceous. Their distinguishing characteristics include "forewing has long Sc, regular venation with distinct intercalaries and hindwing has simple CuP, branched ...
,
Blattidae Blattidae is a cockroach family in the order Blattodea containing several of the most common household cockroaches. Some notable species include: * ''Blatta orientalis'': Oriental cockroach, * Common shining cockroach: (''Drymaplaneta commun ...
, † Caloblattinidae,
Corydiidae Corydiidae, previously known as Polyphagidae, is a family of the order Blattodea ( cockroaches). Many are known as sand cockroaches. The family is divided into five subfamilies, comprising some 40 genera. One prominent species is the desert cockr ...
, Ectobiidae, † Olidae, †
Liberiblattinidae Liberiblattinidae is an extinct family of cockroaches known from the Jurassic to Cretaceous. Some taxa, like '' Cryptoblatta'' and '' Hydrokhoohydra'', are suggested to be semiaquatic. '' Spongistoma'' is suggested to be a nectarivore due to its ...
, † Alienopteridae, †
Manipulatoridae ''Manipulator modificaputis'' is an extinct cockroach which lived during the Upper Cretaceous period. The holotype specimen is fossilized in a 100-million-year-old piece of Burmese amber, which was found in a quarry of volcanoclastic mudstone ( ...
Umenocoleidae ,
Nocticolidae Nocticolidae is a small family in the order Blattodea (cockroaches). It consists of only 32 known species in 9 genera. They are found in Africa, Asia and Australia. Most live in cave habitats, although a few are associated with termites. Cave ada ...
, † Pabuonqedidae Termites († Archeorhinotermitidae, Hodotermitidae, Stolotermitidae, Termitidae and
Mastotermitidae Mastotermitidae is a family of termites with one sole living species, ''Mastotermes darwiniensis'' which is found only in northern Australia. The remaining genera of this family are only known from the fossil record. Fossil record Numerous foss ...
) and mantid '' Burmantis''.


= Mecoptera

= Six families of mecopteran are known, including: Bittacidae, Eomeropidae, Meropeidae, † Orthophlebiidae , † Pseudopolycentropodidae and † Aneuretopsychidae.


= Psocoptera

= Ten families of psocopteran are known, including: †
Archaeatropidae Archaeatropidae is an extinct family of Psocoptera in the suborder Trogiomorpha.A. Baz and V. M. Ortuño. 2000. Archaeatropidae, a new family of Psocoptera from the Cretaceous Amber of Alava, northern Spain. ''Annals of the Entomological Society o ...
,
Compsocidae Compsocidae is a family of Psocodea (formerly Psocoptera) belonging to the suborder Troctomorpha. The family comprises two extant species in two genera, both found in Mesoamerica. ''Compsocus elegans'' is found in Mexico and Central America, wh ...
, †
Cormopsocidae Cormopsocidae is an extinct family of Psocodea. All currently known members are from the mid-Cretaceous Burmese amber of Myanmar. The family is considered either to be the earliest diverging group of the suborder Trogiomorpha, or the sister grou ...
Empheriidae Empheriidae is an extinct family of Psocoptera in the suborder Trogiomorpha. Genera * †'' Burmempheria'' Li et al. 2020 Burmese amber, Myanmar, Cenomanian * †'' Empheria'' Hagen 1856 Baltic amber, Eocene * †'' Empheropsocus'' Baz and Ort ...
, Liposcelididae,
Manicapsocidae Manicapsocidae is a family of Psocodea (formerly Psocoptera). It contains 8 extant species in 4 genera, with most of the species being found in the Neotropics, with one species in the Afrotropics. The extinct family Electrentomidae has been sugg ...
, Pachytroctidae,
Prionoglarididae Prionoglarididae is a family of the order Psocodea (formerly Psocoptera) that are barklice characterized by the reduction or simplification of the lacinia in adults and the specialised form of the male genitalia. It contains the only known genu ...
,
Psyllipsocidae Psyllipsocidae is a family of cave barklice in the order Psocodea. There are about 7 genera and more than 70 described species in Psyllipsocidae. Genera These eight genera belong to the family Psyllipsocidae: * '' Annulipsyllipsocus'' Hakim, Az ...
, Sphaeropsocidae and
Trogiidae Trogiidae is a family (biology), family of granary booklice in the order (biology), order Psocodea (formerly Psocoptera). There are about 11 genera and more than 50 described species in Trogiidae. Genera These 11 genera belong to the family Tro ...
.


= Orthoptera

= Seven families of orthopteran are known, including: †
Elcanidae Elcanidae are an extinct family of Mesozoic and early Cenozoic orthopterans. Members of the family are distinguished by the presence of spurs on the distal part of the metatibia, unique among orthopterans, these have been suggested to have been ...
, Gryllidae, Mogoplistidae,
Ripipterygidae Ripipterygidae is a family of insects in the order Orthoptera. Members of the family are commonly known as mud crickets. Description Ripipterygids are small, often dark-colored, cricket-like orthopterans, between 3 and 14 mm in length. Th ...
, Tetrigidae, Tettigoniidae and
Tridactylidae The Tridactylidae are a family in the insect order Orthoptera. They are small, mole-cricket-like insects, almost always less than long when mature. Generally they are shiny, dark or black, sometimes variegated or sandy-coloured. They commonly ...


= Trichoptera

= Eight families of trichopteran are known, including: Calamoceratidae, †
Dysoneuridae Dysoneuridae is an extinct family of insect in the order Trichoptera, the caddisflies. The family was first described by I.D. Sukacheva (also spelled Sukatsheva) in 1968, and lived from the Middle Jurassic to mid-Cretaceous. In Wichard et al. (20 ...
,
Helicopsychidae Helicopsychidae (snail-case caddisflies) are a family of Trichoptera. The name refers to the helix shaped larval cases and they should not be confused with Limnephilidae which sometimes inhabit the snail shells. Their shells range from and are c ...
, Hydroptilidae, Odontoceridae,
Philopotamidae Philopotamidae is a family of insects in the order Trichoptera, the caddisflies. They are known commonly as the finger-net caddisflies.Polycentropodidae The Polycentropodidae are a family of trumpet-net and tube-making caddisflies. There are at least 30 genera and 720 described species in Polycentropodidae. The type genus for Polycentropodidae is '' Polycentropus'' J. Curtis, 1835. The larvae ...
and
Psychomyiidae The Psychomyiidae are a family of tube-making caddisflies. Members of this family are typically very similar to polycentropodids, most of them can be differentiated by the spur formula is 2-4-4, thyridial cell short, absence of the forewing for ...
.


= Dermaptera

= Five families of dermapteran are known, including:
Anisolabididae Anisolabididae is a family of earwigs, in the suborder Forficulina and the order Dermaptera. It is one of nine families in the suborder Forficulina, and contains thirty-eight genera spread across thirteen subfamilies. Subfamilies The family cont ...
,
Diplatyidae Diplatyidae is a family of earwigs in the suborder Neodermaptera. It contains three subfamilies,Labiduridae Labiduridae, whose members are known commonly as striped earwigs, is a relatively large family of earwigs in the suborder Forficulina.See first entry in external links section for reference. Taxonomy The family contains a total of approximatel ...
and
Pygidicranidae Pygidicranidae is a family of earwigs, formerly placed in the suborder Forficulina, now in the suborder Neodermaptera. The family currently contains twelve subfamilies and twenty six genera. Eight of the subfamilies are monotypic, each containi ...
.


= Embioptera

= Four families of
embiopteran The order Embioptera, commonly known as webspinners or footspinners, are a small group of mostly tropical and subtropical insects, classified under the subclass Pterygota. The order has also been called Embiodea or Embiidina. More than 400 spe ...
are known, including:
Clothodidae Clothodidae is a family of webspinners in the order Embioptera. There are about 5 genera and 18 described species in Clothodidae. Genera These five genera belong to the family Clothodidae: * ''Antipaluria ''Antipaluria'' is a genus of websp ...
, Oligotomidae, Notoligotomidae and † Sorellembiidae.


= Notoptera

= One species of notopteran is known, a nymph ice crawler (
Grylloblattidae Grylloblattidae, commonly known as the icebugs, or ice crawlers, is a family of extremophile (psychrophile) and wingless insects that live in the cold on top of mountains and the edges of glaciers. They belong, along with Mantophasmatidae (rock c ...
) '' Sylvalitoralis cheni.''


= Strepsiptera

= Two families of
strepsipteran The Strepsiptera are an order of insects with eleven extant families that include about 600 described species. They are endoparasites in other insects, such as bees, wasps, leafhoppers, silverfish, and cockroaches. Females of most species neve ...
are known, † Cretostylopidae and † Phthanoxenidae


= Lepidoptera

= Six families of
lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
n are known, including:
Agathiphagidae ''Agathiphaga'' is a genus of moths, known as kauri moths. It is the only living in the family Agathiphagidae. This caddisfly-like lineage of primitive moths was first reported by Lionel Jack Dumbleton in 1952, as a new genus of Micropterigidae ...
,
Douglasiidae Douglasiidae is a small Lepidopteran family including around 28 species of micromoth whose adults are collectively called Douglas moths. The largest genus in the family is ''Tinagma''. They are primarily found in the Palearctic (20 spp.) and Near ...
, Gelechiidae, Gracillariidae,
Lophocoronidae Lophocoronoidea is a superfamily of insects in the order Lepidoptera. There is a single extant genus, ''Lophocorona'', in the family Lophocoronidae. These are small, primitive nocturnal moths restricted to Australia whose biology is largely unkno ...
and Micropterigidae.


= Megaloptera

= One species of
megaloptera Megaloptera is an order of insects. It contains the alderflies, dobsonflies and fishflies, and there are about 300 known species. The order's name comes from Ancient Greek, from ''mega-'' (μέγα-) "large" + ''pteryx'' (πτέρυξ) "wing ...
n is known, '' Haplosialodes liui'' of the family Sialidae.


= Phasmatodea

= Four families of
phasmatodean The Phasmatodea (also known as Phasmida, Phasmatoptera or Spectra) are an order of insects whose members are variously known as stick insects, stick-bugs, walking sticks, stick animals, or bug sticks. They are also occasionally referred to as D ...
are known: † Archipseudophasmatidae,
Phasmatidae The Phasmatidae are a family of the stick insects (order Phasmatodea). They belong to the superfamily Anareolatae of suborder Verophasmatodea. Like many of their relatives, the Phasmatidae are capable of regenerating limbs and commonly reprodu ...
. † Pterophasmatidae and Timematidae


= Thysanoptera

= Five families of thrips are known, including: Aeolothripidae, Melanthripidae, † Rohrthripidae, Thripidae and
Stenurothripidae Stenurothripidae is a family of thrips belonging to the order Thysanoptera Thrips ( order Thysanoptera) are minute (mostly long or less), slender insects with fringed wings and unique asymmetrical mouthparts. Different thrips species feed m ...
.


= Plecoptera

= Two families of stoneflies are known,
Perlidae The Perlidae are a family of stoneflies, with more than 50 genera and 1,100 described species. The majority of the Perlidae are found in eastern North America, but they occur worldwide except for Antarctica and parts of Africa. Their lifecycle ...
and † Petroperlidae.


= Raphidioptera

= One family of Raphidiopteran is known, †
Mesoraphidiidae Mesoraphidiidae is an extinct family (biology), family of snakeflies in the suborder Raphidiomorpha. The family lived from the Late Jurassic through the Late Cretaceous and is known from twenty-five genera. Mesoraphidiids have been found as both ...
.


= †Chresmodidae

= A species of the enigmatic long legged chresmodid insect ''
Chresmoda ''Chresmoda'' is an extinct genus of insects within the family Chresmodidae. Description ''Chresmoda'' are large enigmatic insects with very long specialized legs, probably adapted for skating on the water surface similar to pond skaters. They ...
'' is known.


= †Tarachoptera

= One family of Tarachopteran is known: †
Tarachocelidae Tarachoptera is an extinct order of insects, currently solely known from the mid Cretaceous aged Burmese amber. It belongs to Amphiesmenoptera alongside living Lepidopterans (butterflies and moths) and Trichoptera (caddiesflies), but is outside ...


= †Permopsocida

= One family of Permopsocidan is known: †
Archipsyllidae Permopsocida is an extinct order of insects known from the Early Permian to the Mid-Cretaceous. It is part of Paraneoptera, alongside bark lice (including lice), bugs and thrips. Within Paraneoptera it is considered to be closer to the clade ...


= Zoraptera

= Multiple species of '' Zorotypus'' and the monotypic genus '' Xenozorotypus'' are known.


Nematoda

Five families of
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s are known, including:
Cosmocercidae The Cosmocercidae are a nematode family in the superfamily Cosmoceroidea. Genera Genera within the family Cosmocercidae include: * '' Cosmocercoides'' Wilkie, 1930 * ''Nemhelix'' Morand & Petter, 1986 - with the only species ''Nemhelix bakeri' ...
,
Heterorhabditidae ''Heterorhabditis'' is a genus of nematodes belonging to the order Rhabditida. All species of this genus are obligate parasites of insects, and some are used as biological control agents for the control of pest insects. ''Heterorhabditis'' nemato ...
, Mermithidae,
Thelastomatidae Thelastomatidae is a family of nematodes belonging to the order Oxyurida Oxyurida is an order of nematode worms of the class Secernentea. It consists of four families, one of which contains the human pinworm (''Enterobius vermicularis''). ...
,
Aphelenchoididae Aphelenchoididae is a nematode family in the order Aphelenchida. List of genera Subfamily Anomyctinae * Genus '' Anomyctus'' Subfamily Aphelenchoidinae'' * Genus '' Aphelenchoides'' * Genus '' Ficophagus'' * Genus '' Laimaphelenchus'' * Ge ...


Nematomorpha

One genus of
nematomorph Nematomorpha (sometimes called Gordiacea, and commonly known as horsehair worms, hairsnakes, or Gordian worms) are a phylum of parasitoid animals superficially similar to nematode worms in morphology, hence the name. Most species range in size ...
is known: '' Cretachordodes'' ( Chordodidae, Gordioidea)


Mollusca

Aside from the previously mentioned ammonites and marine gastropod shells, Seven families of terrestrial gastropod are known: Diplommatinidae,
Pupinidae Pupinidae is a taxonomic family of land snails with an operculum, terrestrial gastropod mollusks in the superfamily Cyclophoroidea (according to the taxonomy of the Gastropoda by Bouchet & Rocroi, 2005).MolluscaBase eds. (2021). MolluscaBase ...
, Achatinidae, Punctidae, Valloniidae,
Assimineidae Assimineidae is a family of minute snails, also known as palmleaf snails, with an operculum, gastropod mollusks or micromollusks in the superfamily Rissoidae. Many of these very small snails live in intermediate habitats, being amphibious betwee ...
and Cyclophoridae


Vertebrates

While the deposit is well known for invertebrate inclusions, some vertebrate inclusions have been found as well. One of the more notable discoveries was a well preserved theropod dinosaur tail, with preserved feathers. As well as fossils of
enantiornithine The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and cla ...
birds including juveniles and partial wings and preserved feet, including a diagnostic taxon, ''
Elektorornis ''Elektorornis'' is an extinct genus of enantiornithean bird known from a partial hindlimb and a small amount of wing plumage. It contains a single species, ''Elektorornis chenguangi''. The hindlimb and feathers were preserved in a piece of 99 mi ...
''. A complete skull of the lizard ''
Oculudentavis ''Oculudentavis'' is an extinct genus of lizard of uncertain taxonomic placement,Bolet et al., 2021 originally identified as an avialan dinosaur (bird, in the broad sense). It contains two known species, ''O. khaungraae'' and ''O. naga''. Each ...
'' is known. ''
Electrorana ''Electrorana'' is an extinct genus of frog that lived in what is now Myanmar during the mid-Cretaceous, around 99 million years ago. The type and only species is ''E. limoae''. The generic name is derived from the Latin () and (), while the sp ...
'' is a well preserved frog known from the amber. Other notable specimens include an embryonic snake. Several specimens of lizard have been described from the deposit including a
gecko Geckos are small, mostly carnivorous lizards that have a wide distribution, found on every continent except Antarctica. Belonging to the infraorder Gekkota, geckos are found in warm climates throughout the world. They range from . Geckos ar ...
with preserved toe pads (''
Cretaceogekko ''Cretaceogekko'' (meaning “Cretaceous gecko”) is an extinct monotypic genus of gecko (represented by the type species ''Cretaceogekko burmae'') known from a single partial specimen preserved in Burmese amber from the Cenomanian of Myanmar. T ...
''). and a miniaturised (~2 cm) long possible stem- anguimorph ('' Barlochersaurus'') One of the "lizard" specimens was initially described to be a chamelonid, actually turned out to be an albanerpetontid amphibian. This was described in 2020 as the new genus and species ''
Yaksha perettii ''Yaksha perettii'' is an extinct species of albanerpetontid amphibian, and the only species in the genus ''Yaksha''. It is known from three specimens found in Cenomanian aged Burmese amber from Myanmar. The remains of ''Yaksha perettii'' are the ...
.''


Flora


Angiosperms

Eleven species of
Angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
are known in nine genera, including members of Cornaceae, Cunoniaceae,
Lauraceae Lauraceae, or the laurels, is a plant family that includes the true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genera worldwide (Christenhusz & Byng 2016 ). They are dicotyledons, and occur ma ...
, ? Monimiaceae and Laurales ''incertae sedis''. Poales ''incertae sedis'' and Angiosperm incertae sedis.


Bryopsida

Two genera of Bryopsida in the separate orders
Dicranales Dicranales is an order of haplolepideous mosses in the subclass Dicranidae The Dicranidae are a widespread and diverse subclass of mosses in class Bryopsida, with many species of dry or disturbed areas. They are distinguished by their spores; ...
and Hypnodendrales


Jungermanniopsida

Three families of Jungermanniopsida are known, Frullaniaceae, Lepidolaeanaceae,
Radulaceae ''Radula'' is a genus of liverwort, and is the only genus in family Radulaceae. It is a leafy liverwort. The appearance of the plants are as a scaly, green surface on the trunk of a tree, log or rock in a sheltered, moist out-door environment. ...
.


Pinophyta

Two families of Pinopsida are known: Araucariaceae and
Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdio ...
including '' Metasequoia''.


Pteridopsida

Five families of Pteridopsidan are known: Cystodiaceae, Dennstaedtiaceae,
Lindsaeaceae Lindsaeaceae is a pantropical family of ferns in the order Polypodiales. It contains six or seven genera with about 220 known species, some of which also extend into the more temperate regions of eastern Asia, New Zealand, and South America.
, Pteridaceae, Thyrsopteridaceae, and several genera of Polypodiales ''incertae sedis.''


Amoebozoa


Myxogastria

Sporocarps of extant
myxogastrid Myxogastria/Myxogastrea (myxogastrids, ICZN) or Myxomycetes ( ICN), is a class of slime molds that contains 5 orders, 14 families, 62 genera, and 888 species. They are colloquially known as the ''plasmodial'' or ''acellular' ...
slime mould genus ''
Stemonitis ''Stemonitis'' is a distinctive genus of slime moulds found throughout the world (except Antarctica). They are characterised by the tall brown sporangia, supported on slender stalks, which grow in clusters on rotting wood. The genus was first des ...
'' are known.


Dictyostelia

A possible dictyostelid '' Paleoplastes burmanica'' has been described.


History

The amber is apparently referred to in ancient Chinese sources as originating from Yunnan Province as early as the first century AD according to the ''
Book of the Later Han The ''Book of the Later Han'', also known as the ''History of the Later Han'' and by its Chinese name ''Hou Hanshu'' (), is one of the Twenty-Four Histories and covers the history of the Han dynasty from 6 to 189 CE, a period known as the Later ...
'' and trade with China had been ongoing for centuries. This has been confirmed by Fourier-transform infrared spectroscopy (FTIR) analysis of amber artifacts from the Eastern Han Dynasty (25 - 220 CE). It was first mentioned in European sources by the
Jesuit , image = Ihs-logo.svg , image_size = 175px , caption = ChristogramOfficial seal of the Jesuits , abbreviation = SJ , nickname = Jesuits , formation = , founders ...
Priest Álvaro Semedo who visited China in 1613, it was described as being "digged out of mines, and sometimes in great pieces, it is redder than our amber though not so cleane".Zherikhin, V.V., Ross, A.J., 2000
A review of the history, geology and age of Burmese amber (Burmite).
Bulletin of the Natural History Museum, London (Geology) 56 (1), 3–10.
The locality itself has been known to European explorers since the 1800s with visitation to the Hukawng Valley by Simon Fraser Hannay in 1836–1837. At that time the principle products of the valley mines were salt, gold, and amber, with the majority of gold and amber being bought by Chinese traders. Hannay visited the amber mines themselves on March 21, 1836, and he noted that the last three miles to the mines were marked with numerous abandoned pits, up to in depth, where amber had been dug in the past. The mining had moved over the hill to a series of 10 pits but no visible amber was seen, suggesting that miners possibly hid the amber found that day before the party arrived. Mining was being performed manually at the time through the use of sharpened bamboo rods and small wooden shovels. Finer pieces of amber were recovered from the deeper pits, with clear yellow being recovered from depths of The recovered amber was bought with silver or often exchanged for jackets, hats, copper pots, or
opium Opium (or poppy tears, scientific name: ''Lachryma papaveris'') is dried latex obtained from the seed capsules of the opium poppy ''Papaver somniferum''. Approximately 12 percent of opium is made up of the analgesic alkaloid morphine, which i ...
among other goods. mixed and lower quality amber was sold from around ticals to 4 rupees per Seer (unit), seer. Pieces that were considered high quality or fit or use as ornamentation were described as expensive and price varied depending on clarity and color. Women of the valley were noted to wear amber earrings as part of their jewelry. In 1885 the Konbaung dynasty was annexed to the British Raj and a survey of the area was conducted by Dr. Fritz Noetling on behalf of the Geological Survey of India. The final research before Burmese independence in 1947 was conducted by Dr. H.L. Chhibber in 1934, who provided the most detailed description of Burmite occurrences.


History of research

The first research on the inclusions in the Burmese amber was published in 1916 by Theodore Dru Alison Cockerell, who initially concluded that the amber was Miocene in age. However, he subsequently noted the archaic nature of the insects, and concluded that the amber must be older.


Modern exploitation and controversy

Leeward Capital Corp, a small Canadian mining firm, controlled the deposit from the mid-1990s to c. 2000, though the history of exploitation during the 2000s is obscure. The Kachin Independence Army (KIA), an armed rebel group seeking to secede Kachin province from Myanmar, controlled the area during the early to mid 2010's. During the early 2010s, production rapidly increased. The working conditions at the mines have been described as extremely unsafe, down deep pits barely wide enough to crawl through, with no accident compensation. The KIA controlled amber export via numerous licenses, taxes, restrictions on movement of labor, and enforced auctions. The main amber market in Myanmar is Myitkyina. Most amber is smuggled into China, primarily for jewelry, with estimates of around 100 tonnes passing through to the main market of Tengchong, Yunnan in 2015, with an estimated value between five and seven billion Yuan (currency), yuan. Burmese amber was estimated to make up 30% of Tenchong's gemstone market (the rest being Jade trade in Myanmar, Myanmar Jade), and was declared one of the cities eight main industries by the local government. The presence of calcite veins are a major factor in determining the gem quality of pieces, with pieces with a large number of veins having significantly lower value. In June 2017 the Tatmadaw seized control of the mines from the KIA. Sales of amber were alleged to help fund the Kachin conflict by various news organisations in 2019. Interest in this discussion rose in March 2020 after the highly publicised description of ''
Oculudentavis ''Oculudentavis'' is an extinct genus of lizard of uncertain taxonomic placement,Bolet et al., 2021 originally identified as an avialan dinosaur (bird, in the broad sense). It contains two known species, ''O. khaungraae'' and ''O. naga''. Each ...
,'' which made the cover of Nature (journal), ''Nature''. On April 21, 2020, the Society of Vertebrate Paleontology (SVP) published a letter of recommendation to journal editors asking for “a moratorium on publication for any fossil specimens purchased from sources in Myanmar after June 2017 when the Myanmar military began its campaign to seize control of the amber mining”. On April 23, 2020 ''Acta Palaeontologica Polonica'' stated that it would not accept papers on Burmese amber material collected from 2017 onwards, after the Burmese military took control of the deposit, requiring "certification or other demonstrable evidence, that they were acquired before the date both legally and ethically". On May 13, 2020, the ''Journal of Systematic Palaeontology'' published an editorial stating that it would no longer consider papers based whole or in part on Burmese amber material, regardless of whether in historic collections or not. On 30 June 2020, a statement from the International Palaeoentomological Society was published in response to the SVP, criticising the proposal to ban publishing on Burmese amber material. In August 2020, a comment from over 50 authors was published in ''PalZ'' responding to the SVP statement. The authors disagreed with the proposal of a moratorium, describing the focus on the Burmese amber as "arbitrary" and that "The SVP’s recommendation for a moratorium on Burmese amber affects fossil non-vertebrate research much more than fossil vertebrate research and clearly does not represent this part of the palaeontological community." The conclusion that Burmese amber funded the Tatmadaw was disputed by George Poinar and Sieghard Ellenberger, who found that the supply of amber collapsed after the 2017 takeover of the mines by the Tatmandaw, and that most of the current circulation of amber in Chinese markets was extracted prior to 2017. A story in ''Science'' in 2019 stated: "Two former mine owners, speaking through an interpreter in phone interviews, say taxes have been even steeper since government troops took control of the area. Both shut their mines when they became unprofitable after the government takeover, and almost all deep mines are now out of business, dealers here corroborate. Only shallow mines and perhaps a few secret operations are still running." There were around 200,000 miners working in the Hukawng valley mines prior to the takeover by the Tatmadaw, which shrunk to 20,000 or less after the military operations. Adolf Peretti, a gemologist who owns a museum with Burmese amber specimens, noted that the 2017 cutoff suggested by the SVP does not take into account that the export of Burmese amber prior to 2017 was also funding internal conflict in Myanmar due to the control by the KIA. Much of the amber cutting since 2017 has been done in internally displaced person camps, under humanitarian and non-conflict conditions.


Other Myanmar ambers

Other deposits of amber are known from several regions in Myanmar, with noted deposits in the Shwebo District of the Sagaing Region, from the Pakokku District, Pakokku and Thayet District, Thayet districts of Magway Region and the Bago District of the Bago Region. Unlike the Hukawng deposit, none of these sources have produce notable quantities of amber.


Tilin amber

A 2018 study on an amber deposit from Htilin, Tilin in central Myanmar indicated that deposit to be 27 million years younger than the Hukawng deposit, dating to approximately 72 million years old, placing it in the latest Campanian age. The deposit was associated with an overlying tuffaceous layer, and underlying nodules of brown sandstone yielded remains of the ammonite ''Sphenodiscus.'' Within a number of arthropod specimens were described though much more poorly preserved than specimens in the Hukawng amber. These include members of Hymenoptera ( Braconidae, Diapriidae, Scelioninae, Scelionidae) Diptera ( Ceratopogonidae, Chironomidae) Dictyoptera (Cockroach, Blattaria, Mantodea) planthoppers, Berothidae and bark lice (Lepidopsocidae) as well as extant ant subfamilies Dolichoderinae and tentatively Ponerinae, as well as fragments of moss.


Hkamti amber

The Hkamti site is located ca. 90 km southwest of the Angbamo site and predominantly consists of limestone, interbedded with mudstone and tuff, the amber is found within the unconsolidated mudstone/tuff layers. A crinoid was found attached to one amber specimen, alongside marine plant remains in the surrounding sediment, indicating deposition in a shallow marine setting. The amber is generally red-brown, and yellow colouration is rare, the amber is generally found as angular clasts, indicating short transport distance and is more brittle than other northern Myanmar ambers. Zircon dating has constrained the age of the deposit to the early Albian, c. 110 Ma, significantly older than the dates obtained from other deposits. Fauna found within the amber includes: Archaeognatha, Millipede, Diplopoda, Beetle, Coleoptera, Spider, Araneae, Trichoptera, Neuroptera, Psocodea, Isoptera Diptera, Orthoptera, Pseudoscorpionida, Hymenoptera and Thysanoptera.


See also

*


References

{{Reflist


External links


Blood Amber Military resource grab clears out indigenous peoples in Kachin State’s Hugawng Valley ''Kachin Development Networking Group''
Burmese amber, Paleontological sites of Asia Paleontology in Myanmar