![Burmalindenia imperfecta whole amber](https://upload.wikimedia.org/wikipedia/commons/c/c0/Burmalindenia_imperfecta_whole_amber.png)
Burmese amber, also known as Burmite or Kachin amber, is
amber
Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In M ...
from the
Hukawng Valley
The Hukawng Valley ( my, ဟူးကောင်းချိုင့်ဝှမ်း; also spelt Hukaung Valley) is an isolated valley in Myanmar, roughly in area. It is located in Tanaing Township in the Myitkyina District of Kachin State ...
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 Wells explai ...
. The amber is dated to around 100 million years ago, during the latest
Albian
The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...
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 ...
ages of the mid-
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
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
![Burmese amber geological context PNAS Fig1 A (cropped)](https://upload.wikimedia.org/wikipedia/commons/f/f9/Burmese_amber_geological_context_PNAS_Fig1_A_%28cropped%29.jpg)
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 configu ...
sedimentary basin
Sedimentary basins are region-scale depressions of the Earth's crust where subsidence has occurred and a thick sequence of sediments have accumulated to form a large three-dimensional body of sedimentary rock. They form when long-term subsiden ...
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
In geology, a terrane (; in full, a tectonostratigraphic terrane) is a crust fragment formed on a tectonic plate (or broken off from it) and accreted or " sutured" to crust lying on another plate. The crustal block or fragment preserves its ow ...
, 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 sta ...
during at least the Early
Paleozoic
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ...
, 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, wh ...
to Early Cretaceous. It is also disputed whether the block had
accreted onto the Asian
continental margin
A continental margin is the outer edge of continental crust abutting oceanic crust under coastal waters. It is one of the three major zones of the ocean floor, the other two being deep-ocean basins and mid-ocean ridges. The 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
Paleomagnetism (or palaeomagnetismsee ), is the study of magnetic fields recorded in rocks, sediment, or archeological materials. Geophysicists who specialize in paleomagnetism are called ''paleomagnetists.''
Certain magnetic minerals in rock ...
reconstruction finds that the Burma Terrane formed an island land mass in the
Tethys Ocean during the Mid Cretaceous at a
latitude
In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north ...
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
Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks.
Most sandstone is composed of quartz or feldspar (both silicat ...
, with beds of
silt
Silt is granular material of a size between sand and clay and composed mostly of broken grains of quartz. Silt may occur as a soil (often mixed with sand or clay) or as sediment mixed in suspension with water. Silt usually has a floury feel ...
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, especiall ...
and laterally persistent thin (1–2 mm thick)
coal
Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen.
Coal is formed when ...
horizons. Massive
micritic limestone
Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms w ...
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
** ...
, 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 cuttlefis ...
''
Mortoniceras'' has been found in a sandstone bed 2 metres above the amber horizon, alongside indeterminate gastropods and bivalves.
Lead-uranium dating of
zircon
Zircon () is a mineral belonging to the group of nesosilicates and is a source of the metal zirconium. Its chemical name is zirconium(IV) silicate, and its corresponding chemical formula is Zr SiO4. An empirical formula showing some of t ...
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 ...
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 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
Danai ( my, တနိုင်းမြို့) is a town in Kachin State, in the northernmost part of Myanmar.
Kachin State ( Kachin: ''Wunpawng Mungdan''; Burmese: ကချင်ပြည်နယ်) is the northernmost state of Myanmar. ...
. The Hkamti site SW of the Hukawng basin has been determined to be significantly older, dating to the early
Albian
The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...
around ca. 110 Ma and is therefore considered distinct.
Paleoenvironment
![Puzosia Bhimaites species Burmese amber PNAS Fig2 A](https://upload.wikimedia.org/wikipedia/commons/a/ad/Puzosia_Bhimaites_species_Burmese_amber_PNAS_Fig2_A.jpg)
The Burmese amber paleoforest is considered to have been a
tropical rainforest
Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equator ...
, 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 shell
The gastropod shell is part of the body of a Gastropoda, gastropod or snail, a kind of mollusc. The shell is an exoskeleton, which protects from predators, mechanical damage, and dehydration, but also serves for muscle attachment and calcium s ...
s (including ''
Mathilda'') and
littoral or supralittoral isopods entombed in a piece of amber with shell sand,
along with growth of
Isocrinid crinoids,
coral
Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and se ...
s and
oyster
Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
s 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 environmen ...
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
In geology, a bed is a layer of sediment, sedimentary rock, or pyroclastic material "bounded above and below by more or less well-defined bedding surfaces".Neuendorf, K.K.E., J.P. Mehl, Jr., and J.A. Jackson, eds., 2005. ''Glossary of Geology'' ...
, 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 mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratc ...
veins
Veins are blood vessels in humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated ...
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 ext ...
origin, with a likely
araucarian source tree, based on spectroscopic analysis and wood fragment inclusions, though a
pine
A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
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 ...
, 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
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s. 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.
Invertebrates
Well over 1000 species of invertebrates are known from the deposit, including, notably the oldest members of
Palpigradi
Palpigrades, commonly known as microwhip scorpions, are arachnids belonging to the order Palpigradi.
Description
Palpigrades belong to the arachnid class. They are the sister group to Solifugae, no more than in length, and averaging . They have ...
(''
Electrokoenenia'')
and
Schizomida (''
Mesozomus'') the oldest
Velvet worm
Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...
(''
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 mo ...
'') and the only known fossil members of
Mesothelae
The Mesothelae are a suborder of spiders (order Araneae) that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral ...
and
Ricinulei
Ricinulei is a small order of arachnids. Like most arachnids, they are predatory, eating small arthropods. They occur today in west-central Africa ('' Ricinoides'') and the Neotropics ('' Cryptocellus'' and '' Pseudocellus'') as far north as Texa ...
since the Paleozoic. ''
Chimerarachne
''Chimerarachne'' is a genus of extinct arachnids, sometimes considered as spider itself,Dunlop, J. A., Penney, D. & Jekel, D. 2020A summary list of fossil spiders and their relatives In World Spider Catalog. Natural History Museum Bern, online ...
'' is a unique stem spider still possessing a tail, with similar forms only known from the Paleozoic.
Arachnids
= Araneae
=
Forty-four families of
spiders are known from the Burmese amber, including:
Archaeidae, †
Burmadictynidae, †
Burmascutidae, †
Burmathelidae,
Clubionidae
The sac spiders of the family Clubionidae have a very confusing taxonomic history. Once, this family was a large catch-all taxon for a disparate collection of spiders, similar only in that they had eight eyes arranged in two rows and conical anter ...
,
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,
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 ano ...
,
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 o ...
,
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,
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
Lynx spider (Oxyopidae) is a family of araneomorph spiders first described by Tamerlan Thorell in 1870. Most species make little use of webs, instead spending their lives as hunting spiders on plants. Many species frequent flowers in particular, ...
,
Palpimanidae, †
Parvithelidae,
Pholcidae, †
Pholcochyroceridae, †
Plumorsolidae, †
Praearaneidae, †
Praeterleptonetidae,
Psechridae
Psechridae is a family of 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 belongs to the RTA clade of spiders ...
,
Psilodercidae,
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,
Tetrablemmidae
Tetrablemmidae, sometimes called armored spiders, is a family of tropical araneomorph spiders first described by Octavius Pickard-Cambridge in 1873. It contains 126 described species in 29 genera from southeast Asia, with a few that occur in Afri ...
,
Tetragnathidae,
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
Theridiidae, also known as the tangle-web spiders, cobweb spiders and comb-footed spiders, is a large family of araneomorph spiders first described by Carl Jakob Sundevall in 1833. This diverse, globally distributed family includes over 3,000 sp ...
,
Thomisidae
The Thomisidae are a family of spiders, including about 170 genera and over 2,100 species. The common name crab spider is often linked to species in this family, but is also applied loosely to many other families of spiders. Many members of th ...
,
Uloboridae and †
Vetiaroridae.
= Acariformes
=
Twenty families of
acariformes are known from the Burmese amber, including:
Anystidae,
Archaeorchestidae,
Bdellidae,
Caeculidae,
Cheyletidae,
Enantioppiidae,
Eremaeidae,
Erythraeidae,
Eupodidae,
Gymnodamaeidae,
Malaconothridae,
Microtrombidiidae,
Neoliodidae,
Oribatellidae,
Oribotritiidae,
Resinacaridae,
Smarididae
Smarididae is a family of mites belonging to the order Trombidiformes. These large predatory mites have long oval bodies, distinctively pointed in front. They are usually red and densely hairy with slender legs, sometimes very long. They have e ...
,
Trombellidae,
Trombidiidae
Trombidiidae, also known as red velvet mites, true velvet mites, or rain bugs, are small arachnids (eight-legged arthropods) found in plant litter and are known for their bright red color.
While adults are typically in length, some, such as t ...
and
Tuckerellidae.
= Opiliones
=
Nine families of
opiliones are known from the Burmese amber, including:
Beloniscidae,
Epedanidae, †
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 ...
and
Tithaeidae.
= Pseudoscorpiones
=
![Procheiridium judsoni holotype Fig1 D](https://upload.wikimedia.org/wikipedia/commons/c/c4/Procheiridium_judsoni_holotype_Fig1_D.png)
Twelve families of
pseudoscorpions are known from the Burmese amber, including:
Atemnidae,
Cheiridiidae,
Cheliferidae,
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'' Max Bei ...
,
Garypinidae,
Hyidae Ideoroncidae,
Neobisiidae,
Pseudocheiridiidae and
Withiidae.
= Scorpiones
=
![Betaburmesebuthus bellus Fig1 B](https://upload.wikimedia.org/wikipedia/commons/c/ca/Betaburmesebuthus_bellus_Fig1_B.jpg)
Seven families of
scorpions are known from the Burmese amber, including:
Buthidae,
Chaerilidae
Chaerilidae is a family of scorpion
Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic fo ...
, †
Chaerilobuthidae, †
Palaeoburmesebuthidae, †
Palaeoeuscorpiidae, †
Palaeotrilineatidae and †
Sucinolourencoidae.
= Parasitiformes
=
Seven families of
parasitiformes are known from the Burmese amber, including:
Argasidae, †
Deinocrotonidae, †
Khimairidae,
Ixodidae,
Opilioacaridae
Opilioacaridae is the sole family of mites in the order Opilioacarida, made up of about 13 genera. The mites of this family are rare, large (1.5 to 2.5 mm) mites, and are widely considered primitive, as they retain six pairs of eyes, and ab ...
,
Polyaspididae 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
Burmese amber, also known as Burmite or Kachin amber, is ...
.
= Schizomida
=
One family of
schizomida is known from the Burmese amber:
Hubbardiidae
= Palpigradi
=
One genus of
palpigradi
Palpigrades, commonly known as microwhip scorpions, are arachnids belonging to the order Palpigradi.
Description
Palpigrades belong to the arachnid class. They are the sister group to Solifugae, no more than in length, and averaging . They have ...
is known: ''
Electrokoenenia'', 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 is known: ''
Cushingia'', which does not belong to any extant family.
= Thelyphonida
=
Two genera of
whip scorpion are known: ''
Mesothelyphonus'', which belongs to
Thelyphonidae and ''
Burmathelyphonia,'' which does not belong to any extant family.
= Ricinulei
=
Three genera of
Ricinulei
Ricinulei is a small order of arachnids. Like most arachnids, they are predatory, eating small arthropods. They occur today in west-central Africa ('' Ricinoides'') and the Neotropics ('' Cryptocellus'' and '' Pseudocellus'') as far north as Texa ...
are known: ''
Hirsutisoma'', ?''
Poliochera'' (an otherwise Carboniferous taxon) and ''
Primoricinuleus,'' none of which belong to extant families
Myriapoda
![Burmanopetalum inexpectatum female holotype](https://upload.wikimedia.org/wikipedia/commons/3/32/Burmanopetalum_inexpectatum_female_holotype.png)
Sixteen families of
Myriapods are known, including:
Anthroleucosomatidae, †
Electrocambalidae,
Tingupidae,
Glomeridellidae,
Andrognathidae
Andrognathidae is a family of millipede in the order Platydesmida
Platydesmida (Greek for ''platy'' "flat" and ''desmos'' "bond") is an order of millipedes containing two families and over 60 species. Some species practice paternal care, i ...
,
Paradoxosomatidae,
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,
Polyzoniidae,
Siphoniulidae,
Siphonophoridae
Siphonophoridae is a family of millipede in the order Siphonophorida
Siphonophorida (Greek for "tube bearer") is an order of millipedes containing two families and over 100 species.
Description
Millipedes in the order Siphonophorida are lo ...
,
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,
Cambalidae,
Scolopendrellidae and †
Burmanopetalidae.
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 place ...
,
Tomoceridae,
Neanuridae
The family Neanuridae contains pudgy short-legged springtails of the order Poduromorpha. It was established by Carl Börner in 1901.
Systematics
There are six subfamilies currently recognized:
* Caputanurininae
* Frieseinae
* Morulinina ...
,
Odontellidae 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
archaeognatha
The Archaeognatha are an order of apterygotes, known by various common names such as jumping bristletails. Among extant insect taxa they are some of the most evolutionarily primitive; they appeared in the Middle Devonian period at about the sa ...
ns are known from the Burmese amber:
Machilidae
The Machilidae are a family of insects belonging to the order Archaeognatha (the bristletails). There are around 250 described species worldwide. These insects are wingless, elongated and more or less cylindrical with a distinctive humped thorax ...
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 la ...
= Zygentoma
=
One family of
Zygentoma
Zygentoma are an order in the class Insecta, and consist of about 550 known species. The Zygentoma include the so-called silverfish or fishmoths, and the firebrats. A conspicuous feature of the order are the three long caudal filaments. The tw ...
n is known:
Lepismatidae
= Ephemeroptera
=
![Vetuformosa buckleyi holotype](https://upload.wikimedia.org/wikipedia/commons/9/9a/Vetuformosa_buckleyi_holotype.jpg)
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 orde ...
are known: †
Australiphemeridae,
Baetidae,
Ephemeridae
Ephemeridae is a family of mayflies with about 150 described species found throughout the world except Australia and Oceania. These are generally quite large mayflies (up to 35 mm) with either two or three very long tails. Many species hav ...
,
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
The Aeshnidae, also called aeshnids, hawkers, or darners, is a family of dragonflies. The family includes the largest dragonflies found in North America and Europe and among the largest dragonflies on the planet.
Description
Common worldwide o ...
, †
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. E ...
,
Lindeniidae,
Megapodagrionidae, †
Mesomegaloprepidae, †
Paracoryphagrionidae,
Perilestidae,
Platycnemididae,
Platystictidae.
= Hymenopterans
=
![Diversinitus attenboroughi-1 (cropped)](https://upload.wikimedia.org/wikipedia/commons/9/94/Diversinitus_attenboroughi-1_%28cropped%29.png)
Over fifty families of
hymenopteran
Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic.
Females typica ...
s have been described beginning with the papers of
Cockerell who described a group of
Bethylidae and
Aulacidae species between 1917 and 1920. The monotypic family Melittosphecidae is only known from the Burmese amber species ''
Melittosphex burmensis'' and eight species belonging to ''
Aptenoperissus'' of the monotypic family Aptenoperissidae are also known. Originally described as an
Aneuretinae
Aneuretinae is a subfamily of ants consisting of a single extant species, ''Aneuretus simoni'' ( Sri Lankan relict ant), and 9 fossil species. Earlier, the phylogenetic position of ''A. simoni'' was thought to be intermediate between primitive ...
ant ''
Burmomyrma rossi'' was moved to the extinct
Chrysidoidea
The superfamily Chrysidoidea is a very large cosmopolitan group (some 6,000 described species, and many more undescribed) , all of which are parasitoids or cleptoparasites of other insects. There are three large, common families (Bethylidae, Chr ...
family
Falsiformicidae. A number of Formicidae species known, belonging to ''
Baikuris'' (indet) ''
Camelomecia janovitzi,
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
''Haidomyrmex'' is an extinct genus of ants in the formicid subfamily Haidomyrmecinae, and is one of nine genera placed in the subfamily Haidomyrmecinae. The genus contains three described species ''Haidomyrmex cerberus'', ''Haidomyrmex scimit ...
'', ''
Linguamyrmex vladi'', 2 species of ''
Zigrasimecia
''Zigrasimecia'' is an extinct genus of ants which existed in the Cretaceous period approximately 98 million years ago. The first specimens were collected from Burmese amber in Kachin State, west of Myitkyina town in Myanmar. In 2013, palaeo ...
'', ''
Dhagnathos autokrator'', ''
Chonidris insolita'', ''
Aquilomyrmex huangi'', ''
Protoceratomyrmex revelatus'' and ''
Linguamyrmex brevicornis''. Other families include
Ampulicidae,
Braconidae
The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
,
Cephidae
Cephidae is a family of stem sawflies in the order Hymenoptera. There are about 27 genera and more than 160 described species in Cephidae.
Genera
These 27 genera belong to the family Cephidae:
* '' Athetocephus'' Benson, 1935
* '' Australcep ...
,
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
The Chalcididae are a moderate-sized family within the Chalcidoidea, composed mostly of parasitoids and a few hyperparasitoids. The family is apparently polyphyletic, though the different subfamilies may each be monophyletic, and some may be elev ...
,
Chrysididae
Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group (over 3000 described species) of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic colors ...
,
Crabronidae
The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over 9 ...
,
Diapriidae
The Diapriidae are a family of parasitoid wasps. These tiny insects have an average length of 2–4 mm and never exceed 8 mm. They typically attack larvae and pupae of a wide range of insects, especially flies. The about 2,300 described ...
,
Dryinidae
Dryinidae is a cosmopolitan family of solitary wasps. Its name comes from the Greek ''drys'' for oak: Latreille named the type genus ''Dryinus'' because the first species was collected in an oak plant in Spain. The larvae are parasitoids of the ...
,
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
Evaniidae is a family of parasitoid wasps also known as ensign wasps, nightshade wasps, hatchet wasps, or cockroach egg parasitoid wasps. They number around 20 extant genera containing over 400 described species, and are found all over the world ...
,
Gasteruptiidae,
Geoscelionidae,
Heloridae,
Ichneumonidae
The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25,000 species cur ...
,
Megalyridae,
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
The Mymaridae, commonly known as fairyflies or fairy wasps, are a family of chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species.
Fairyflies are very tiny insects ...
,
Mymarommatidae
The Mymarommatidae, sometimes referred to as false fairy wasps. are a very small family of microscopic parasitic wasps. Only about half of the known species are living taxa (the others are fossils), but they are found worldwide.Gibson, G.A.P.; R ...
,
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
The hymenopteran family Platygastridae (sometimes incorrectly spelled Platygasteridae) is a large group (over 4000 species) of exclusively parasitoid wasps, mostly very small (1–2 mm), black, and shining, with geniculate (elbowed) anten ...
,
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,
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,
Scolebythidae,
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
The Sierolomorphidae are a family of 13 extant species of wasps, in the genera ''Sierolomorpha'' and ''Proscleroderma'', mostly found in the Northern Hemisphere. They are rare and very little is known of their biology. A fossil species ''Loreisom ...
,
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,
Stephanidae,
Tiphiidae
The Tiphiidae (also known as the tiphiid wasps) are a family of large, solitary wasps whose larvae are parasitoids of various beetle larvae, especially those in the superfamily Scarabaeoidea. Until recently, this family contained several addition ...
,
Vespidae
The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
,
Xiphydriidae
Xiphydriidae are a family of wood wasps that includes around 150 species. They are located all over the world including North and South America, Australia, Europe, and others. Xiphydriidae larvae are wood borers in dead trees or branches of a ran ...
, †
Angarosphecidae
Angarosphecidae is an extinct family of Mesozoic and early Cenozoic wasps in the superfamily Apoidea.
Genera
* †'' Angarosphex'' Rasnitsyn 1975 Weald Clay, United Kingdom, Hauterivian/Barremian La Pedrera de Rúbies Formation, Spain, Barrem ...
, †
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, †
Burmorussidae †
Burmusculidae, †
Chrysobythidae, †
Dipterommatidae, †
Diversinitidae, †
Falsiformicidae, †
Gallorommatidae
The Gallorommatidae is an extinct family of microscopic parasitoid wasps, belonging to the Mymarommatoidea. It is known from several species found in Cretaceous aged amber.Engel, M.S.; Grimaldi, D.A. (2007) New false fairy wasps in Cretaceous amb ...
, †
Ohlhoffiidae, †
Panguidae, †
Plumalexiidae, †
Maimetshidae, †
Myanmarinidae, †
Othniodellithidae, †
Peleserphidae, †
Praeaulacidae, †
Proterosceliopsidae, †
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, †
Syspastoxyelidae and several ''incertae sedis'' taxa.
= Dipterans
=
Over forty families of
dipterans
Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced m ...
are known from the Burmese amber, including:
Acroceridae
The Acroceridae are a small family of odd-looking flies. They have a hump-backed appearance with a strikingly small head, generally with a long proboscis for accessing nectar. They are rare and not widely known. The most frequently applied common ...
,
Anisopodidae
The Anisopodidae are a small cosmopolitan family of gnat-like flies known as wood gnats or window-gnats, with 154 described extant species in 15 genera, and several described fossil taxa. Some species are saprophagous or fungivorous. They are m ...
,
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
The Asilidae are the robber fly family, also called assassin flies. They are powerfully built, bristly flies with a short, stout proboscis enclosing the sharp, sucking hypopharynx. The name "robber flies" reflects their notoriously aggressive pre ...
,
Atelestidae,
Blephariceridae,
Bombyliidae
The Bombyliidae are a family of flies, commonly known as bee flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae are mostly parasitoids of other insects.
Overview
The Bombyliidae are a large family of fl ...
,
Cecidomyiidae
Cecidomyiidae is a family of flies known as gall midges or gall gnats. As the name implies, the larvae of most gall midges feed within plant tissue, creating abnormal plant growths called galls. Cecidomyiidae are very fragile small insects us ...
,
Ceratopogonidae
Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally in length. The family includes more than 5,000 species, distributed worldwide, apart from the Antarctic and the Arctic.
Ceratopogonidae are holomet ...
,
Chaoboridae
Chaoboridae, commonly known as phantom midges or glassworms, is a family of fairly common midges with a cosmopolitan distribution. They are closely related to the Corethrellidae and Chironomidae; the adults are differentiated through peculiarit ...
,
Chironomidae
The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many specie ...
,
Corethrellidae
Corethrellidae are a family of biting midges, small flying insects belonging to the order Diptera, females of which feed on the blood of frogs. The members of the family are sometimes known as frog-biting midges. The family currently consists of ...
,
Culicidae
Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning "gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "litt ...
,
Diadocidiidae
The Diadocidiidae are a family of flies (Diptera), containing one extant genus with over 20 species and one extinct genus. Diadocidiidae are found worldwide, except in Africa and Antarctica. They are usually considered close to the Keroplatidae, B ...
,
Dolichopodidae
Dolichopodidae, the long-legged flies, are a large, cosmopolitan family of true flies with more than 7,000 described species in about 230 genera. The genus ''Dolichopus'' is the most speciose, with some 600 species.
Dolichopodidae generally are ...
,
Empididae
__NOTOC__
Empididae is a family of flies with over 3,000 described species occurring worldwide in all the biogeographic realms but the majority are found in the Holarctic. They are mainly predatory flies like most of their relatives in the Empid ...
,
Hybotidae,
Ironomyiidae,
Keroplatidae
The Keroplatidae are a family of small fly, flies known as fungus gnats. About 950 species are described, but the true number of species is undoubtedly much higher. They are generally forest dwellers found in the damp habitats favoured by their h ...
,
Limoniidae,
Lygistorrhinidae
Lygistorrhinidae is a family of long-beaked fungus gnats in the order Diptera. There are about 7 genera and at least 30 described species in Lygistorrhinidae.
Genera
*†'' Archaeognoriste'' Blagoderov & Grimaldi, 2004
*'' Asiorrhina'' Bl ...
,
Mycetophilidae
The Mycetophilidae are a family of small flies, forming the bulk of those species known as fungus gnats. About 3000 described species are placed in 150 genera, but the true number of species is undoubtedly much higher. They are generally found i ...
,
Mythicomyiidae
Mythicomyiidae, commonly called mythicomyiids, are very tiny flies (0.5–5.0 mm) found throughout most parts of the world, especially desert and semi-desert regions, except the highest altitudes and latitudes. They are not as common in the t ...
,
Nemestrinidae
Nemestrinidae, or tangle-veined flies is a family of flies in the superfamily Nemestrinoidea, closely related to Acroceridae. The family is small but distributed worldwide, with about 300 species in 34 genera. Larvae are endoparasitoids of eith ...
,
Phoridae
The Phoridae are a family of small, hump-backed flies resembling fruit flies. Phorid flies can often be identified by their escape habit of running rapidly across a surface rather than taking to the wing. This behaviour is a source of one of thei ...
,
Pipunculidae
Pipunculidae is a family of flies (Diptera) commonly termed big-headed flies, a reference to the large ( holoptic) eyes, which cover nearly the entire head. The family is found worldwide and more than 1300 species have been described.
The larva ...
,
Platypezidae
Platypezidae is a family of true flies of the superfamily Platypezoidea. The more than 250 species are found worldwide primarily in woodland habitats. A common name is flat-footed flies, but this is also used for the closely related Opetii ...
,
Psychodidae
Psychodidae, called drain flies, sink flies, filter flies, sewer flies, or sewer gnats, is a family of Fly, true flies. Some genera have short, hairy bodies and wings giving them a "furry" moth-like appearance, hence one of their common names, m ...
,
Ptychopteridae
The Ptychopteridae, phantom crane flies, are a small family (three extant genera) of nematocerous Diptera. Superficially similar in appearance to other "tipuloid" families, they lack the ocelli of the Trichoceridae, the five-branched radial vein ...
,
Rachiceridae,
Rhagionidae
Rhagionidae or snipe flies are a small family of flies.
They get their name from the similarity of their often prominent proboscis that looks like the beak of a snipe.
Description
Rhagionidae are medium-sized to large flies with slender bodies a ...
,
Scatopsidae
__NOTOC__
The minute black scavenger flies or "dung midges", are a family, Scatopsidae, of nematoceran flies. Despite being distributed throughout the world, they form a small family with only around 250 described species in 27 genera, although m ...
,
Sciaridae
The Sciaridae are a family of flies, commonly known as dark-winged fungus gnats. Commonly found in moist environments, they are known to be a pest of mushroom farms and are commonly found in household plant pots. This is one of the least studie ...
,
Stratiomyidae,
Tabanidae
Horse-flies or horseflies are true flies in the family Tabanidae in the insect order Diptera. They are often large and agile in flight, and only the female horseflies bite animals, including humans, to obtain blood. They prefer to fly in s ...
,
Tanyderidae
Tanyderidae, sometimes called primitive crane flies, are long, thin, delicate flies with spotted wings, superficially similar in appearance to some Tipulidae, Trichoceridae, and Ptychopteridae. Most species are restricted in distribution. They ...
,
Tipulidae
Crane fly is a common name referring to any member of the insect family Tipulidae. Cylindrotominae, Limoniinae, and Pediciinae have been ranked as subfamilies of Tipulidae by most authors, though occasionally elevated to family rank. In the m ...
,
Valeseguyidae
Valeseguyidae is a family of flies, belonging to Scatopsoidea. It contains only one known extant species, ''Valeseguya rieki'', known from a single male specimen found in Victoria, Australia, described in 1990. It was initially classified as a m ...
,
Xylomyidae
Xylomyidae is a family of flies known commonly as the wood soldier flies. They are xylophagous and are associated with dead or dying wood.
Description
For terms see Morphology of Diptera.
These flies are 4 to 14 millimeters long. Their colorat ...
, †
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, †
Tethepomyiidae, †
Zhangsolvidae
Zhangsolvidae is an extinct family of brachyceran flies known from the Cretaceous period. Members of the family possess a long proboscis, varying in length between 1.3 and 7 mm depending on the species, and were probably nectarivores. A spec ...
and several ''incertae sedis'' taxa.
= Coleopterans
=
![Electroxyra 1](https://upload.wikimedia.org/wikipedia/commons/a/a7/Electroxyra_1.png)
Over ninety families of
coleopterans
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 described ...
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
The Anthicidae are a family of beetles that resemble ants. They are sometimes called ant-like flower beetles or ant-like beetles. The family comprises over 3,500 species in about 100 genera.
Description
Their heads constrict just in front of ...
,
Anthribidae
Anthribidae is a family of beetles also known as fungus weevils. The antennae are not elbowed, may occasionally be longer than the body and thread-like, and can be the longest of any members of Curculionoidea. As in the Nemonychidae, the labrum ...
, †
Apotomouridae,
Belidae
Belidae is a family of weevils, called belids or primitive weevils because they have straight antennae, unlike the "true weevils" or Curculionidae which have geniculate (elbowed) antennae. They are sometimes known as "cycad weevils", but this pro ...
,
Boganiidae
Boganiidae is a family of beetles, in the superfamily Cucujoidea. Members of the family are found in southern Africa, Australia and New Caledonia. Adults and larvae are pollenivorous, feeding on the pollen of cycads and flowering plants of the f ...
,
Bostrichidae
The Bostrichidae are a family of beetles with more than 700 described species. They are commonly called auger beetles, false powderpost beetles, or horned powderpost beetles. The head of most auger beetles cannot be seen from above, as it is down ...
,
Brachypsectridae
The Brachypsectridae are a family of beetles commonly known as the Texas beetles. There are only two extant genera, '' Brachypsectra'' and '' Asiopsectra. Brachypsectra'' has a cosmopolitan distribution, mostly in arid regions, while ''Asiopsectr ...
,
Buprestidae,
Cantharidae
The soldier beetles (Cantharidae) are relatively soft-bodied, straight-sided beetles. They are cosmopolitan in distribution. One of the first described species has a color pattern reminiscent of the red coats of early British soldiers, hence the ...
,
Carabidae
Ground beetles are a large, cosmopolitan family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it is one of the 10 most species-rich animal fami ...
,
Caridae
Caridae is a small Gondwanan family of weevils. They are considered part of the primitive weevil group, because they have straight rather than geniculate (elbowed) antennae. The insertion of the antennae on the rostrum cannot be seen from above. ...
,
Cerambycidae
The longhorn beetles (Cerambycidae), also known as long-horned or longicorns, are a large family of beetles, with over 35,000 species described. Most species are characterized by extremely long antennae, which are often as long as or longer than ...
,
Cerophytidae
The Cerophytidae are a family of beetles belonging to Elateroidea. Larvae are associated with rotting wood, on which they are presumed to feed.Costa, Cleide, Vanin, Sergio A., Lawrence, John F. and Ide, Sergio. "4.4. Cerophytidae Latreille, 1834" ...
,
Cerylonidae,
Chrysomelidae,
Ciidae
The minute tree-fungus beetles, family Ciidae, are a sizeable group of beetles which inhabit Polyporales bracket fungi or coarse woody debris. Most numerous in warmer regions, they are nonetheless widespread and a considerable number of species ...
,
Clambidae
Clambidae is a family of beetles. They are known commonly as the minute beetlesMajka, C. G., & Langor, D. (2009)Clambidae (Coleoptera) of Atlantic Canada.''Journal of the Acadian Entomological Society'' 5(7), 32-40. or the fringe-winged beetles.< ...
,
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 fe ...
,
Cucujidae,
Cupedidae
The Cupedidae are a small family of beetles, notable for the square pattern of "windows" on their elytra (hard forewings), which give the family their common name of reticulated beetles.
The family consists of about 30 species in 9 genera, with ...
,
Curculionidae,
Cyclaxyridae
Cyclaxyridae are a family of beetles in the superfamily Cucujoidea. The only living genus is '' Cyclaxyra'', with two species endemic to New Zealand. Other species have been named from fossils. They are also known as sooty mould beetles due to the ...
,
Dascillidae,
Dermestidae
Dermestidae are a family of Coleoptera that are commonly referred to as skin beetles. Other common names include larder beetle, hide or leather beetles, carpet beetles, and khapra beetles. There are over 1,100 species described.
Dermestids have ...
,
Dytiscidae
The Dytiscidae – based on the Greek ''dytikos'' (δυτικός), "able to dive" – are the predaceous diving beetles, a family of water beetles. They occur in virtually any freshwater habitat around the world, but a few species live ...
,
Elateridae
Elateridae or click beetles (or "typical click beetles" to distinguish them from the related families Cerophytidae and Eucnemidae, which are also capable of clicking) are a family of beetles. Other names include elaters, snapping beetles, sp ...
,
Elmidae
Elmidae, commonly known as riffle beetles, is a family of beetles in the superfamily Byrrhoidea described by John Curtis in 1830. Both adults and larvae are usually aquatic, living under rocks in fast-flowing shallow areas of streams, such as r ...
,
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
Eucinetidae is a family of beetles, notable for their large coxal plates that cover much of the first ventrite of the abdomen, sometimes called plate-thigh beetles. The family is small for beetles, with about 50 species in 11 genera, but are ...
,
Eucnemidae
Eucnemidae, or false click beetles, are a family of elateroid beetles including about 1700 species distributed worldwide.
Description
Closely related to the family Elateridae, specimens of Eucnemidae can reach a length of . Bodies are slight ...
,
Geotrupidae
Geotrupidae (from Greek γῆ ''(gē)'', earth, and τρῡπητής ''(trȳpētēs)'', borer) is a family of beetles in the order Coleoptera. They are commonly called earth-boring dung beetles or dor beetles. Most excavate burrows in which t ...
,
Glaresidae
''Glaresis'' is a genus of beetles, sometimes called "Enigmatic scarab beetles", in its own family, the Glaresidae. It is closely related to, and was formerly included in, the family Scarabaeidae. Although its members occur in arid and sandy area ...
,
Gyrinidae
The whirligig beetles are water beetles, comprising the family Gyrinidae that usually swim on the surface of the water if undisturbed, though they swim underwater when threatened. They get their common name from their habit of swimming rapidly i ...
,
Heteroceridae
Heteroceridae, the variegated mud-loving beetles, are a widespread and relatively common family of beetles found on every continent except for Antarctica.
Around two hundred and fifty species of heterocerids are known to occur worldwide. They a ...
,
Histeridae
Histeridae is a family of beetles commonly known as clown beetles or Hister beetles. This very diverse group of beetles contains 3,900 species found worldwide. They can be easily identified by their shortened elytra that leaves two of the seven ...
,
Hybosoridae
Hybosoridae, sometimes known as the scavenger scarab beetles, is a family of scarabaeiform beetles. The 690 species in 97 genera occur widely in the tropics, but little is known of their biology.
Hybosorids are small, 5–7 mm in length a ...
,
Hydraenidae
Hydraenidae is a family of very small aquatic beetles, sometimes called "Minute moss beetles", with a worldwide distribution. They are around 0.8 to 3.3 mm in length. The adults store air on the underside of the body as well as beneath the el ...
,
Hydrophilidae
Hydrophilidae, also known colloquially as water scavenger beetles, is a family of beetles. Aquatic hydrophilids are notable for their long maxillary palps, which are longer than their antennae. Several of the former subfamilies of Hydrophilidae ...
,
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
Kateretidae also known as short-winged flower beetles are a family of beetles in the superfamily Cucujoidea. There are 10 extant and 4 extinct genera, and at least 40 described species. They are found worldwide except in New Zealand. Adults are ...
,
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
The Lampyridae are a family of elateroid beetles with more than 2,000 described species, many of which are light-emitting. They are soft-bodied beetles commonly called fireflies, lightning bugs, or glowworms for their conspicuous production ...
,
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
Stag beetles are a family of about 1,200 species of beetles in the family Lucanidae, currently classified in four subfamilies.Smith, A.B.T. (2006). A review of the family-group names for the superfamily Scarabaeoidea (Coleoptera) with corrections ...
,
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
Melandryidae is a family of beetles in the superfamily Tenebrionoidea. Members of the family are found worldwide, with around 420 species in 60 genera. Larvae and adults are generally associated with rotting wood and wood-decomposing fungi.
Ge ...
,
Meloidae
Blister beetles are beetles of the family Meloidae, so called for their defensive secretion of a blistering agent, cantharidin. About 7,500 species are known worldwide. Many are conspicuous and some are aposematically colored, announcing their ...
,
Melyridae
Melyridae (common name: soft-winged flower beetles) are a family of beetles of the superfamily Cleroidea.
Description
Most are elongate-oval, soft-bodied beetles 10 mm long or less. Many are brightly patterned in black and brown, yellow, ...
, †
Mesophyletidae
Mesophyletidae is an extinct family of weevils known from a number of genera preserved in Cretaceous amber. The family was first described as a subfamily in the extant family Caridae, and subsequently raised to family status in 2018.
Distribut ...
,
Micromalthidae,
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,
Nemonychidae
Nemonychidae is a small family of weevils, placed within the ''primitive weevil'' group because they have straight rather than geniculate (elbowed) antennae. They are often called pine flower weevils. As in the Anthribidae, the labrum appears ...
,
Nitidulidae
The sap beetles, also known as Nitidulidae, are a family (biology), family of beetles.
They are small (2–6 mm) ovoid, usually dull-coloured beetles, with knobbed antenna (biology), antennae. Some have red or yellow spots or bands. They fe ...
,
Oedemeridae
The family Oedemeridae is a cosmopolitan group of beetles commonly known as false blister beetles, though some recent authors have coined the name pollen-feeding beetles. There are some 100 genera and 1,500 species in the family, mostly associate ...
,
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
Passalidae is a family of beetles known variously as "bessbugs", "bess beetles", "betsy beetles" or "horned passalus beetles". Nearly all of the 500-odd species are tropical; species found in North America are notable for their size, ranging fro ...
, †
Parandrexidae, †
Passalopalpidae,
Passandridae,
Phloeostichidae,
Prostomidae
Prostomidae is a family of beetles with no vernacular common name, though recent authors have coined the name jugular-horned beetles. They are often found in dead wood. The family consist of two extant genera with about 20 species. '' Prostomis a ...
,
Psephenidae
Water-penny beetles are a family (the Psephenidae) of 272 species (in 35 genera) of aquatic beetles found on all continents except Antarctica, in both tropical and temperate areas. The young, which live in water, resemble tiny pennies. The larv ...
,
Ptiliidae
Ptiliidae is a family of very tiny beetles with a cosmopolitan distribution. This family contains the smallest of all beetles, with a length when fully grown of . The weight is approximately 0.4 milligrams. They are colloquially called feat ...
,
Ptinidae
Ptinidae is a family of beetles in the superfamily Bostrichoidea. There are at least 220 genera and 2,200 described species in Ptinidae worldwide. The family includes spider beetles and deathwatch beetles.
The Ptinidae family species are hard ...
,
Ptilodactylidae
Ptilodactylidae is a family of beetles belonging to the Elateriformia. There around 500 extant species in 35 genera. They are generally associated with riparian and aquatic habitats. The larvae generally live associated with rotting wood or veg ...
,
Ripiphoridae
Ripiphoridae (formerly spelled Rhipiphoridae) is a cosmopolitan family of some 450 described species of beetles sometimes called "wedge-shaped beetles". Ripiphoridae are unusual among beetle families in that many species are hypermetamorphic pa ...
,
Salpingidae
Salpingidae or narrow-waisted bark beetles is a family of beetles in the superfamily Tenebrionoidea. The species are small, about 1.5 – 7 mm in length. The family is globally distributed and consists of about 45 genera and 300 species, w ...
,
Scarabaeidae
The family Scarabaeidae, as currently defined, consists of over 30,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change in recent years. Several sub ...
,
Scirtidae
Scirtidae is a Taxonomic rank, family of beetles (Coleoptera). These beetles are commonly referred to as marsh beetles, as the larvae are typically associated with stagnant water, but can be found in flowing water. Adults prefer decomposing plan ...
,
Scraptiidae,
Silphidae
Silphidae is a family of beetles that are known commonly as large carrion beetles, carrion beetles or burying beetles. There are two subfamilies: Silphinae and Nicrophorinae. Nicrophorines are sometimes known as sexton beetles. The number of ...
,
Silvanidae
Silvanidae, "silvan flat bark beetles", is a family of beetles in the superfamily Cucujoidea,Thomas, M. C., and R.A. B. Leschen. 2010. Silvanidae Kirby, 1837. p. 346-350. In: Leschen, R.A.B., R.G. Beutel, and J.F. Lawrence. Coleoptera, Beetles. Vo ...
,
Smicripidae,
Sphaeriusidae
''Sphaerius'' is a genus of beetles, comprising 23 species, which are the only living members of the family Sphaeriusidae. They are typically found along the edges of streams and rivers, where they feed on algae; they occur on all continents ex ...
,
Staphylinidae,
Tenebrionidae
Darkling beetle is the common name for members of the beetle family Tenebrionidae. The number of species in the Tenebrionidae is estimated at more than 20,000 and the family is cosmopolitan in distribution.
Taxonomy
''Tenebrio'' is the Latin g ...
,
Tetratomidae
Tetratomidae is a small family of beetles sometimes called polypore fungus beetles. The family consists of several genera, most of which used to be in the family Melandryidae. Tetratomidae can be found worldwide.
Their food consists of fruiti ...
,
Thanerocleridae
Thanerocleridae is a family of beetles belonging to the superfamily Cleroidea. It was formerly considered a subfamily of Cleridae, but was recently elevated to the rank of family. The family has 36 living species in 10 genera, which are found glo ...
,
Throscidae
Throscidae is a family of elateroid beetles found worldwide (except New Zealand) with around 150 species in 5 extant genera. The larvae are soil-dwelling, siphoning fluid from mycorrhizae attached to trees. The adults are short-lived, with the ...
,
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, Rentoni ...
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
neuropterans 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
The Berothidae are a family of winged insects of the order Neuroptera. They are known commonly as the beaded lacewings. The family was first named by Anton Handlirsch in 1906. The family consists of 24 genera and 110 living species distributed ...
,
Chrysopidae
Green lacewings are insects in the large family Chrysopidae of the order Neuroptera. There are about 85 genera and (differing between sources) 1,300–2,000 species in this widespread group. Members of the genera ''Chrysopa'' and ''Chrysoper ...
,
Coniopterygidae, †
Corydasialidae,
Dilaridae
Dilaridae is a family of Euneuropteran insects in the order Neuroptera, known as "pleasing lacewings". They were formerly placed in the paraphyletic superfamily Hemerobioidea, though the group is currently placed in the monophyletic superfamily ...
, †
Dipteromantispidae,
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
Ithonidae, commonly called moth lacewings and giant lacewings, is a small family of winged insects of the insect order Neuroptera. The family contains a total of ten living genera, and over a dozen extinct genera described from fossils. The mo ...
,
Kalligrammatidae
Kalligrammatidae, sometimes known as kalligrammatids or kalligrammatid lacewings, is a family of extinct insects in the order Neuroptera (lacewings) that contains twenty genera and a number of species. The family lived from the Middle Jurassic t ...
,
Mantispidae
Mantispidae, known commonly as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a family of small to moderate-sized insects in the order Neuroptera. There are many genera with around 400 species worldwide, especially in ...
, †
Mesochrysopidae,
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 larvae ...
,
Nemopteridae
Nemopteridae, the spoonwings, are a family of neuropteran insects. They are also called thread-winged antlions. They are found in the Ethiopian, Palearctic, Australasian and Neotropical realms but absent in North America (though a fossil has bee ...
,
Nevrorthidae
The Nevrorthidae, (often incorrectly spelled "Neurorthidae"), are a small family of winged insects of the order Neuroptera. Extant species may be described as living fossils. There are 19 extant species in four genera, with a geographically di ...
,
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
Osmylidae are a small family of winged insects of the net-winged insect order Neuroptera. The osmylids, also called lance lacewings, stream lacewings or giant lacewings, are found all over the world. There are around 225 extant species.
Descr ...
,
Psychopsidae
Psychopsidae is a family (biology), family of Pterygota, winged insects of the order (biology), order Neuroptera. They are commonly called silky lacewings.
The silky lacewings are distinguishable in their adult stage by their spectacularly patte ...
,
Rachiberothidae,
Sisyridae
Sisyridae, commonly known as spongeflies or spongillaflies, are a family of winged insects in the order Neuroptera. There are approximately 60 living species described, and several extinct species identified from the fossil record.
Description
...
and several ''incertae sedis'' taxa.
= Hemiptera
=
![Mimaplax ekrypsan Fig1 A](https://upload.wikimedia.org/wikipedia/commons/b/b4/Mimaplax_ekrypsan_Fig1_A.jpg)
Over sixty families of
hemipterans are known from the Burmese amber, including:
Achilidae
Achilidae is a family of achilid planthoppers in the order Hemiptera
Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, ass ...
, †
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
The Aphrophoridae or spittlebugs are a family of insects belonging to the order Hemiptera. There are at least 160 genera and 990 described species in Aphrophoridae.
European genera
* ''Aphrophora'' Germar 1821
* '' Lepyronia'' Amyot & Servill ...
,
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, †
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, Cercopidae, Cicadellidae, Cicadidae, Cimicidae, Cixiidae, Coccidae, Coreidae, Cydnidae, Dictyopharidae, †Dinglidae Dipsocoridae, †Dorytocidae, Enicocephalidae, Fulgoridae, Gelastocoridae, Gerridae, †Hodgsonicoccidae, Hydrometridae, Issidae, †Jubisentidae, †Juraphididae, †Katlasidae, Kinnaridae, †Kozariidae, †Lalacidae, Leptopodidae, †Liadopsyllidae, †Macrodrilidae, Margarodidae, Matsucoccidae, †Mimarachnidae, †Minlagerrontidae, Miridae, Monophlebidae, Naucoridae, Nabidae, †Neazoniidae, Ochteridae, Ortheziidae,†Palaeoleptidae, †Parvaverrucosidae, †Perforissidae, †Protopsyllidiidae, †Procercopidae, Pseudococcidae, Reduvidae, Schizopteridae, †Sinoalidae, †Tajmyraphididae, Tettigarctidae, Tingidae, Tropiduchidae, Velocipedidae, Veliidae, †Weitschatidae, Xylococcidae, †Yetkhatidae, †Yuripopovinidae and several ''incertae sedis'' taxa such as ''Mesophthirus'', formerly believed to be an ectoparasite but since determined to be a basal scale insect.
= Dictyoptera
=
Twenty one families of dictyopterans are known from the Burmese amber, including: Blaberidae, †Blattulidae, Blattidae, †Caloblattinidae, Corydiidae, Ectobiidae, †Olidae, †Liberiblattinidae, †Alienopteridae, †Manipulatoridae †Umenocoleidae , Nocticolidae, †Pabuonqedidae Termites (†Archeorhinotermitidae, Hodotermitidae, Stolotermitidae, Termitidae and Mastotermitidae) 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, Compsocidae, †Cormopsocidae †Empheriidae, Liposcelididae, Manicapsocidae, Pachytroctidae, Prionoglarididae, Psyllipsocidae, Sphaeropsocidae and Trogiidae.
= Orthoptera
=
Seven families of orthopteran are known, including: †Elcanidae, Gryllidae, Mogoplistidae, Ripipterygidae, Tetrigidae, Tettigoniidae and Tridactylidae
= Trichoptera
=
Eight families of Caddisfly, trichopteran are known, including: Calamoceratidae, †Dysoneuridae, Helicopsychidae, Hydroptilidae, Odontoceridae, Philopotamidae, Polycentropodidae and Psychomyiidae.
= Dermaptera
=
![Astreptolabis_laevis_fig1_A](https://upload.wikimedia.org/wikipedia/commons/6/6f/Astreptolabis_laevis_fig1_A.jpg)
Five families of Earwig, dermapteran are known, including: Anisolabididae, Diplatyidae, Labiduridae and Pygidicranidae.
= Embioptera
=
Four families of embiopteran are known, including: Clothodidae, Oligotomidae, Notoligotomidae and †Sorellembiidae.
= Notoptera
=
One species of notopteran is known, a nymph ice crawler (Grylloblattidae) ''Sylvalitoralis, Sylvalitoralis cheni.''
= Strepsiptera
=
Two families of strepsipteran are known, †Cretostylopidae and †Phthanoxenidae
= Lepidoptera
=
Six families of lepidopteran are known, including: Agathiphagidae, Douglasiidae, Gelechiidae, Gracillariidae, Lophocoronidae and Micropterigidae.
= Megaloptera
=
One species of megalopteran is known, ''Haplosialodes liui'' of the family Sialidae.
= Phasmatodea
=
Four families of phasmatodean are known: †Archipseudophasmatidae, Phasmatidae. †Pterophasmatidae and Timematidae
= Thysanoptera
=
Five families of thrips are known, including: Aeolothripidae, Melanthripidae, †Rohrthripidae, Thripidae and Stenurothripidae.
= Plecoptera
=
Two families of stoneflies are known, Perlidae and †Petroperlidae.
= Raphidioptera
=
![Nanoraphidia electroburmica EMTG BA- 001202 Fig1 A](https://upload.wikimedia.org/wikipedia/commons/8/8a/Nanoraphidia_electroburmica_EMTG_BA-_001202_Fig1_A.png)
One family of Raphidiopteran is known, †Mesoraphidiidae.
= †Chresmodidae
=
A species of the enigmatic long legged Chresmodidae, chresmodid insect ''Chresmoda'' is known.
= †Tarachoptera
=
One family of Tarachopteran is known: †Tarachocelidae
= †Permopsocida
=
One family of Permopsocidan is known: †Archipsyllidae
= Zoraptera
=
Multiple species of ''Zorotypus'' and the monotypic genus ''Xenozorotypus'' are known.
Nematoda
Five families of nematodes are known, including: Cosmocercidae, Heterorhabditidae, Mermithidae, Oxyurida, Thelastomatidae, Aphelenchoididae
Nematomorpha
One genus of Nematomorpha, nematomorph is known: ''Cretachordodes'' (Chordodidae, Gordioidea)
Mollusca
![Archaeocyclotus plicatula Fig1 B](https://upload.wikimedia.org/wikipedia/commons/e/e9/Archaeocyclotus_plicatula_Fig1_B.jpg)
Aside from the previously mentioned ammonites and marine gastropod shells, Seven families of terrestrial gastropod are known: Diplommatinidae, Pupinidae, Achatinidae, Punctidae, Valloniidae, Assimineidae 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 Enantiornithes, enantiornithine birds including juveniles and partial wings and preserved feet, including a diagnostic taxon, ''Elektorornis''. A complete skull of the lizard ''Oculudentavis'' is known. ''Electrorana'' 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 with preserved gecko feet, toe pads (''Cretaceogekko''). and a miniaturised (~2 cm) long possible stem-Anguimorpha, anguimorph (''Barlochersaurus'')
One of the "lizard" specimens was initially described to be a chamelonid, actually turned out to be an Albanerpetontidae, albanerpetontid amphibian. This was described in 2020 as the new genus and species ''Yaksha perettii.''
Flora
Angiosperms
Eleven species of Angiosperm are known in nine genera, including members of Cornaceae, Cunoniaceae, Lauraceae, ?Monimiaceae and Laurales ''incertae sedis''. Poales ''incertae sedis'' and Angiosperm incertae sedis.
Bryopsida
Two genera of Bryopsida in the separate orders Dicranales and Hypnodendrales
Jungermanniopsida
Three families of Jungermanniopsida are known, Frullaniaceae, Lepidolaeanaceae, Radulaceae.
Pinophyta
Two families of Pinophyta, Pinopsida are known: Araucariaceae and Cupressaceae including ''Metasequoia''.
Pteridopsida
Five families of Leptosporangiate fern, Pteridopsidan are known: Cystodiaceae, Dennstaedtiaceae, Lindsaeaceae, Pteridaceae, Thyrsopteridaceae, and several genera of Polypodiales ''incertae sedis.''
Amoebozoa
Myxogastria
Sporocarp (fungi), Sporocarps of extant myxogastrid slime mould genus ''Stemonitis'' 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, Yunnan Province as early as the first century AD according to the ''Book of the Later Han'' 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, Han Dynasty (25 - 220 Common Era, CE). It was first mentioned in European sources by the Society of Jesus, Jesuit 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 among other goods. mixed and lower quality amber was sold from around Thai baht, 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
![Noje Bum amber](https://upload.wikimedia.org/wikipedia/commons/7/76/Noje_Bum_amber.jpg)
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,'' 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
The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
,
Diapriidae
The Diapriidae are a family of parasitoid wasps. These tiny insects have an average length of 2–4 mm and never exceed 8 mm. They typically attack larvae and pupae of a wide range of insects, especially flies. The about 2,300 described ...
, Scelioninae, Scelionidae) Diptera (
Ceratopogonidae
Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally in length. The family includes more than 5,000 species, distributed worldwide, apart from the Antarctic and the Arctic.
Ceratopogonidae are holomet ...
,
Chironomidae
The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many specie ...
) Dictyoptera (Cockroach, Blattaria, Mantodea) planthoppers,
Berothidae
The Berothidae are a family of winged insects of the order Neuroptera. They are known commonly as the beaded lacewings. The family was first named by Anton Handlirsch in 1906. The family consists of 24 genera and 110 living species distributed ...
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