The species figured and/or listed on this page have been officially described in the scientific literature.
They include all of the species named from the Kishenehn Formation. Most have been described by me and/or my collaborators, but other laboratories have also named new species of Kishenehn Formation insects.
Occasionally, specimens from other fossil sites around the world, that are related to Kishenehn Formation fossils, have been named and described by me and/or my collaborators. These are also included here.
They include all of the species named from the Kishenehn Formation. Most have been described by me and/or my collaborators, but other laboratories have also named new species of Kishenehn Formation insects.
Occasionally, specimens from other fossil sites around the world, that are related to Kishenehn Formation fossils, have been named and described by me and/or my collaborators. These are also included here.
Diptera (Flies)
The fossil fly Bolitohilia warreni
Bolitophila warreni (Diptera: Bolitophilidae)
Named after Jill Warren, a Corvallis, Montana middle school science teacher who participated in Kishenehn Formation fieldwork for many years.
Named after Jill Warren, a Corvallis, Montana middle school science teacher who participated in Kishenehn Formation fieldwork for many years.
A fossil fly of the genus Bolitophila
Bolitophila rohdendorfi (Diptera: Bolitophilidae)
Described from Baltic amber by Vladimir Blagoderov, Principal Curator of Invertebrates at the National Museum Scotland in Edinburgh.
Described from Baltic amber by Vladimir Blagoderov, Principal Curator of Invertebrates at the National Museum Scotland in Edinburgh.
The genitalia of two similar fossil species, one in shale and one in amber.
A comparison of the genitalia of the two related fossil species, Bolitophila warreni and Bolitophila rohdendorfi,
one in amber and the other in shale rock, documents the beautiful preservation of the Kishenehn Formation insects.
one in amber and the other in shale rock, documents the beautiful preservation of the Kishenehn Formation insects.
A fossil mosquito with beautifully preserved wings.
Culiseta lemniscata (Diptera: Culicidae). Common name: Mosquitos
This specimen was identified to the genus Culiseta based on the presence of a small group of scales at the base of the wings.
This specimen was identified to the genus Culiseta based on the presence of a small group of scales at the base of the wings.
Note the very long proboscis of this fossil mosquito.
Culiseta kishenehn (Diptera: Culicidae). Common name: Mosquitos
Four species of fossil flies in the family Dixidae, a family closely related to mosquitos (Culicidae) have been described from the Kishenehn Formation. They are Dixella intacta, Dixella eomarginata, Dixella spinilobata and Dixella curvistyla, from left to right in the slideshow below.
The study of these four specimens, and a comparative study of other dixids in the genera Dixella and Dixa, demonstrated that there is no single wing venation character or combination of characters that is unique to either genus and that the current taxonomic definition of these two genera is inaccurate.
The study of these four specimens, and a comparative study of other dixids in the genera Dixella and Dixa, demonstrated that there is no single wing venation character or combination of characters that is unique to either genus and that the current taxonomic definition of these two genera is inaccurate.
Fossil insect wings contain important taxonomic information.
Synneuron eomontana (Diptera: Canthyloscelidae) 1 & 2
Synneuron jelli (Diptera: Canthyloscelidae) 3 & 4
Both of these rare specimens, the only fossils of the genus known to exist, are single wings. S. jelli was collected in Lower Cretaceous Koonwarra Fossil Bed near Melbourne, Australia.
Only two species of the genus Synneuron live today. The geographic distribution of these two fossil specimens suggest that the genus was much more diverse in deep time.
Synneuron jelli (Diptera: Canthyloscelidae) 3 & 4
Both of these rare specimens, the only fossils of the genus known to exist, are single wings. S. jelli was collected in Lower Cretaceous Koonwarra Fossil Bed near Melbourne, Australia.
Only two species of the genus Synneuron live today. The geographic distribution of these two fossil specimens suggest that the genus was much more diverse in deep time.
An anamalous fossil fly from the Eocene.
Eoanomala melas (Diptera: Bombyliidae). Common name: Bee fly
Given its unusual morphological characteristics, this specimen was designated as belonging to a new genus.
Given its unusual morphological characteristics, this specimen was designated as belonging to a new genus.
A fossil bee fly, beautifully preserved in amber.
Anthrax succini (Diptera: Bombyliidae). Common name: Bee fly
This specimen was found in the NMNH’s marvelous collection of amber from the Dominican Republic.
This specimen was found in the NMNH’s marvelous collection of amber from the Dominican Republic.
Helius constenius (Diptera: Limoniidae). Common name: crane fly
Described by Wiesław Krzemiński in 1991, this was the first fossil insect from the Kishenehn Formation to be identified to genus and given a new species name.
Described by Wiesław Krzemiński in 1991, this was the first fossil insect from the Kishenehn Formation to be identified to genus and given a new species name.
A fossil male crane fly with an intact phallus.
Tipula fji (Tipulidae). Common name: Crane flies
There is some disagreement as to whether or not Tipulidae, Limoniidae and Cylindrotomidae (see specimens below) are families or the subfamilies Tipulinae, Limoniinae and Cylindrotominae all within the family Tipulidae.
This particular specimen is unusual in that it has preserved the long thin aedeagus, the male’s phallus, within the insect’s abdomen. Note the dark heavily sclerotized spines at the terminus of the genitalia.
There is some disagreement as to whether or not Tipulidae, Limoniidae and Cylindrotomidae (see specimens below) are families or the subfamilies Tipulinae, Limoniinae and Cylindrotominae all within the family Tipulidae.
This particular specimen is unusual in that it has preserved the long thin aedeagus, the male’s phallus, within the insect’s abdomen. Note the dark heavily sclerotized spines at the terminus of the genitalia.
A fossil crane fly in the family Limoniidae.
Ellipteroides kishenehn (Limoniidae). Common name: Crane flies
Ellipteroides kishenehn is the first known fossil in genus
Ellipteroides kishenehn is the first known fossil in genus
This fossil cranefly is in the family Cylindrotomidae.
Cyttaromyia lynnae (Cylindrotomidae). Common name: Crane flies
This fossil fly has the setae (hairs) on its wing veins preserved.
Sylvicola silibrarius (Anisopodidae). Common name: Wood gnats
The species name is in honor of the employees of the library at the National Museum of Natural History.
The species name is in honor of the employees of the library at the National Museum of Natural History.
The wing venation and antennal morphology of this fossil fly helped to identify it to the genus Efcookella.
Efcookella nigra (Scatopsidae). Common name: Dung midges
Efcookella nigra is the first known fossil in genus. Of the 17 fossils in the family, this is the only one that is not an amber inclusion.
Efcookella nigra is the first known fossil in genus. Of the 17 fossils in the family, this is the only one that is not an amber inclusion.
The wing venation of this fossil fly is unusual relative to other genera in the family Bibionidae.
Bibiodes kishenehnensis (Bibionidae). Common name: March flies; Lovebugs
The mouthparts of this fossil fly are beautifully preserved.
Eosciarites hermes (Sciaridae). Common name: Dark-winged fungus gnats
Eosciarites is a new genus. The suffix “-ites” indicates a degree of uncertainty relative to the accuracy of this placement.
Eosciarites is a new genus. The suffix “-ites” indicates a degree of uncertainty relative to the accuracy of this placement.
Count them. All six legs are preserved in this fossil gnat.
Rymosia hypnolithica (Mycetophilidae). Common name: Fungus gnats
This specimen is a male; its genitalia, which are exceedingly important in the taxonomy of flies, is beautifully preserved.
This specimen is a male; its genitalia, which are exceedingly important in the taxonomy of flies, is beautifully preserved.
The genitalia of this fossil fly, important in its classification, are beautifully preserved.
Litoleptis araeostylus (Rhagionidae). Common name: Snipe flies
A photograph of this fossil robber fly hangs in the museum's library.
Kishenehnoasilus bhl (Asilidae). Common name: Robber flies
Dr. Torsten Dikow named this species bhl, an acronym for the Biodiversity Heritage Library. This online library of 16th – 19th century scientific publications is an invaluable resource to scientists around the world.
Dr. Torsten Dikow named this species bhl, an acronym for the Biodiversity Heritage Library. This online library of 16th – 19th century scientific publications is an invaluable resource to scientists around the world.
In several families of flies, including the Hybotidae, the male genitalia has rotated to one side.
Drapetis adelomedos (Hybotidae). Common name: Dance flies
All other fossils of this genus are amber inclusions.
All other fossils of this genus are amber inclusions.
The legs of this fossil fly are preserved despite their lack of sclerotization.
Salishomyia eocenica (Dolichopodidae). Common name: Long-legged flies
Salishomyia is a new genus
Salishomyia is a new genus
The mouthparts of this fossil fly have been preserved.
Agathomyia eocenica sp. nov. (Platypezidae). Common name: Flat-footed flies
Agathomyia eocenica is the first known fossil in genus. The family name is an apt one as the legs are dorsolaterally flattened (wide and thin). The Greek words “platy” and “peza” mean flat and foot, respectively.
Agathomyia eocenica is the first known fossil in genus. The family name is an apt one as the legs are dorsolaterally flattened (wide and thin). The Greek words “platy” and “peza” mean flat and foot, respectively.
Despite the loss of a large portion of this fossil fly's thorax, it was easily identified to its genus.
Lonchoptera eocenica (Lonchopteridae). Common name: Spear-winged flies
The family and generic names are based on the shape of the fly’s wing, the apex of which is distinctly pointed. The Greek words “lonch” and “pteron” mean spearhead and wing, respectively.
Lonchoptera eocenica is the only known fossil in genus and, given that the entire family consists of only this one exant genus, the only fossil in the entire family. Morphological analysis indicates that it is sister to all extant species in the family; that is, it is a stem Lonchoptera. It differs from living species by having wings that are not as elongated and are less pointed at the apex.
The family and generic names are based on the shape of the fly’s wing, the apex of which is distinctly pointed. The Greek words “lonch” and “pteron” mean spearhead and wing, respectively.
Lonchoptera eocenica is the only known fossil in genus and, given that the entire family consists of only this one exant genus, the only fossil in the entire family. Morphological analysis indicates that it is sister to all extant species in the family; that is, it is a stem Lonchoptera. It differs from living species by having wings that are not as elongated and are less pointed at the apex.
Genitalic morphology is exceedingly important in the study of fossil insects.
Aenigmatias kishenehnensis (Phoridae). Common name: Scuttle flies
Extant females of the 14 species of modern Aenigmatias lay their eggs in ant larvae. They have evolved to spend their lives in ant colonies and have, over time, lost their wings. Did Aenigmatias kishenehnensis parasitize ants? As we have seen there were plenty of ants around the shores of Lake Kishenehn. But Aenigmatias kishenehnensis still has wings.
Aenigmatias kishenehnensis does have female genitalia (2), with peglike setae, that are characteristic of modern species (3) and diagnostic of the genus.
Extant females of the 14 species of modern Aenigmatias lay their eggs in ant larvae. They have evolved to spend their lives in ant colonies and have, over time, lost their wings. Did Aenigmatias kishenehnensis parasitize ants? As we have seen there were plenty of ants around the shores of Lake Kishenehn. But Aenigmatias kishenehnensis still has wings.
Aenigmatias kishenehnensis does have female genitalia (2), with peglike setae, that are characteristic of modern species (3) and diagnostic of the genus.
The head of this fossil fly is as wide as its body.
Pipunculinae (Pipunculidae). Common name: Big-headed flies
Given that only the ventral aspect of his specimen is visible, and the terminus of its abdomen is missing, it could only be identified to subfamily.
The antennae are nicely preserved with long filamentous flagella, bulbous basal segments and their setae, both large and small.
Coleoptera (Beetles)
This fossil beetle is a member of the family Ptiliidae, the smallest species of which are less than a third of a millimeter in length.
Ptenidium Kishenehnsis (Coleoptera: Ptiliidae). Common name: featherwing beetles.
The scale bar is a tenth of a millimeter long – this tiny fossil beetle is only 0.65 mm in length!! Species in this family are so small that the female has only a single ovary. The preservation is amazing – the feather-like inner wings are readily visible.
The scale bar is a tenth of a millimeter long – this tiny fossil beetle is only 0.65 mm in length!! Species in this family are so small that the female has only a single ovary. The preservation is amazing – the feather-like inner wings are readily visible.
This fossil insect is a rove beetle, a group characterized by very short outer wings.
Tympanophorus greenwalti (Coleoptera: Staphylinidae). Common name: Rove beetle
At first glance, this fossil appears to be poorly preserved; its abdomen is compressed and distorted and a large portion of it is missing. But look closely – a great deal of important morphological detail is present.
This specimen is the oldest definitive representative of the Staphylinini Propria group of rove beetles.
At first glance, this fossil appears to be poorly preserved; its abdomen is compressed and distorted and a large portion of it is missing. But look closely – a great deal of important morphological detail is present.
This specimen is the oldest definitive representative of the Staphylinini Propria group of rove beetles.
Hymenoptera (Wasps and Bees)
The fossil insects of the Kishenehn Formation are usually very small. This wasp is less than a millimeter in length.
Eoeustochus kishenehn (Hymenoptera: Mymaridae). Common name: Fairy wasps.
These exceedingly tiny parasitic wasps, many less than a millimeter in length, are characterized by wings with fringes of long hairs.
An additional five species of Mymaridae, Eoeustochus borchersi, Eoanaphes stethynioides, Gonatocerus greenwalti, Gonatocerus kootenai and Gonatocerus rasnitsyni, are figured, left to right, in the slideshow below.
These exceedingly tiny parasitic wasps, many less than a millimeter in length, are characterized by wings with fringes of long hairs.
An additional five species of Mymaridae, Eoeustochus borchersi, Eoanaphes stethynioides, Gonatocerus greenwalti, Gonatocerus kootenai and Gonatocerus rasnitsyni, are figured, left to right, in the slideshow below.
This fossil wasp is one of the most important specimens in the Kishenehn Formation collection.
Phasmatopelecinus leonae (Hymenoptera: Pelecinidae)
The long slender abdomen of this specimen is characteristic of the females of this family which lay their eggs in the larvae of June beetles that live buried in soil. But this specimen is much shorter than the few species that live today, which are all about 3 inches long.
The pelecinid wasps were much more diverse a hundred million years ago. Phasmatopelecinus leonae, named after Leona Constenius who collected the specimen, is a very rare and important specimen relative to our understanding of the evolution of this group of wasps.
The long slender abdomen of this specimen is characteristic of the females of this family which lay their eggs in the larvae of June beetles that live buried in soil. But this specimen is much shorter than the few species that live today, which are all about 3 inches long.
The pelecinid wasps were much more diverse a hundred million years ago. Phasmatopelecinus leonae, named after Leona Constenius who collected the specimen, is a very rare and important specimen relative to our understanding of the evolution of this group of wasps.
This fossil wasp has a very characteristic ovipositor.
Scelioninae (Hymenoptera: Platygastridae)
According to Dr. Elijah Talamas, curator of Hymenoptera at the Florida State Collection of Arthropods, this specimen is the oldest record of the Scelio-type ovipositor system. He states that “near the tip of the ovipositor is a dark area that corresponds to the typical location of sclerotized tergite 7+8 when it is extruded with the ovipositor.” and makes the observation that important information can be obtained from specimens that are not designated to a genus and species.
According to Dr. Elijah Talamas, curator of Hymenoptera at the Florida State Collection of Arthropods, this specimen is the oldest record of the Scelio-type ovipositor system. He states that “near the tip of the ovipositor is a dark area that corresponds to the typical location of sclerotized tergite 7+8 when it is extruded with the ovipositor.” and makes the observation that important information can be obtained from specimens that are not designated to a genus and species.
This fossil wasp was identified to a genus but not given a species name.
Fidiobia sp. (Hymenoptera: Platygastridae)
This fossil ant is a reproductive - only mating-capable males and females have wings.
Ktunaxia jucunda (Formicidae). Common name: Ants
Ktunaxia jucunda belongs to a new genus. It is morphologically distinct from other ants due to the unusually short and stocky terminal half of its antennae.
John LaPolla of Towson University has described 11 additional species of ants, in 10 different genera, from the Kishenehn Formation. They are Crematogaster aurora, Pseudomyrmex saxulum, Dolichoderus dlusskyi, Proiridomyrmex rotundatus, Protazteca eocenica, Formica annosa, Lasius glom, Solenopsites abdita, Ponerites kishenehne, Eoformica brevipetiola and Eoformica latimedia, from left to right, in the slideshow below.
Two of the new species from the Kshenehn Formation, Crematogaster aurora and Pseudomyrmex saxulum, are the oldest known species in their respective genus.
Almost all of the fóssil ants of the Kishenehn Formation have wings; very few are worker ants. Most are alates (male and female reproductives) which were seeking mates. Flying over algae-covered water in lake Kishenehn, they fell and became trapped on the sticky mat surface – the first step in the fossilization process.
Dr. LaPolla states that “The Eocene is of particular interest for understanding ant evolution because it is during this period that many present-day speciose and ecologically dominant clades of ants apparently emerged.”
Ktunaxia jucunda belongs to a new genus. It is morphologically distinct from other ants due to the unusually short and stocky terminal half of its antennae.
John LaPolla of Towson University has described 11 additional species of ants, in 10 different genera, from the Kishenehn Formation. They are Crematogaster aurora, Pseudomyrmex saxulum, Dolichoderus dlusskyi, Proiridomyrmex rotundatus, Protazteca eocenica, Formica annosa, Lasius glom, Solenopsites abdita, Ponerites kishenehne, Eoformica brevipetiola and Eoformica latimedia, from left to right, in the slideshow below.
Two of the new species from the Kshenehn Formation, Crematogaster aurora and Pseudomyrmex saxulum, are the oldest known species in their respective genus.
Almost all of the fóssil ants of the Kishenehn Formation have wings; very few are worker ants. Most are alates (male and female reproductives) which were seeking mates. Flying over algae-covered water in lake Kishenehn, they fell and became trapped on the sticky mat surface – the first step in the fossilization process.
Dr. LaPolla states that “The Eocene is of particular interest for understanding ant evolution because it is during this period that many present-day speciose and ecologically dominant clades of ants apparently emerged.”
Orthoptera (Grasshoppers and Katydids)
This is the largest fossil insect in the Kishenehn Formation collection.
Pseudotettigonia leona (Orthoptera: Tettigoniidae). Common name: katydid.
Katydids are known for their mating calls. This Katydid wing has preserved its sound-making apparatus – a rare event in the fossil record. Occupying the bottom left corner of the wing, the “stridulatory field” contains a long narrow row of teeth called the “file” (bottom right), over which the sharp edge of a “scraper” on another wing is rubbed - as if playing a violin.
This is the largest insect ever found in the Kishenehn Formation; the wing is over 5 cm in length.
Katydids are known for their mating calls. This Katydid wing has preserved its sound-making apparatus – a rare event in the fossil record. Occupying the bottom left corner of the wing, the “stridulatory field” contains a long narrow row of teeth called the “file” (bottom right), over which the sharp edge of a “scraper” on another wing is rubbed - as if playing a violin.
This is the largest insect ever found in the Kishenehn Formation; the wing is over 5 cm in length.
This fossil katydid was collected in Europe.
Tettigonia bricei (Orthoptera: Tettigoniidae).
This katydid specimen is from the late Miocene site of la Montagne d’Andance (the Mountain of Andance) in Saint-Bauzile, France. It is named for the scientist who described but did not name the specimen.
This katydid specimen is from the late Miocene site of la Montagne d’Andance (the Mountain of Andance) in Saint-Bauzile, France. It is named for the scientist who described but did not name the specimen.
Odonata (Damselflies and Dragonflies)
Fossil damselflies exist in the Kishenehn Formation only as isolated wings.
Eolestes ramosus (Odonata: Eolestidae)
The description of this species of damselfly led to the establishment of the new family Eolestidae.
The description of this species of damselfly led to the establishment of the new family Eolestidae.
The shiny objects that surround this specimen are fossil ostracods.
Lutetialestes uniformis (Odonata: family unknown)
Blattaria (Roaches)
The wing venation pattern of this fossil cockroach was all that was required for its identification.
Latiblattella avita (Blattaria: Ectobiidae). Common name: Cockroach
A wing and a leg but enough for its identification. L. avita is the first fossil in its genus.
A wing and a leg but enough for its identification. L. avita is the first fossil in its genus.
Raphidioptera (Snakeflies)
P. borealis consisted of a front wing; this specimen is a hind wing - might the two be from the same species?
Paraksenocellia sp. cf. P. borealis (Raphidioptera: Inocelliidae). Common name: snakefly; relictual, predatory, related to Neuroptera (e.g. lacewings)
Sometimes, even if just a single wing is preserved, there is still enough information to identify the fossil to a genus.
Sometimes, even if just a single wing is preserved, there is still enough information to identify the fossil to a genus.
Hemiptera (True Bugs)
The wing venation patterns of fossil insects may be the paleoentomologist's most important tool.
Eogyropsylla paveloctogenarius (Hemiptera: Aphalaridae). Common name: Jumping plant lice
This specimen is the oldest known representative of the superfamily Psylloidea sensu stricto.
This specimen is the oldest known representative of the superfamily Psylloidea sensu stricto.