Coxoplectoptera or "chimera wings" is an extinct

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

stem-group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

mayflies 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 order ...

containing one family, Mickoleitiidae. Together with mayflies ( Ephemeroptera), Coxoplectoptera are assigned to the clade Heptabranchia. Two adult and more than 20 nymphal fossils of ''Mickoleitia'' have been scientifically described from Mesozoic outcrops, mainly from the

Lower Cretaceous Lower may refer to: * ''Lower'' (album), 2025 album by Benjamin Booker * Lower (surname) * Lower Township, New Jersey *Lower Receiver (firearms) * Lower Wick Gloucestershire, England See also * Nizhny {{Disambiguation ...

Crato Formation The Crato Formation is a geologic formation (stratigraphy), formation of Early Cretaceous (Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätten, Lagerstätte (undisturbed fossil accumulation) for palaeontology, pa ...

Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...

(in total, around 40 fossil nymphs have been found). Both the winged adults and the aquatic nymphs were predators with raptorial forelegs, which are reminiscent to those of

praying mantids Mantidae is one of the largest families in the order of praying mantises, based on the type species ''Mantis religiosa''; most genera are tropical or subtropical. Historically, this was the only family in the order, and many references still use ...

. The nymphs had a peculiar

freshwater shrimp Freshwater shrimp are any shrimp which live in fresh water. This includes: *Any Caridea (shrimp) which live in fresh water, especially the family Atyidae *Species in the genus '' Macrobrachium'' :*'' Macrobrachium ohione'', the Ohio River shrimp :* ...

-like habitus.

Etymology

The genus ''Mickoleitia'' and family Mickoleitiidae was named in honor of German zoologist Gerhard Mickoleit from the

University of Tübingen The University of Tübingen, officially the Eberhard Karl University of Tübingen (; ), is a public research university located in the city of Tübingen, Baden-Württemberg, Germany. The University of Tübingen is one of eleven German Excellenc ...

, who was among the first proponents of

Willi Hennig Emil Hans Willi Hennig (20 April 1913 – 5 November 1976) was a German biologist and zoologist who is considered the founder of phylogenetic systematics, otherwise known as cladistics. In 1945 as a prisoner of war, Hennig began work on his th ...

's "

Phylogenetic Systematics Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...

". The scientific name of the order Coxoplectoptera refers to the prolonged coxal segment of the nymphal and adult legs, and the old scientific name Plectoptera for mayflies (not to be confused with

Plecoptera Plecoptera is an order (biology), order of insects commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to b ...

for stoneflies). The common name "chimera wings" was coined in reference to the strange combination of characters in the morphology of the adult animal, which looks like a kind of

chimera Chimera, Chimaera, or Chimaira (Greek for " she-goat") originally referred to: * Chimera (mythology), a fire-breathing monster of ancient Lycia said to combine parts from multiple animals * Mount Chimaera, a fire-spewing region of Lycia or Cilicia ...

built from unrelated insects, with their oblique thorax and broad hind wing shape like a dragonfly, their wing venation like a primitive mayfly ancestor, and their raptorial forelegs like a mantis.

History of discovery

The fossil nymphs of the genus ''Mickoleitia'' are not especially rare in the limestones of the Crato Formation; the local brick workers even have a common Brazilian name for them ("Abacaxi" =

pineapple The pineapple (''Ananas comosus'') is a Tropical vegetation, tropical plant with an edible fruit; it is the most economically significant plant in the family Bromeliaceae. The pineapple is indigenous to South America, where it has been culti ...

). These nymphs were scientifically discovered and first mentioned by Bechly (2001: fig. 36), who also pointed to their strange morphology. Staniczek (2002, 2003) discussed the larvae as well and claimed that they arguably had been a kind of

living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...

in the Lower Cretaceous. The German biologist Rainer Willmann described the nymphs in a chapter in Martill, Bechly & Loveridge (2007) and erroneously attributed them to the extinct stem group mayfly family Cretereismatidae that he described based on adult specimens from the same locality. During the work for this monograph on the Crato Formation the German palaeoentomologist Günter Bechly and entomologist Arnold H. Staniczek discovered in the fossil collection of the Stuttgart State Museum of Natural History the very adult specimen that later would become the

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

of ''Mickoleitia longimanus''. They figured this fossil in Martill, Bechly & Loveridge 2007 (Fig. 11.90i,j) as undescribed stem group mayfly and indicated in a brief figure legend the possible relationship to the erratic nymphs. The detailed scientific description of Coxoplectoptera and the demonstration of the relationship of fossil adult and nymphs was finally published by Staniczek, Bechly & Godunko (2011) in a special issue on Cretaceous insects of the journal "Insect Systematics & Evolution". The authors also determined that two fossil nymphs (''Mesogenesia petersae'' = ''Archaeobehnigia edmundsi'') that had been erroneously described by Tshernova (1977) as modern mayfly nymphs from the

Middle Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ...

Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...

Transbaikal Transbaikal, Trans-Baikal, Transbaikalia ( rus, Забайка́лье, r=Zabaykal'ye, p=zəbɐjˈkalʲjɪ), or Dauria (, ''Dauriya'') is a mountainous region to the east of or "beyond" (trans-) Lake Baikal at the south side of the eastern Si ...

s, can be attributed the order Coxoplectoptera. The discovery of Coxoplectoptera represented one of the more spectacular findings of paleontology in 2011 and was heavily covered by news media around the globe.

Description

Adult

The adult stage of the type species ''Mickoleitia longimanus'' had a wing length of 28–29 mm and a probable body length of ca. 35–40 mm (the

abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...

is not preserved in the single known fossil holotype specimen). A second unnamed species of the genus ''Mickoleitia'' was only of half this size, and is only known by a single adult specimen from a private fossil collection in

Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...

. The head of ''Mickoleitia'' was provided with large

compound eyes A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...

and functional mouthparts (preserved are 3-segmented labial palps). The

thoracic The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main ...

segments are obliquely tilted backwards as in dragonflies, so that the raptorial forelegs are shifted forwards. All legs have a strongly prolonged and free coxal segment. The forelegs are developed as subchelate raptorial devices with a single-segmented tarsus with an unpaired claw. Most likely the abdomen was provided with three caudal filaments (two lateral cerci and the median epiproct) as in modern mayflies and their

Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...

stem group representatives ( Permoplectoptera, e.g. Protereismatidae). Since males of modern mayflies and of Permoplectoptera have gonopods on the 9th abdominal segment that are developed as genital claspers to grip the female for copulation, such a character state and behavior is also likely for Coxoplectoptera, who have an intermediate position as

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

link between these two groups.

Nymph

The more than 20 described nymphs of different stages have a body length of . Their laterally compressed body is unique among all known fossil and Recent

aquatic insect Aquatic insects or water insects live some portion of their life cycle in the water. They feed in the same ways as other insects. Some ''diving'' insects, such as predatory diving beetles, can hunt for food underwater where land-living insects ...

nymphs, and rather resembles the body of

gammarid Gammaridae is a family (biology), family of Amphipoda, amphipods. In North America they are included among the folk taxonomy, folk taxonomic category of "Scud (crustacean), scuds", and otherwise gammarids is usually used as a common name. They ...

freshwater shrimps. Many of the fossil nymphs are preserved in a characteristic posture with arched back, erect antennae and terminal filaments, and forelegs always in catching position similar to a praying mantis. The head was strongly armored and provided with horn- or shovel-like projections. Of the mouthparts only the crossed, sabre-like

mandibles In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...

and the spoon-shaped labium are known. All legs have a strongly prolonged and free coxal segment as in the adult. Likewise, the forelegs are developed as slender subchelate raptorial legs with nearly identical segment proportions as in the adult stage, but with a shorter tibia that may have been fused with the single-segmented tarsus, which ended in an unpaired claw. Styliform and ventrally directed abdominal gills are developed on abdominal segments 1-7. These gills are composed of a broader, more strongly sclerotized basal part and a slender and rather membranous distal part. The gills articulate dorsally within the abdominal

tergites A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'margin'. ...

that are distinctly separated from the ventral

sternites The sternum (: sterna) is the ventral portion of a segment of an arthropod thorax or abdomen. In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the sub ...

. The caudal filaments are formed by the two lateral cerci and the slightly longer medial terminal filament. All three appendages are lined with dense rows of long and thin setae.

Ecology and behavior

Adult

Because in the adult holotype specimen well-preserved mouthparts (palps) are visible, the adult animals almost certainly were able to feed. In direct contrast, the adult form of modern mayflies has dramatically reduced, non-functional mouthparts, and lives solely to reproduce. The raptorial forelegs and oblique thorax indicate that ''Mickoleitia'' was a predator. The large and broad hinds suggest that they were ecologically similar to dragonflies, in that they were swift, flying predators of other flying insects.

Nymph

The abundance of fossils, the circumstances of preservation and special anatomical

adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...

s (7 pairs of abdominal gills, 3 caudal filaments with dense rows of swimming hairs) prove that the larvae have been living in freshwater of streams and rivers, just like those of modern mayflies. They were washed in as

allochthon upright=1.6, Schematic overview of a thrust system. The hanging wall block is (when it has reasonable proportions) called a window. A klippe is a solitary outcrop of the nappe in the middle of autochthonous material. An allochthon, or an alloc ...

ous elements into the brackish Crato lagoon, were the limestones were deposited. The raptorial forelegs, sabre-like mandibles, large eyes and long antennae indicate that the nymphs were predators like the adults. On the other hand, the strong, shortened and broadened mid- and hind legs, the strong body armature, and shovel-like projections on the head all suggest that the animals were burrowing. Staniczek, Bechly & Godunko (2011) therefore assumed that the nymphs were

ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture their prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey u ...

s that were hiding, partly burrowed in the

river bed A streambed or stream bed is the bottom of a stream or river and is confined within a channel or the banks of the waterway. Usually, the bed does not contain terrestrial (land) vegetation and instead supports different types of aquatic vegeta ...

, and waiting for small prey passing by.

Evolution and phylogeny

The nymphs of Coxoplectoptera provided new clues to the disputed question of the evolutionary origin of insect wings. Before this discovery the paranotal-hypothesis and the leg-exite-hypothesis have been considered as incompatible alternative explanations, which have both been supported by a set of evidences from the

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

record,

comparative morphology Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing in t ...

developmental biology Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of Regeneration (biology), regeneration, asexual reproduction, metamorphosis, and the growth and di ...

and

genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...

. The expression of leg

gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...

s in the

ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the ovum, egg to adult. The term can also be used to refer to t ...

of the insect wing has been universally considered as conclusive evidence in favour of the leg-exite-hypothesis, which proposes that insect wings are derived from mobile leg appendages (exites). However, the larvae of Coxoplectoptera show that the abdominal gills of mayflies and their ancestors, which are generally considered as corresponding structures to insect wings, articulated within the dorsal tergite plates. This cannot be seen in modern mayfly nymphs, because their abdominal tergites and sternites are fused, without any traces of separation left even in embryonic development. If nymphal gills and wings are corresponding ("serial homologous") structures and thus share the same evolutionary origin, the new results from Coxoplectoptera demonstrate that also wings are of tergal origin, as proposed by the classical paranotal-hypothesis. Staniczek, Bechly & Godunko (2011) therefore suggested a new hypothesis that could reconcile the apparently conflicting evidence from

paleontology Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure ge ...

and

developmental genetics Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of stem ...

: wings originated as stiff outgrowths of tergal plates ( paranota), and only later in evolution became mobile, articulated appendages through secondary recruiting of leg genes. Within

pterygote Pterygota ( ) is a subclass of insects that includes all winged insects and groups who lost them secondarily. Pterygota group comprises 99.9% of all insects. The orders not included are the Archaeognatha (jumping bristletails) and the Zygent ...

insects the Coxoplectoptera represent the

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

of modern mayflies ( Ephemeroptera). This relationship is indicated by several

synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...

, such as: adult wing venation with costal brace (absent in other winged insects), nymphs with 7 pairs of abdominal gills (compared to still 9 pairs in Permoplectoptera like '' Protereisma'' nymphs), and with single-segmented tarsus with unpaired claw (compared to 3-segmented tarsus with paired claw in Permoplectoptera like ''Protereisma'' larvae). Together with mayflies and dragonflies they belong to the clade

Palaeoptera The name Palaeoptera (from Greek ( 'old') + ( 'wing')) has been traditionally applied to those ancestral groups of winged insects (most of them extinct) that lacked the ability to fold the wings back over the abdomen as characterizes the Neopt ...

, which is characterized by a derived wing articulation with fused sclerites, a vertical resting position of the wings in the

groundplan Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...

, and a

wing venation Insect wings are adult outgrowths of the insect exoskeleton that enable insect flight, insects to fly. They are found on the second and third Thorax (insect anatomy), thoracic segments (the mesothorax and metathorax), and the two pairs are often ...

with intercalary veins between the main longitudinal veins (esp. IR1+ between RP1- and RP2-, and IR2+ between RP2- and RP3/4-). Because of some very primitive character states, the Coxoplectoptera rather looked like early

Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...

ancestors of mayflies, e.g. in the wing venation of the adult stage they still had the elongate costal brace that is not fused to the costal margin, and in the nymphal stage they still had articulated lateral wing pads. The large and broad hind wings are a further

plesiomorphy In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral Phenotypic trait, character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorph ...

compared to the small hind wing of modern mayflies, and even compared to the slender hind wing of

stem group mayflies like ''Protereisma''. The monophyly of Coxoplectoptera is demonstrated by several autapomorphic characters in the adult stage, such as the raptorial forelegs and single-segmented tarsi with unpaired claw, as well as in the larval stage by the laterally compressed body, the body armature, the raptorial forelegs and burrowing mid- and hind legs, and the styliform shape of the ventrally directed abdominal gills. Coxoplectoptera are only known from the Jurassic and the Lower Cretaceous. It is not yet known why and when they went

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

Systematics

The

Coxoplectoptera contains a single

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Mickoleitiidae with two

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

from the

Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...

: ''Mickoleitia'' (

Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...

): * ''Mickoleitia longimanus'' (

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

) * ''Mickoleitia spec.'' (smaller unnamed species, represented by a single adult specimen in a private fossil collection in Japan) '' Mesogenesia'' (Middle or

Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...

s): * ''Mesogenesia petersae'' (= ''Archaeobehnigia edmundsi'')

References

Bibliography

* (1977): Distinctive new mayfly nymphs (Ephemeroptera; Palingeniidae, Behningiidae) from the Jurassic of Transbaikal. ''Paleontologicheskii Zhurnal'', 2: 91-96. (in Russian). * et al. (Hrsg.) (2001): Ur-Geziefer - Die faszinierende Evolution der Insekten. ''Stuttgarter Beiträge zur Naturkunde Serie C'', 49: 96 pp., Stuttgart. ISSN 0341-0161
PDF fulltext
. * (2002): Fossile Eintagsfliegen - Einblicke in die Welt urtümlicher Fluginsekten. ''Fossilien'', 19: 297-302. * (2003): Eintagsfliegen - Manna der Flüsse. ''Stuttgarter Beiträge zur Naturkunde Serie C'', 53: 80 pp., Stuttgart. ISSN 0341-0161. * (Hrsg.) (2007): ''The Crato Fossil Beds of Brazil - Window into an Ancient World.'' Cambridge University Press, Cambridge etc. . * (2011): Coxoplectoptera, a new fossil order of Palaeoptera (Arthropoda: Insecta), with comments on the phylogeny of the stem group of mayflies (Ephemeroptera). ''Insect Systematics & Evolution'', 42(2): 101-138, Brill, Leiden. ISSN 1399-560X

.

External links

*

* ttp://www.livescience.com/15100-insect-frakenstein-fossil-order-coxoplectoptera.html LiveScience: Ancient 'Frankenstein' Insect Discovered
Scientific American: Odd Insect Fossils Suggest Early Carnivorous Lifestyle

MSNBC: Fossil reveals an ancient 'Frankenstein' insect

Fox News: Ancient 'Frankenstein' Insect Discovered

YouTube: feature in German L-TV news magazine, incl. interviews

YouTube: feature in German Regio-TV news journal, incl. interviews
{{Taxonbar, from1=Q1073158, from2=Q21447579 Extinct insect orders Tithonian first appearances Early Cretaceous extinctions