Blattoptera, or proto-cockroaches, is a name given to various "roachid" fossil insects related to cockroaches, mantises and termites, and of general cockroach-like appearance and possibly habit.[1] The group is on the rank of an order, though being paraphyletic is most often given without formal taxonomic rank. Several alternative names have been suggested for this fossil group, including Blattodea,[1] a name currently used for the group including the modern cockroaches as well as their fossil relatives.[2]

Temporal range: Carboniferous–Late Permian
Scientific classification

Systematic position

Cockroaches are popularly thought to be an ancient order of insects, with their origins in the Carboniferous.[3] However, since the middle of the 20th century it has been known that the primitive cockroach insects found fossilized in Palaeozoic strata are the forerunners not only of modern cockroaches but also of mantises and termites.[2] The origin of these groups from a blattopteran stock are now generally thought to be in the early Jurassic; the earliest modern cockroach found is from the Cretaceous. Thus the “Palaeozoic cockroaches” are not cockroaches per se, but a paraphyletic assemblage of primitive relatives.[4]

Anatomy and habits

The fossils assigned to the "roachids" are of general cockroach-like build, with a large disc-like pronotum covering most of the head, long antennae, legs built for running, flattened body and heavily veined wings with the distinct arched CuP-vein so typical of modern cockroach wings.[5] Like modern cockroaches, the roachids were probably swift litter inhabitants living on a wide range of dead plant and animal matter.

Contrary to modern forms, the female roachids all have a well-developed external ovipositor, a primitive insect trait.[6] They probably inserted eggs singly into soil or crevices. The egg pods, called ootheca, seen in modern cockroaches and their relatives is a new shared trait separating them from their primitive ancestors. Some of the roachid species could reach relatively large sizes compared to most of their modern relatives, like the Carboniferous Archimylacris and the Permian Apthoroblattina, the latter who could reach 50 mm in body length. However, none that we know of reached near the size of today's largest tropical species, the modern 'giant cockroaches', such as the famous Hissing Cockroach.


  1. ^ a b Henning, W. (1981): Insect Phylogeny. Wiley, Chichester, Britain
  2. ^ a b Grimaldi, D (1997): A fossil mantis (Insecta: Mantoidea) in Cretaceous amber of New Jersey, with comments on early history of Dictyoptera. American Museum Novitates 3204: 1-11
  3. ^ Guthrie, D. M. & A. R. Tindal (1968): The Biology of the Cockroach. St. Martin's Press, New York
  4. ^ Grimaldi, D. & M. S. Engel, Michael (2005): Evolution of the Insects, Cambridge University Press, ISBN 0-521-82149-5
  5. ^ Schneider, J. (1983): Die Blattodea (Insecta) des Paleozoicums, Teil II, Morphogenese des Flügelstrukturen und Phylogenie. Freiberger Forchnungshefte, Reie C 391. pp 5-34
  6. ^ Grimaldi, D. & M. S. Engel, Michael (2005): Evolution of the Insects, Cambridge University Press, p 227

Amphiesmenoptera is an insect superorder, established by S. G. Kiriakoff, but often credited to Willi Hennig in his revision of insect taxonomy for two sister orders: Lepidoptera (butterflies and moths) and Trichoptera (caddisflies). In 2017, a third fossil order was added to the group, the Tarachoptera.Trichoptera and Lepidoptera share a number of derived characters (synapomorphies) which demonstrate their common descent:

Females, rather than males, are heterogametic (i.e. their sex chromosomes differ).

Dense setae are present in the wings (modified into scales in Lepidoptera).

There is a particular venation pattern on the forewings (the double-looped anal veins).

Larvae have mouth structures and glands to make and manipulate silk.Thus these two extant orders are sisters, with Tarachoptera basal to both groups. Amphiesmenoptera probably evolved in the Jurassic. Lepidoptera differ from the Trichoptera in several features, including wing venation, form of the scales on the wings, loss of the cerci, loss of an ocellus, and changes to the legs.Amphiesmenoptera are thought to be the sister group of Antliophora, a proposed superorder comprising Diptera (flies), Siphonaptera (fleas) and Mecoptera (scorpionflies). Together, Amphiesmenoptera and Antliophora compose the group Mecopterida.


Archodonata is an extinct order of palaeozoic paleopterous insects, sometimes included in Odonata.


Coniopterygoidea is a lacewing superfamily in the suborder Hemerobiiformia. In some classifications, Coniopterygoidea is an expansion used to signify that the spongillaflies (Sisyridae) and the dustywings (Coniopterygidae) are each other's closest relatives.


The Diaphanopterodea or Paramegasecoptera are an extinct order of moderate to large-sized Palaeozoic insects. They are first known from the Middle Carboniferous (late Serpukhovian or early Bashkirian in age), and include some of the earliest known flying insects.


The Dicondylia are a taxonomic group (taxon) that includes all insects except the jumping bristletails (Archaeognatha). Dicondylia have a mandible attached with two hinges to the head capsule (dicondyl), in contrast to the original mandible with a single ball joint (monocondyl).


Dictyoptera (from Greek δίκτυον diktyon "net" and πτερόν pteron "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the order Mantodea (mantises), along with one extinct order, the Alienoptera. While all modern Dictyoptera have short ovipositors, the oldest fossils of Dictyoptera have long ovipositors, much like members of the Orthoptera.


Endopterygota (from Ancient Greek endon “inner” + pterón, “wing” + New Latin -ota “having”), also known as Holometabola, is a superorder of insects within the infraclass Neoptera that go through distinctive larval, pupal, and adult stages. They undergo a radical metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour. This is called holometabolism, or complete metamorphism.

The Endopterygota are among the most diverse insect superorders, with over 1 million living species divided between 11 orders, containing insects such as butterflies, flies, fleas, bees, ants, and beetles.They are distinguished from the Exopterygota (or Hemipterodea) by the way in which their wings develop. Endopterygota (meaning literally "internal winged forms") develop wings inside the body and undergo an elaborate metamorphosis involving a pupal stage. Exopterygota ("external winged forms") develop wings on the outside their bodies and do not go through a pupal stage. The latter trait is plesiomorphic, however, and not exclusively found in the exopterygotes, but also in groups such as Odonata (dragonflies and damselflies), which are not Neoptera, but more basal among insects.

The earliest endopterygote fossils date from the Carboniferous.


Eumetabola is an unranked category of Neoptera. Two large unities known as the Paurometabola and Eumetabola are probably from the adelphotaxa of the Neoptera after exclusion of the Plecoptera. The monophyly of these unities appears to be weakly justified.


The Ithonioidea are a superfamily of lacewing insects in the order Hemerobiiformia.


Myrmeleontoidea is a lacewing superfamily in the suborder Myrmeleontiformia.


Neoptera is a classification group that includes most orders of the winged insects, specifically those that can flex their wings over their abdomens. This is in contrast with the more basal orders of winged insects (the "Palaeoptera" assemblage), which are unable to flex their wings in this way.


The Odonatoptera are a superorder (sometimes treated as an order) of ancient winged insects, placed in the Palaeoptera which probably form a paraphyletic group however. The dragonflies and damselflies are the only living members of this group, which was far more diverse in the late Paleozoic and contained gigantic species, including the griffinflies (colloquially called "giant dragonflies", although they were not dragonflies in the strict sense) of the order Protodonata. This lineage dates back at least to the Bashkirian, not quite 320 million years ago.


The name Palaeoptera 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 Neoptera. The Diaphanopterodea, which are palaeopteran insects, had independently and uniquely evolved a different wing-folding mechanism. Both mayflies and dragonflies lack any of the smell centers in their brain found in Neoptera.


Panorpida or Mecopterida is a proposed superorder of Endopterygota. The conjectured monophyly of the Panorpida is historically based on morphological evidence, namely the reduction or loss of the ovipositor and several internal characteristics, including a muscle connecting a pleuron and the first axillary sclerite at the base of the wing, various features of the larval maxilla and labium, and basal fusion of CuP and A1 veins in the hind wings. The monophyly of the Panorpida is also supported by recent molecular data.


Paraneoptera is a monophyletic superorder of insects which includes four orders, the bark lice, true lice, thrips, and hemipterans, the true bugs. The mouthparts of the Paraneoptera reflect diverse feeding habits. Basal groups are microbial surface feeders, whereas more advanced groups feed on plant or animal fluids.


Protelytroptera is an extinct order of insects thought to be a stem group from which the modern Dermaptera evolved. These insects, which resemble modern Blattodea, or Cockroaches, are known from the Permian of North America, Europe and Australia, from the fossils of their shell-like forewings and the large, unequal, anal fan. None of their fossils are known from the Triassic when the morphological changes from Protelytroptera to Dermaptera presumably took place.


Protodiptera is an extinct order of insects containing the two genera Permotipula and Permila.


Psocodea is a taxonomic group of insects comprising the bark lice, book lice and true lice. It was formerly considered a superorder, but is now generally considered by entomologists as an order. Despite the greatly differing appearance of lice, they are believed to have evolved from within the former order "Psocoptera", which contained the bark lice and book lice. Psocodea contains around 11,000 species, divided among seven suborders.


The Pterygota are a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless (that is, insect groups whose ancestors once had wings but that have lost them as a result of subsequent evolution).The pterygotan group comprises almost all insects. The insect orders not included are the Archaeognatha (jumping bristletails) and the Zygentoma (silverfishes and firebrats), two primitively wingless insect orders. Also not included are the three orders no longer considered to be insects: Protura, Collembola, and Diplura.

Insect orders


This page is based on a Wikipedia article written by authors (here).
Text is available under the CC BY-SA 3.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.