The name Apterygota is sometimes applied to a subclass of small, agile

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...

s, distinguished from other insects by their lack of wings in the present and in their evolutionary history; notable examples are the silverfish, the

firebrat The firebrat (''Thermobia domestica'') is a small hexapod (typically 1–1.5 cm), in the order Zygentoma. Habitat Firebrats prefer higher temperatures and require some humidity, and can be found in bakeries and near boilers or f ...

, and the jumping bristletails. Their first known occurrence in the fossil record is during the Devonian period, 417–354 million years ago. The group Apterygota is not a clade; it is paraphyletic, and not universally recognized. As defined, the group contains two separate

clades A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

of wingless insects:

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 ...

comprises jumping bristletails, while

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 t ...

comprises silverfish and firebrats. The Zygentoma are in the clade

Dicondylia 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 a hypothetical ancestral m ...

with winged insects, a clade that includes all other insects, while Archaeognatha is sister to this lineage.A. Blanke, M. Koch, B. Wipfler, F. Wilde, B. Misof (2014) Head morphology of ''Tricholepidion gertschi'' indicates monophyletic Zygentoma. Frontiers in Zoology 11:16 doi:10.1186/1742-9994-11-16 The

nymphs A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...

(younger stages) go through little or even no metamorphosis, hence they resemble the adult specimens. Their skin is thin, making them appear translucent. Currently, no

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

are listed as being at conservation risk.

Characteristics

The primary characteristic of the apterygotes is they are primitively wingless. While some other insects, such as

flea Flea, the common name for the order Siphonaptera, includes 2,500 species of small flightless insects that live as external parasites of mammals and birds. Fleas live by ingesting the blood of their hosts. Adult fleas grow to about long, a ...

s, also lack wings, they nonetheless descended from winged insects but have lost them during the course of evolution. By contrast, the apterygotes are a primitive group of insects that diverged from other ancient orders before wings evolved. Apterygotes, however, have the demonstrated capacity for directed, aerial gliding descent from heights. It has been suggested by researchers that this evolved gliding mechanism in apterygotes might have provided an evolutionary basis from which winged insects would later evolve the capability for powered flight. Apterygotes also have a number of other primitive features not shared with other insects. Males deposit sperm packages, or

spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...

s, rather than fertilizing the female internally. When hatched, the young closely resemble adults and do not undergo any significant metamorphosis, and lack even an identifiable nymphal stage. They continue to molt throughout life, undergoing multiple

instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...

s after reaching sexual maturity, whereas all other insects undergo only a single instar when sexually mature. Apterygotes possess small unsegmented appendages, referred to as "styli", on some of their

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

segments, but play no part in locomotion. They also have long, paired abdominal cerci and a single median, tail-like caudal filament, or telson.

History of the concept

The composition and classification of Apterygota changed over time. By the mid-20th century, the subclass included four orders (

Collembola Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...

Protura The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. The Protura constitute an order of hexapods that were previously ...

Diplura The order Diplura ("two-pronged bristletails") is one of three orders of non-insect hexapods within the class Entognatha (alongside Collembola (springtails) and Protura). The name "diplura", or "two tails", refers to the characteristic pair of ...

, and

Thysanura Thysanura is the now deprecated name of what was, for over a century, recognised as an order in the class Insecta. The two constituent groups within the former order, the Archaeognatha (jumping bristletails) and the Zygentoma (silverfish and fireb ...

). With the advent of a more rigorous

cladistic Cladistics (; ) 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 typically shared derived char ...

methodology, the subclass was proven paraphyletic. While the first three groups formed a monophyletic group, the

Entognatha The Entognatha are a class of wingless and ametabolous arthropods, which, together with the insects, makes up the subphylum Hexapoda. Their mouthparts are entognathous, meaning that they are retracted within the head, unlike the insects. Entog ...

, distinguished by having mouthparts submerged in a pocket formed by the lateral and ventral parts of the head capsule, the Thysanura (

plus

) appeared to be more closely related to winged insects. The most notable

synapomorphy 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 hav ...

proving the

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

of Thysanura+Pterygota is the absence of intrinsic antennal muscles, which connect the antennomeres in entognaths,

myriapods Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, ...

, and

crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...

s. For this reason, the whole group is often termed the Amyocerata, meaning "lacking antennal muscles". However, the Zygentoma are now considered more closely related to the Pterygota than to the Archaeognatha, thus rendering even the amyocerate apterygotes paraphyletic, and resulting in the dissolution of Thysanura into two separate monophyletic orders.

References

* ''Firefly Encyclopedia of Insects and Spiders'', edited by Christopher O'Toole, , 2002 {{Use dmy dates, date=April 2020 Insect taxonomy Arthropod subclasses Extant Devonian first appearances Paraphyletic groups