Centipede

Etymology and names

The scientific name Chilopoda derives from Ancient Greek kheilos, meaning lip, and podos, meaning foot, referencing the mouth-covering forcipules formed from modified legs.^[4][5][6] The common English name centipede originates from the Neo-Latin prefix centi-, meaning hundred, and pes, pedis, meaning foot.^[1] Despite the literal meaning of hundred feet, no species possesses exactly 100 legs, as the total number of leg-bearing segments in centipedes is always an odd number.^[1][7][8] Colloquial names in English include forty-legs and hundred-legs.^[9][10]

In Japanese and Chinese, centipedes are historically written with several kanji characters such as 百足, 蜈蜙, 蜈蚣, and 蝍蛆.^[11][12] In East Asia, scutigeromorphs are often distinguished under distinct vernacular terms, such as geji in Japanese and youyan in Chinese, separating them from other centipedes in popular usage.^[13] In traditional Chinese literature, they were also recorded under names including wugong, baizuchong (hundred-footed insect), baijiaochong, and tianlong (heavenly dragon).^[14][15][16]

Anatomy and morphology

Centipedes possess elongated bodies composed of two tagmata: a distinct head and an extended, segmented trunk.^[17] Body sizes range from a few millimetres in dwarf lithobiomorphs and geophilomorphs up to 30 cm in giant scolopendromorphs such as Scolopendra gigantea, though most species measure between 1 and 10 cm.^[2][18][19] The cuticle lacks the heavy calcification seen in millipedes and is comparatively soft.^[20] Typical pigmentation includes shades of drab brown, yellow, and reddish hues.^[20] Subterranean and troglobitic species frequently lack pigmentation entirely, while many tropical scolopendromorphs exhibit bright aposematic color patterns in red, orange, green, black, or violet.^[6][21]

Head and cephalic appendages

The head capsule is covered dorsally by a sclerotized cephalic plate, which is flattened in most orders but domed in Scutigeromorpha.^[22][23] The cephalic capsule is formed by the fusion of the acron (bearing eyes and upper lip) and five post-oral segments.^[17][24] As in other myriapods and hexapods, the second segment is an intercalary segment lacking appendages.^[24] The forward margin of the head bears a single pair of multi-articulated antennae.^[21][25] In Scutigeromorpha, these are flagelliform, whiplike, and subdivided into hundreds of minute annulations, whereas in other orders they are moniliform or beadlike, with segment counts varying from 14 (a fixed number in Geophilomorpha) up to more than 100.^[21][22][26]

Mouthparts situated ventrally comprise an anterior labrum attached to the clypeus, a pair of mandibles, and two pairs of maxillae.^[22][25][27] The mandibles are slender, triangular, and equipped with teeth and fine setae, enclosed within the mouth cavity and concealed from view by the maxillae.^[21][22] The first pair of maxillae forms a lower lip and carries short sensory palps.^[21][25] The coxae of the maxillae are frequently fused with the ventral sternite into a coxosternite that shields the posterior lower head.^[20][22] The second pair of maxillae is leglike with three to five segments; in scutigeromorphs, they are long and slender, while in other groups they are shorter, stouter, and tipped with small apical claws or dense brush-like bristles.^[20][21][22]

Visual organs vary considerably across the class. Scutigeromorphs possess a pair of true, faceted compound eyes featuring distinct crystalline cones analogous to those of insects and chelicerates.^[28][29][30] All other orders have lateral ocelli consisting of lenses formed from the cornea without crystalline cones, or lack eyes altogether.^[21] Lithobiomorpha possess from one to 49 pairs of ocelli, Scolopendromorpha usually have four pairs or one pair (in Mimops), and Craterostigmomorpha possess a single pair.^[31] All geophilomorphs and numerous cave-dwelling or burrowing species of other orders are entirely blind.^[2][31][32] These non-compound ocelli discern only light from dark rather than resolving distinct visual images.^[25][30]

At the base of the antennae, many lithobiomorphs and scutigeromorphs bear a specialized sensory structure termed the organ of Tömösváry.^[23][25] This consists of a discoid cuticular structure surrounding a central pore encircled by sensitive cells.^[25][33] It is rudimentary in Craterostigmomorpha and absent in Geophilomorpha and Scolopendromorpha.^[23][34] While historically interpreted as an auditory or vibration receptor, it has also been evaluated as a receptor for humidity, atmospheric pressure, and carbon dioxide concentrations.^[25][27][35]

Forcipules and venom apparatus

The defining synapomorphy of Chilopoda is the modification of the first pair of post-cephalic limbs into heavy, pincer-like venom claws known as forcipules, toxicognaths, or maxillipeds.^[1][2][36] Located on the first trunk segment directly beneath the head, they do not constitute oral mouthparts derived from the cephalon, but rather highly specialized raptorial thoracic limbs.^{[24][25][34][37]} The large coxae of the forcipules are fused with the sternite of the first segment into an expansive ventral plate called the coxosternite.^[20][38] In Scutigeromorpha, the coxae retain mobility from a vestigial sternite and bear long marginal setae, whereas in all other orders (Pleurostigmophora) the coxosternite is fully fused, immobile or barely movable, and armed anteriorly with stout chitinous teeth.^{[21][36][39][40]}

The movable appendage consists of four functional articles: the basal trochanteroprefemur (a fused trochanter and prefemur), the intermediate femur and tibia, and a terminal tarsungulum (a fusion of the tarsus and ungulum) that tapers to a sharp piercing claw.^[20][21][36] In Scutigeromorpha, the femur and tibia are complete cylindrical segments permitting vertical, stabbing movements, whereas in Scolopendromorpha and Geophilomorpha these middle articles are reduced to incomplete hemicylindrical rings lacking lateral walls, concentrating articulation onto a single pair of pivots that force the fangs to close horizontally in an opposable pincer-like motion.^[34][36][41] A venom gland resides inside the appendage (extending into the trunk in some geophilomorphs) and empties through a narrow duct opening just before the subterminal tip of the claw.^{[21][25][36][37]}

Trunk and locomotory appendages

Behind the forcipular segment, the body consists of a series of segments enclosed by dorsal sclerotized plates (tergites), ventral plates (sternites), and flexible lateral pleural membranes studded with small pleurites and subcoxal plates.^[20][21] In many orders, the trunk displays heterotergy, an alternation of longer and shorter tergites that provides longitudinal rigidity and dampens lateral undulations during rapid locomotion.^[40][42][43] In Lithobiomorpha, tergites over segments 2, 4, 6, 9, 11, 13, and 15 are visibly shortened.^[23] In Scutigeromorpha, only eight large dorsal plates cover the 15 leg-bearing segments.^[23] In Geophilomorpha, the tergites are homonomous, but each leg-bearing segment (except the first) is subdivided into an anterior pretergite (intercalary tergite) and a posterior metatergite, with corresponding subdivisions of the sternites into presternites and metasternites, conferring extensive telescopic flexibility for burrowing through substrate.^[21][22][23] In some European geophilomorphs, muscular contraction and elastic pleura allow the trunk to expand its length by up to 70 percent.^[44][45]

Each walking leg consists of six primary podomeres: coxa, trochanter, prefemur, femur, tibia, and tarsus, ending in a pointed apical pretarsal claw.^[21][23] The joint between the trochanter and prefemur is immovable, and in all orders except Geophilomorpha the trochanter possesses a pre-formed cleavage line enabling autotomy.^[21][46][47] In Scutigeromorpha, the legs are long and stilt-like with the tarsi subdivided into hundreds of flexible tarsomeres that can curl around objects or prey, whereas in other centipedes they are shorter, stouter, and conical.^[21][48][49] Successive leg pairs increase slightly in length toward the posterior end of the animal, which reduces the chance that adjacent legs collide and trip the centipede while it runs.^[25]

The final pair of appendages is modified into ultimate legs, also termed anal, caudal, or terminal legs, which project backward and are not used for locomotion.^{[49][50][51][52]} Ultimate legs exhibit diverse specializations: they may be elongated and antenna-like, thickened into defensive pincers, or flattened into leaf-like structures.^[49][52] In the African genus Alipes, these leaf-shaped legs can be stridulated against each other to produce a hissing, rattling sound that may distract or threaten predators, while Rhysida immarginata togoensis produces a faint creaking sound when swinging its rear limbs.^[53] In many species, ultimate legs bear glandular coxal organs, sensory pores, or distinctive tubercles and spines that are often sexually dimorphic, functioning in courtship, defense, or holding prey.^{[51][52][54][55]} In species of Lithobiomorpha, coxal pores situated on the ventral surface of coxae 12 through 15 produce sticky secretions and participate in osmoregulation and pheromone dissemination.^[27][54]

Posterior to the ultimate leg-bearing segment lie the reduced postpedal segments, consisting of two genital segments and an unsegmented terminal telson bearing the anal valves.^[17][21][56] The first genital segment carries the gonopods, which are typically much better developed in females as claspers or pincers for handling eggs, especially in Scutigeromorpha and Lithobiomorpha.^[23][56] The second genital segment houses the median gonopore, representing the vulva in females or the penis in males.^[20][56] In Scolopendromorpha, these genital segments are highly retracted and normally concealed entirely within the final pedigerous segment.^[21]

Internal anatomy and physiology

Centipedes breathe by means of an internal tracheal system opening to the exterior via spiracles.^[25] In the Notostigmophora (Scutigeromorpha), the spiracles are unpaired, slit-like, and located dorsally near the posterior margins of seven large tergites, opening into an internal atrium from which radiating bundles of short tracheae arise in an arrangement termed a tracheal lung, with hemocyanin dissolved in the blood performing oxygen transport.^{[57][58][59][60]} In all other centipedes (Pleurostigmophora), the spiracles are paired and situated laterally on the pleural membranes above the leg bases.^[23][61] These lateral spiracles typically occur on segments with long tergites (segments 3, 5, 8, 10, 12, and 14 in 15-legged forms) and lead to long, branching tracheae and tracheoles.^[21][25] In Geophilomorpha, paired spiracles are present on every leg-bearing segment except the first and ultimate pairs, and their internal branches form repeating X-shaped chiasmata across the midline.^[21][26] Some species are able to close their spiracles, and a few species in dry environments have evolved a waterproof cuticle.^[62][63]

The circulatory system is open, consisting of a tubular dorsal vessel located in the pericardial sinus that functions as the heart and pumps hemolymph forward toward the head.^[45][64] The heart extends through almost the entire trunk, containing over 20 chambers with lateral ostia in each segment, contracting at rates between 30 and 54 beats per minute depending on temperature, accelerated by adrenaline and acetylcholine and slowed by histamine.^[64] At its anterior end, the heart tapers into a cephalic aorta that enters the head capsule, where lateral vessels branch off and loop through a maxillipedal arch to connect with a ventral supraneural vessel running backward above the nerve cord.^[27][64] The hemolymph is colorless and contains five types of hemocytes that participate in immune defense, cellular nutrition, and clotting.^[64]

The digestive tract forms a straight tube running from the mouth to the posterior anus, divided into a foregut with esophagus and salivary glands, a midgut, and a hindgut.^[25] Waste excretion is carried out by a single pair (or occasionally two pairs) of slender Malpighian tubules that arise at the junction between the midgut and hindgut and empty into the proctodeum.^[25][60] Unlike terrestrial insects and arachnids that excrete insoluble uric acid, centipedes excrete the majority of their nitrogenous waste in the form of ammonia, an excretory pathway that requires extra water and adds to the water loss that ties them to moist microhabitats.^[60]

The nervous system conforms to the typical arthropod ladder plan, featuring a dorsal tripartite brain (protocerebrum, deutocerebrum, and tritocerebrum) connected via circumesophageal connectives to a ventral nerve cord.^[45][60] The protocerebrum integrates photoreception from the eyes but is severely reduced in blind subterranean species; the deutocerebrum innervates the antennae; and the tritocerebrum innervates the labrum and stomatogastric system.^[27][45][60] A large subesophageal ganglion, formed by the fusion of ganglia from the mouthpart segments, governs the mandibles and maxillae, followed immediately by an enlarged forcipular ganglion that directs the predatory strikes of the venom claws.^[27][45][60] Each subsequent body segment possesses a pair of fused ventral ganglia linked by longitudinal and transverse commissures.^[45]

Distinction from millipedes

Although both centipedes and millipedes belong to the subphylum Myriapoda and share multi-segmented bodies with postantennal sensory organs, they differ in morphology, ecology, and evolutionary history, having diverged between 450 and 475 million years ago.^[65][66] Centipedes bear a single pair of legs on each body segment attached laterally, whereas millipedes (Diplopoda) have two pairs of legs on most segments attached ventrally, resulting from the developmental fusion of adjacent somites into diplosegments.^[65][67] Centipedes have dorsoventrally flattened bodies, although house centipedes (Scutigeromorpha) have cylindrical bodies.^{[26][27][68][69]}

Ecologically, centipedes are venomous predators equipped with forcipules on their first trunk segment, whereas millipedes lack venomous fangs and are primarily slow-moving detritivores feeding on decomposing organic matter.^[65][67] Cephalic structures diverge markedly: millipede antennae are short and elbowed (geniculate) and their second maxillae are fused into a broad platelike gnathochilarium, whereas centipedes have long, threadlike or beadlike antennae, separate maxillae, and forcipules.^[67] Furthermore, millipedes are progoneate, bearing their genital openings on the third body segment, whereas centipedes are opisthogoneate, possessing their gonopores on the final genital segment at the rear of the body.^[70]

Reproduction and life cycle

Centipedes are dioecious arthropods that reproduce without copulation.^[71] Instead, sperm transfer occurs via an external spermatophore produced by the male.^[71] In many species, the male spins a small web of silk from glandular setae in the genital atrium, deposits the spermatophore onto it, and engages in a courtship ritual to guide the female.^[27] The courtship display often involves the partners circling each other for up to an hour while touching the other's terminal appendages with their antennae.^[49] The male deposits a spermatophore for the female to take up, and she receives it with her terminal genitalia.^[27][49][71] Parthenogenetic populations, composed exclusively of females, are known in several species.^[45][71]

In temperate regions, oviposition occurs in spring and summer, whereas tropical and subtropical species exhibit little reproductive seasonality.^[45][71][72] Centipedes display two distinct modes of egg-laying and maternal care.^[73] Females of Lithobiomorpha and Scutigeromorpha lay between 10 and 50 eggs singly in soil crevices, coat them with soil particles and mucus using their articulated gonopods to disguise them, and provide no subsequent parental care.^[45][48] In contrast, females of Geophilomorpha, Scolopendromorpha, and Craterostigmomorpha deposit clutches of 15 to 60 eggs in chambers hollowed out of rotting wood or soil.^[45] The mother curls her body around the clutch, remaining with the eggs and newly hatched young for weeks or months, continually grooming and licking them to prevent fungal infestation and desiccation.^[45][74] If disturbed by severe stress, the female may abandon or consume the clutch, and in several scolopendromorph species the young exhibit matriphagy, consuming their mother before dispersing.^[75]

Post-embryonic development follows either an anamorphic or an epimorphic pattern.^[2][76] In anamorphic development, characteristic of Scutigeromorpha, Lithobiomorpha, and Craterostigmomorpha, larvae hatch with fewer segments and leg pairs than the adult form, adding more segments and limbs during successive moults.^[2][76] In the house centipede Scutigera coleoptrata, the first-instar larva hatches with only four pairs of legs, passing through successive larval stadia with 5, 7, 9, 11, and finally 15 leg pairs, followed by post-larval stadia that develop gonopods and sensory organs.^[60][76][77] Lithobiomorph larvae typically hatch with seven pairs of legs (rarely six or eight) before reaching 15 pairs.^[2][60] Craterostigmomorpha represents a single-phase anamorphosis where the embryo hatches with 12 leg pairs and attains the adult count of 15 at the first moult.^[1][78][79]

In the clade Epimorpha, comprising Scolopendromorpha and Geophilomorpha, development is epimorphic: the embryos develop all body segments and leg pairs prior to hatching, emerging with the full complement of adult walking limbs.^[2][80] In Geophilomorpha, segment numbers vary widely within species and between sexes, with females typically possessing more segments than males.^[80][81] Segmentation in centipedes occurs in two distinct genetic phases: first, an odd number of anterior segments is established by Hox gene expression, and subsequently pairs of posterior segments are generated sequentially by a prepattern double-segment unit regulated by an oscillatory clock mechanism based on the Notch signaling pathway, which then cleaves into two segments.^[82][83][84] Centipedes are long-lived relative to insects; Lithobius forficatus can live five to six years, and Scolopendra subspinipes can exceed ten years.^[85][86][87] The combination of a small number of eggs, a long gestation period, and a long time of development to reproduction has led authors to label lithobiomorph centipedes as K-selected.^[88][89]

Ecology and behavior

Centipedes are predominantly nocturnal predators found in terrestrial environments across every continent except Antarctica, ranging from tropical rainforests to arid deserts, and extending north of the Arctic Circle.^[3][44][72] Because their cuticular exoskeleton lacks the protective outer wax layer present in insects and arachnids, they lose moisture rapidly through evaporation.^[90][91][92] They depend on humid microhabitats, sheltering during daylight beneath rotting logs, forest leaf litter, loose bark, compost piles, and stones.^[6][91] Some geophilomorphs inhabit coastal littoral zones, enduring tidal submersion by trapping air reserves around their coxal pores while foraging for barnacles and marine worms.^[93][94][95] Others are specialized troglobites with elongated antennae and reduced pigmentation, while a few species inhabit human structures.^[96] In Nigerian savanna, Scolopendra morsitans was recorded at densities of 0.16 individuals per square metre with a biomass of 140 mg per square metre wet weight, while small geophilomorphs in temperate forest soils can reach densities of up to 600 individuals per square metre with a biomass of 500 mg per square metre.^[97]

Centipedes are generalist carnivores that locate prey primarily through tactile and chemical cues detected by their antennae.^[25][98][99] Common prey items include earthworms, fly larvae, springtails, beetles, slugs, snails, and spiders.^{[27][100][101]} While plant matter is sometimes ingested by starving individuals in laboratory settings, gut analyses suggest that plant tissues do not form an important component of their natural diet.^{[72][98][102]} When prey is encountered, the centipede lashes forward in a rapid snake-like strike, grasping the quarry with its legs and sinking its forcipules to inject paralyzing venom.^[34][103] Digestion begins extra-orally as salivary secretions and venom proteases break down the victim's tissues before the food is cut into pieces by the mandibles and transferred into the mouth cavity.^[103]

Large scolopendromorph centipedes, particularly in the genera Scolopendra and Ethmostigmus, capture sizable vertebrates.^{[104][105][106]} The South American Scolopendra gigantea preys on tarantulas, scorpions, tree frogs, lizards, rodents, snakes, and even bats, which it captures by dangling from cave ceilings using its clawed rear legs.^{[107][108][109]} On an uninhabited island south of Norfolk Island, Cormocephalus coynei was found to derive 48 percent of its dietary intake from vertebrates, and it has been suggested that it functions as an apex predator in the local island ecosystem.^[106] Three tropical species, Scolopendra cataracta, Scolopendra paradoxa, and Scolopendra alcyona, are amphibious, swimming with undulating lateral movements to hunt aquatic invertebrates and small vertebrates in freshwater streams.^{[110][111][112]} Specialization is rare, though the genus Edentistoma is reported to feed exclusively on millipedes.^[34]

Centipedes are preyed upon by mongooses, rodents, shrews, salamanders, toads, predatory beetles, chickens, and birds such as the European roller.^{[102][113][114]} Certain predators specialize on centipedes, notably the South African Cape black-headed snake Aparallactus capensis, which feeds almost exclusively on them, and the African ant Amblyopone pluto, whose diet consists entirely of geophilomorphs.^[113][115] In response, centipedes employ a range of defensive adaptations. Species of Geophilomorpha, Lithobiomorpha, and Scolopendromorpha possess defensive repugnatorial glands.^{[116][117][118]} Geophilomorphs secrete viscous, odorous fluids containing hydrogen cyanide and benzoic acid from ventral sternal glands to foul attackers, and Scolopendra secretions similarly contain cyanide compounds.^{[119][120][121]} Lithobiomorphs produce sticky secretions from glands on their posterior legs that gum up the mouthparts of attacking ants and spiders.^[49][116] Many species assume a threat posture, elevating and splaying their ultimate legs to display spines or flash aposematic warning coloration, as seen in the contrasting black, orange, and yellow patterns of Scolopendra polymorpha.^{[53][74][122]} Scutigeromorphs and lithobiomorphs readily shed legs via autotomy along trochanteral fracture lines to escape grasping predators.^[47]

Evolution and phylogeny

The fossil record of centipedes dates to the Late Silurian, approximately 430 to 418 million years ago.^{[46][123][124]} The earliest confirmed centipede fossils consist of isolated leg remains attributed to the scutigeromorph genus Crussolum from the Late Silurian of Britain and the Early Devonian Rhynie chert of Scotland.^{[123][125][126]} The fossil head material of Rhyniognatha from the Rhynie chert, originally described as the earliest insect, has been reinterpreted as possibly belonging to a scutigeromorph myriapod.^[127][128] From the Middle Devonian Panther Mountain Formation in New York, two distinct taxa are documented: Crussolum and Devonobius delta, the latter placed in its own extinct order Devonobiomorpha.^[125][129] Carboniferous strata, including the Mazon Creek fossil beds (309–307 Ma), have yielded Latzelia (Scutigeromorpha), Mazoscolopendra, and Palenarthrus (Scolopendromorpha).^[130][131] By the Cretaceous period, fossil centipedes belonging to modern families of Scutigeromorpha and Scolopendromorpha appear preserved in amber.^[132][133]

Centipedes represent the most basal extant branch within the subphylum Myriapoda, which forms part of the Mandibulata alongside Pancrustacea (crustaceans and hexapods).^[134][135] All living centipedes form a monophyletic group united by four distinct synapomorphies: transformation of the first post-cephalic appendages into venomous forcipules, presence of an egg tooth on the embryonic cuticle of the second maxilliped, an immovable trochanter-prefemur articulation on the walking legs, and a helical spiral ridge along the nucleus of the spermatozoon.^{[136][137][138]} Molecular and developmental analyses place Chilopoda as the sister group to the Progoneata (Diplopoda, Pauropoda, and Symphyla), distinguished by the rearward position of the genital openings (opisthogony).^[70]

Chilopoda is divided into two primary subclasses: Notostigmophora, containing only the Scutigeromorpha, and Pleurostigmophora, containing the remaining four extant orders and Devonobiomorpha.^[65] Pleurostigmophora is diagnosed by paired lateral spiracles, flattened head capsules, fused forcipular coxosternites, and the presence of coxal organs.^[41][54] Within Pleurostigmophora, morphological phylogenies traditionally recognized the clade Phylactometria, comprising Craterostigmomorpha, Scolopendromorpha, and Geophilomorpha, united by maternal care of eggs and young and ventral glandular pores.^[139] Scolopendromorpha and Geophilomorpha form the well-supported monophyletic clade Epimorpha, characterized by epimorphic development, hemicylindrical forcipular articles, and 21 or more leg pairs.^{[32][36][136]} While traditional morphology viewed the anamorphic orders as a grade (Anamorpha), modern phylogenetics treats Anamorpha as paraphyletic.^[136][140] Some molecular transcriptomic studies challenge Phylactometria, proposing the clade Amalpighiata in which Lithobiomorpha is recovered as the sister group to Epimorpha, placing Craterostigmomorpha in a more basal position within Pleurostigmophora.^{[141][142][143]}

The evolution of centipede venom represents one of the oldest venom systems among terrestrial animals, originating prior to the divergence of scorpions and spiders over 420 million years ago.^{[32][144][145]} Over the first 50 million years of chilopod history, the venom cocktail appears to have comprised approximately four ancestral components, diversifying into complex, specialized toxins only after the modern orders diverged.^[146] Centipede venom evolution has incorporated foreign genes via horizontal gene transfer from bacteria, fungi, and oomycetes.^[146]

Classification and diversity

The class Chilopoda encompasses five extant orders containing 21 to 24 recognized families, approximately 339 genera, and over 3,000 described species, with global species richness estimated at 7,000 to 8,000 species.^{[44][147][148]} An extinct sixth order, Devonobiomorpha, is known from the Devonian.^[46][125] Extant diversity is distributed unevenly among orders with distinct morphology and habits.^[2][44]

Scutigeromorpha

Scutigeromorpha (house centipedes) represents the only living order of the subclass Notostigmophora, encompassing approximately 100 species across three families: Scutigeridae, Psellioididae, and Scutigerinidae.^[2] They are fast-running, anamorphic centipedes with 15 pairs of extraordinarily long, multi-tarsate walking legs, a domed cephalic plate, true compound eyes, long multi-segmented whiplike antennae, and seven unpaired dorsal spiracles opening from large tergites.^[2][57] Scutigera coleoptrata is a well-known cosmopolitan species native to the Mediterranean basin that lives in human dwellings, where it preys on flies, moths, and cockroaches.^[2][76]

Lithobiomorpha

Lithobiomorpha (stone centipedes) includes approximately 1,200 to 1,500 described species in two primary families, Lithobiidae and Henicopidae (with some classifications adding Anopsobiidae and Cermatobiidae).^[26][149] Adults measure 3 to 6 cm in length, possess 15 leg-bearing segments covered by alternating long and short tergites, paired lateral spiracles, beadlike antennae of 15 to 111 segments, and lateral clusters of ocelli (up to 49 per side) or no eyes.^[21] They are anamorphic, hatching with seven (or six to eight) leg pairs, and mature females carry three-segmented clawed gonopods used to handle individual eggs.^[2][21][60] Common European and holarctic species include Lithobius forficatus and Lithobius melanops.^[85]

Craterostigmomorpha

Craterostigmomorpha is the least diverse extant order, containing a single family, Craterostigmidae, with only two described species in the genus Craterostigmus: Craterostigmus tasmanianus in Tasmania and Craterostigmus crabilli in New Zealand.^[2][79] They possess 15 leg-bearing segments covered by 21 distinct dorsal tergites due to secondary subdivision of long tergites, a vertically elongated head with a single pair of ocelli, and prominent forward-projecting forcipules.^[2][78][150] They exhibit an intermediate developmental mode with a single anamorphic moult, hatching with 12 pairs of legs and acquiring 15 pairs at the first moult.^[1][79] Maternal brooding unites them with Epimorpha in the clade Phylactometria.^[151]

Scolopendromorpha

Scolopendromorpha (tropical or bark centipedes) comprises approximately 800 described species classified into five families: Scolopendridae, Cryptopidae, Scolopocryptopidae, Plutoniumidae, and Mimopidae.^[2][26] They possess 21 or 23 leg-bearing segments, with the sole exception of the Neotropical species Scolopendropsis duplicata, which possesses 39 or 43 pairs of legs.^[152][153] Scolopendromorphs typically have four pairs of ocelli, although some species have one pair and some are blind.^[30][31] Development is epimorphic, and females brood clutches of eggs.^[45][80] This order includes the largest living centipedes, such as Scolopendra gigantea (reaching 30 cm) and Scolopendra heros.^[18][26] In Plutonium zwierleini, spiracles are present on every leg-bearing segment, an exception to the normal alternation in the order.^[21]

Geophilomorpha

Geophilomorpha (soil centipedes) is the most species-rich order, with approximately 1,260 to 1,800 described species across seven to sixteen families.^[26] Morphologically and molecularly, they are divided into two suborders: Placodesmata (containing only Mecistocephalidae) and Adesmata (containing all other families, including Geophilidae, Himantariidae, Schendylidae, and Oryidae).^[26] All geophilomorphs are blind, subterranean burrowers with slender, worm-like bodies bearing between 27 and 191 pairs of legs, antennae fixed at 14 segments, and paired spiracles on all leg-bearing segments except the first and ultimate.^[1][26] In species such as Himantarium gabrielis, leg counts vary intraspecifically between 87 and 177 segments.^[26] Females care for the eggs and young, and development is epimorphic.^[2][45][80]

Devonobiomorpha

The extinct order Devonobiomorpha is established solely from fossil specimens of Devonobius delta discovered in the Middle Devonian Panther Mountain Formation of Gilboa, New York.^{[46][125][129]} The fossils display a flattened, eyeless cephalic plate, multi-segmented antennae with an elongated apical article, alternating tergite lengths, and forcipules featuring paired internal apodemes on the coxosternite, setting them apart from all extant centipedes.^[40] The number of leg-bearing segments exceeded 16, but complete specimens remain unknown.^[40] Sternal pores identified on Devonobius indicate affinities with Phylactometria.^[151]

Interaction with humans

Some species can inflict painful bites on humans using their forcipules.^[154] While bites from small centipedes cannot pierce human skin, bites from large scolopendromorphs cause intense localized pain, significant edema, erythema, chills, fever, headache, nausea, and localized necrosis.^[154][155] Centipede venom contains cardiotoxic, neurotoxic, and myotoxic compounds along with pain-inducing serotonin, histamine, and cytolysins, which in rare instances provoke anaphylactic shock in sensitized individuals or children.^[154][156] Confirmed fatalities are rare; documented cases include a girl in the Philippines and a 39-year-old Thai man who died after swallowing a live centipede.^{[156][157][158][159][160]} Standard first aid involves washing the wound with soap and water, applying cold compresses or ice to alleviate swelling, and administering analgesics, antihistamines, and tetanus prophylaxis if required.^{[140][161][162]}

In Chinese culture, large centipedes are prepared as street food, skewered and deep-fried or grilled, at vendors in Donghuamen and Wangfujing markets in Beijing.^{[163][164][165][166]} In China, Laos, Thailand, and Cambodia, large centipedes are steeped in high-proof alcohol to make centipede wine or vodka, believed in traditional Chinese medicine to possess medicinal properties for dispelling wind, arresting convulsions, and detoxification.^[167][168] In Chinese pharmacopeia, the dried whole body of Scolopendra subspinipes mutilans is recognized under the medicinal name Wugong.^[169] Indigenous Australians also traditionally used scolopendrid centipedes as an ingredient in seasonings.^[170] Large scolopendrid species are traded internationally as terrarium pets, with the largest specimens commanding high prices among hobbyists.^[19]

Centipedes hold prominent roles in folklore, religion, and military heraldry. In Sengoku-period Japan, samurai adopted centipede motifs on helmet crests (maedate), flags, and armor because centipedes were believed to never retreat, were associated with aggression, and had many legs symbolizing an army with numerous soldiers.^[171] According to the Kōyō Gunkan, the gold-mining sappers of the Takeda clan, who specialized in tunnel warfare, are said to have been called the Mukade-shū (centipede brigade).^[172] In Japanese mythology, centipedes are associated with the mountain god of Mount Akagi and the warrior Fujiwara no Hidesato, who defeated the giant centipede of Mount Mikami.^[171][173] Merchants also used centipede emblems on store banners because many legs suggested high customer foot traffic (kyakuashi).^[171][174] A 19th-century Tibetan Buddhist poem cautioned that anyone who enjoys frightening others will be reborn as a centipede.^[175][176]

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