Centipede


Centipedes are predatory arthropods belonging to the class Chilopoda of the subphylum Myriapoda, an arthropod group which includes millipedes and other multi-legged animals. Centipedes are elongated segmented animals with one pair of legs per body segment. All centipedes are venomous and can inflict painful stings, injecting their venom through pincer-like appendages known as forcipules or toxicognaths, which are actually modified legs instead of fangs. Despite the name, no species of centipede has exactly 100 legs; the number of pairs of legs is an odd number that ranges from 15 pairs to 191 pairs.
Centipedes are predominantly generalist carnivores, hunting for a variety of prey items that can be overpowered. They have a wide geographical range, which can be found in terrestrial habitats from tropical rainforests to deserts. Within these habitats, centipedes require a moist microhabitat because they lack the waxy cuticle of insects and arachnids, causing them to rapidly lose water. Accordingly, they avoid direct sunlight by staying under cover or by being active at night.

Description

Centipedes have a rounded or flattened head, bearing a pair of antennae at the forward margin. They have a pair of elongated mandibles, and two pairs of maxillae. The first pair of maxillae form the lower lip, and bear short palps. The first pair of limbs stretch forward from the body over the mouth. These limbs, or forcipules, end in sharp claws and include venom glands that help the animal to kill or paralyze its prey.
Their size ranges from a few millimetres in the smaller lithobiomorphs and geophilomorphs to about in the largest scolopendromorphs.

Sensory organs

Many species of centipedes lack eyes. Some lack one only, but some possess a variable number of ocelli, sometimes clustered together to form true compound eyes. However, these eyes are only capable of discerning light from dark, and provide no true vision. In some species, the first pair of legs can function as sensory organs, similar to antennae; unlike the antennae of most other invertebrates, these point backwards. An unusual clustering of sensory organs found in some centipedes is the organ of Tömösváry. The organs, at the base of the antennae, consist of a disc-like structure and a central pore, with an encircling of sensitive cells. They are likely used for sensing vibrations, and may provide a weak form of hearing.

Forcipules

Forcipules are unique to centipedes. The forcipules are modifications of the first pair of legs, forming a pincer-like appendage, just behind the head. Forcipules are not oral mouthparts, though they are used to subdue prey by injecting venom and gripping the prey animal. Venom glands run through a tube, from inside the head to the tip of each forcipule.

Body

Behind the head, the body consists of at least fifteen segments. Most of the segments bear a single pair of legs; the maxillipeds project forward from the first body segment, while the final two segments are small and legless. Each pair of legs is slightly longer than the pair preceding them, ensuring that they do not overlap, which reduces the chance that they will collide and trip the animal. The last pair of legs may be as much as twice the length of the first pair. The final segment bears a telson, and includes the openings of the reproductive organs.
Centipedes mainly use their antennae to seek out their prey. The digestive tract forms a simple tube, with digestive glands attached to the mouthparts. Like insects, centipedes breathe through a tracheal system, typically with a pair of openings, or spiracles, on each body segment. The spiracles lead to long tracheae and tracheoles that mediate oxygen transport to the organs, except for in Scutigeromorpha. In this order the spiracles are unpaired and the tracheae short, and oxygen supply is performed exclusively by the respiratory pigment hemocyanin. Some species are able to close their spiracles, and a few others in dry environments have evolved a waterproof cuticle. They excrete waste through a single pair of malpighian tubules.

Ultimate legs

Just as the first pair of legs are modified into forcipules, the back legs are modified into "ultimate legs", also called anal legs, caudal legs, and terminal legs. Their use varies between species, but does not include locomotion. The ultimate legs may be elongated and thin, thickened, or pincer-like. They are frequently sexually dimorphic, and may play a role in mating rituals. Because glandular pores occur more frequently on ultimate legs than on the "walking" legs, they may serve a sensory role. They are sometimes used in defensive postures, and some species use them to capture prey, defend themselves against predators, or suspend themselves from objects such as branches, using the legs as pincers. Several species use their ultimate legs upon encountering another centipede, trying to grab the body of the other centipede.
Members of the genus Alipes can stridulate their leaf-like ultimate legs to distract or threaten predators. Rhysida immarginata togoensis makes a faint creaking sound when it swings its ultimate legs.

Distinction from millipedes

There are many differences between millipedes and centipedes. Both groups of myriapods have long, multi-segmented bodies, many legs, a single pair of antennae, and the presence of postantennal organs. Centipedes have one pair of legs per segment, while millipedes have two. Their heads differ in that millipedes have short, elbowed antennae, a pair of robust mandibles and a single pair of maxillae fused into a lip; centipedes have long, threadlike antennae, a pair of small mandibles, two pairs of maxillae and a pair of large venom claws.
TraitMillipedesCentipedes
LegsTwo pairs on most body segments; attached to underside of bodyOne pair per body segment; attached to sides of body; last pair extends backwards
LocomotionGenerally adapted for burrowing or inhabiting small crevices; slow-movingGenerally adapted for running, except for the burrowing soil centipedes
FeedingPrimarily detritivores, some herbivores, few carnivores; no venomPrimarily carnivores with front legs modified into venomous fangs
SpiraclesOn underside of bodyOn the sides or top of body
Reproductive openingsThird body segmentLast body segment
Reproductive behaviourMale generally inserts spermatophore into female, using his gonopodsMale produces spermatophore that is usually picked up by female

Life cycle

Reproduction

Centipede reproduction does not involve copulation. Males deposit a spermatophore for the female to take up. In temperate areas, egg laying occurs in spring and summer. A few parthenogenetic species are known. Females provide parental care, both by curling their bodies around eggs and young, and by grooming them, probably to remove fungi and bacteria.
Centipedes are longer-lived than insects; the European Lithobius forficatus may live for 5 to 6 years, and the wide-ranging Scolopendra subspinipes can live for over 10 years. The combination of a small number of eggs laid, long gestation period, and long time of development to reproduction has led authors to label lithobiomorph centipedes as K-selected.

Development

Centipedes grow their legs at different points in their development. In the primitive condition, seen in the orders Lithobiomorpha, Scutigeromorpha, and Craterostigmomorpha, development is anamorphic: more segments and pairs of legs are grown between moults. For example, Scutigera coleoptrata, the house centipede, hatches with only four pairs of legs and in successive moults has 5, 7, 9, 11, 15, 15, 15 and 15 pairs respectively, before becoming a sexually mature adult. Life stages with fewer than 15 pairs of legs are called larval stadia. After the full complement of legs is achieved, the now postlarval stadia develop gonopods, sensory pores, more antennal segments, and more ocelli. All mature lithobiomorph centipedes have 15 leg-bearing segments. The Craterostigmomorpha only have one phase of anamorphosis, with embryos having 12 pairs, and adults 15.
The clade Epimorpha, consisting of the orders Geophilomorpha and Scolopendromorpha, is epimorphic, meaning that all pairs of legs are developed in the embryonic stages, and offspring do not develop more legs between moults. This clade contains the longest centipedes. In the Geophilomorpha, the number of thoracic segments usually varies within species, often on a geographical basis, and in most cases, females bear more legs than males. The number of leg-bearing segments varies within each order, but the total number of pairs is always odd, so there are never exactly 100 legs or 100 pairs, despite the group's common name.
Centipede segments are developed in two phases. Firstly, the head gives rise to a fixed but odd number of segments, driven by Hox genes as in all arthropods. Secondly, pairs of segments are added at the tail end by the creation of a prepattern unit, a double segment, which is then always divided into two. The repeated creation of these prepattern units is driven by an oscillator clock, implemented with the Notch signalling pathway. The segments are homologous with the legs of other arthropods such as trilobites; it would be sufficient for the Notch clock to run faster, as it does in snakes, to create more legs.

Ecology

Diet

Centipedes are predominantly generalist predators, which means they are adapted to eat a broad range of prey. Common prey items include lumbricid earthworms, dipteran fly larvae, collembolans, and other centipedes. They are carnivorous; study of gut contents suggests that plant material is an unimportant part of their diets, although they eat vegetable matter when starved during laboratory experiments.
Species of Scolopendromorpha, noticeably members from the genera Scolopendra and Ethmostigmus, are able to hunt for substantial prey items, including large invertebrates and sizable vertebrates, which could be larger than the myriapod itself. For instance, Scolopendra gigantea preys on tarantulas, scorpions, lizards, frogs, birds, mice, snakes, and even bats, catching them in midflight. Three species are amphibious, believed to hunt aquatic or amphibious invertebrates.