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18 Phylum Arthropoda:
The Myriapods (Centipedes,
Millipedes, and Their Kin)

fpo
It inhabits dwellings and appears at night. So rapidly
can it travel that one often just gets a glimpse of it as
it traverses a floor or wall. It is so fragile that no
damage can be done by it and much good is gained
through its capture of flies, cockroaches, clothes moths,
and other household pests.
Edward Essig,
speaking of the common house centipede, Scutigera

T he arthropod subphylum Myriapoda includes four groups, or classes:

Chilopoda (centipedes), Diplopoda (millipedes), Symphyla (symphy-
lans), and Pauropoda (pauropodans). All modern myriapods are terres-
trial, but their ancestry probably lies in the marine realm. The first fossil records
of millipedes are from the late Ordovician/early Silurian, and some of these are
thought to represent marine species. Fossil evidence suggests that myriapods
(millipedes) did not make their first appearance on land until the mid-Silurian.
Figure 18.1 illustrates a variety of myriapod types. About 11,460 living species
have so far been described.
Almost everyone is familiar with centipedes and millipedes, which, despite
their diversity, conform to a basic bauplan and external appearance. Myriapods
are quickly distinguished by a body divided into just two tagmata, the cephalon
and the long, homonomous, many-segmented trunk. As in the Hexapoda, there
are just four pairs of cephalic appendages: antennae, mandibles, first maxillae,
and second maxillae.
Millipedes are particular favorites with many people, and their harmless an-
tics have provided safe entertainment and lessons in biology for generations of
students of all ages. Millipedes are slow-moving detritivores that spend their
time burrowing through soil and litter, consuming plant remains and converting
vegetable matter into humus. In tropical environments, where earthworms are
often scarce, millipedes may be the major soil-forming animals. Their trunk seg-
ments are actually pairs of two fused segments (called diplosegments) and thus
bear two pairs of legs (see Figure 18.2F). It is thought that the diplosegment con-
dition evolved in conjunction with burrowing habits, as the pushing force of the
legs was more efficient when alternating body joints were made rigid and incom-
pressible. When threatened, many millipedes roll up into a flat coil, and some can
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(A) (B) (C)

(G)
(D)

(E)

Figure 18.1 Representative myri-

apods. (A) The New World centipede,
Nyssodesmus. (B) Scutigera coleoptrata, (F)
the common “house centipede.”
(C) The Southwestern desert centi-
pede Scolopendra heros. (D) A milli-
pede from East Africa. (E) The South-
western desert millipede Orthoporus
ornatus. (F) Pauropus silvaticus
(Pauropoda). (G) Scutigerella
(Symphyla).

roll up into perfect balls, like pillbugs (oniscid isopods). exposed to these defensive chemicals simply go to sleep).
Millipedes typically have many trunk segments, but One common household species in North America (intro-
some African species have only a few segments and are duced long ago from Asia), Oxidus gracilis, releases
often mistaken for pillbugs (especially when they roll strong-smelling defensive chemicals when injured. Mem-
up). Although popular myth says millipedes have a bers of the subclass Penicillata have evolved a porcupine-
thousand legs (milli–pedes), none do, although the record like defense, throwing stiff bristles from the posterior end
holder (Illacme plenipes, a California species), with 375 at ants and other predators. Given their range of defen-
pairs of legs, is still impressive. sive tactics, it is not surprising that many millipedes have
Even though millipedes are docile creatures, many of aposematic (warning) coloration, usually bright reds, yel-
their diplosegments bear lateral repugnatorial glands lows, and oranges, and that some soil-dwelling verte-
that secrete volatile toxic liquids used for defense (see brates (lizards and worm snakes) have evolved look-alike
Figure 18.2F). The defensive chemicals of this group are coloration (Batesian mimicry). Species of the California
surprisingly diverse, and they include quinones, phenols, genus Motyxia (Polydesmida) even utilize biolumines-
and even hydrogen cyanide produced by the common cence, possibly for warning away predators.
flat-backed millipedes of the order Polydesmida. A few In contrast to millipedes, most centipedes are fast-
tropical species have toxins powerful enough to raise running predators, with one pair of legs per segment,
blisters on the skin of humans. The European species and with poison claws (= fangs) on the first legs. They
Glomeris marginata produces quinazolinones, which be- never roll into a coil when threatened; instead, they usu-
long to the same class of substances as the synthetic drug ally strike out with the poison claws with which they
Quaalude, a powerful sedative (attacking wolf spiders can inflict a painful bite. The bite of the American giant
 PHYLUM ARTHROPODA: THE MYRIAPODS 3

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desert centipede, Scolopendra heros, which reaches 8 Synopses of Myriapod Taxa
inches in length, is known to cause local necrosis (simi-
lar to the sting of the bark scorpion, Centruroides). Scolo- Subphylum Myriapoda
pendra also has warning coloration—black and red One pair of antennae; mandibles with articulating en-
banding (Figure 18.1C). One remarkable species, the ap- dite; first maxillae free or fused; second maxillae partly
propriately named Henia vesuviana, secretes copious (or wholly) fused, or absent; postcephalic segments nu-
amounts of proteinaceous glue in response to attack merous, constituting an undifferentiated trunk; all ap-
from predators. The glue hardens within a few seconds pendages of head and trunk uniramous; living species
of exposure to air and immobilizes even the largest in- apparently without compound eyes; exoskeleton with-
sect predators. Some lithobiomorphs bear large num- out well-developed wax layer (except in some desert
bers of unicellular repugnatorial glands on the last four species); many with lateral repugnatorial glands on
pairs of legs, which they kick in the direction of an trunk segments; without entodermally derived diges-
enemy—a behavior that throws out sticky droplets of tive ceca; ectodermally derived (proctodeal) Malpighian
noxious secretions. tubules assist in excretion; with organs of Tömösvary;
copulation indirect; development direct.

Class Diplopoda
Myriapod Classification
Trunk segments fused into pairs called diplosegments;
SUBPHYLUM MYRIAPODA most diplosegments with two pairs of legs, spiracles,
ganglia, and heart ostia; each diplosegment with one
CLASS DIPLOPODA
tergite, two pleura, and one to three sternites; anterior-
SUBCLASS PENICILLATA most segments often with legs suppressed except for
ORDER POLYXENIDA some internal musculature; first trunk segment legless,
modified as collum; antennae simple, 7-jointed; first
SUBCLASS CHILOGNATHA
maxillae fused into a gnathochilarium; second maxillae
ORDER CALLIPODIDA absent; gonopores open anteriorly, on or near coxae of
ORDER CHORDEUMATIDA second pair of legs (third trunk segment); cuticle usual-
ly calcified; many capable of rolling into a tight coil;
ORDER GLOMERIDA
spiracles located ventrally, typically in front of leg
ORDER GLOMERIDESMIDA coxae, and never valvular (cannot be closed); gono-
ORDER JULIDA pores open in anterior region of body; with 11 to 192
leg-bearing body diplosegments (the first and last
ORDER PLATYDESMIDA
diplosegments are always legless, and diplosegments
ORDER POLYDESMIDA 2–4 have one pair of legs each); legs ventrally posi-
ORDER POLYZONIIDA tioned and short, such that the body is carried close to
the ground. Although millipedes lack claws or poisons,
ORDER SIPHONOPHORIDA
many have lateral repugnatorial glands on the trunk
ORDER SIPHONIULIDA segments that secrete noxious chemicals that can irri-
ORDER SPHAEROTHERIIDA tate the skin and eyes. About 8,000 species of millipedes
have been described.
ORDER SPIROBOLIDA
Millipede classification is unstable, with many fami-
ORDER SPIROSTREPTIDA lies of uncertain validity and relationship. The subclass
ORDER STEMMIULIDA Penicillata contains soft-bodied millipedes, without cal-
cification in the exoskeleton, which is covered by tufts of
CLASS CHILOPODA
bristles; males lack copulatory appendages, and repro-
SUBCLASS NOTOSTIGMOPHORA duction occurs without contact between the sexes. The
ORDER SCUTIGEROMORPHA subclass Chilognatha contains millipedes with a hard,
calcified exoskeleton, only scattered setae, and male re-
SUBCLASS PLEUROSTIGMOPHORA
productive appendages; reproduction requires contact
ORDER CRATEROSTIGMOMORPHA between the sexes. Glomeridesmids are flattened milli-
ORDER GEOPHILOMORPHA pedes with 22 diplosegments that cannot roll into a ball.
Glomerids and sphaerotheriids are short, round, 12- or
ORDER LITHOBIOMORPHA
13-segmented millipedes that can roll into a perfect ball.
ORDER SCOLOPENDROMORPHA The most colorful millipedes are the polydesmids,
CLASS PAUROPODA whose bright red, orange, and blue pigmentations warn
of their cyanide defensive secretions.
CLASS SYMPHYLA
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Class Pauropoda Approximately 2,800 living species of chilopods have
Nearly microscopic (0.5–1.5 mm); eyeless; 9–11 pairs of been described.
legs; some trunk segments partly fused (but not as true
diplosegments), plus free telson; without limbs on first Class Symphyla
trunk segment; mouthparts poorly developed; first maxil- Small (0.5–8.0 mm); eyeless; trunk with 14 segments, last
lae fused into a gnathochilarium; second maxillae absent; fused to telson; first 12 trunk segments each with a pair
antennae branched; most without tracheal or circulatory of legs; penultimate segment with spinnerets and a pair
systems; gonopores on third trunk segment; terga often of long sensory hairs; dorsal surface with 15–22 tergal
large and extended over two segments; cuticle soft, uncal- plates; soft uncalcified cuticle; antennae long, simple,
cified. Although found in all parts of the world, pauropo- threadlike; first maxillae medially coalesced; second
dans occur mainly in moist soils and woodland litter; they maxillae completely fused as complex labium; one pair
are not common. About 500 species have been described. of spiracles (on head); tracheae supply first 3 trunk seg-
ments; gonopores open on third trunk segment. Sym-
Class Chilopoda phylans are generally uncommon arthropods that occur
With numerous unfused trunk segments, each with one in soil and rotting vegetation. About 160 species have
pair of legs, the first pair modified as large poison claws been described in two families.
(called prehensors, or forcipules) and held under the
head like mouthparts (also sometimes referred to as
“maxillipeds”); antennae simple, of varying segmenta- The Myriapod Bauplan
tion; both pairs of maxillae may be medially coalesced;
cuticle stiff but uncalcified; gonopores on last true body In Chapter 15 we discussed the various advantages and
segment; spiracles lateral or middorsal (but on the constraints imposed by arthropodization, and in Chap-
pleural side walls), and in some species valvular (can be ter 17 we discussed the suitability of many features of
closed); with 15 to 193 leg-bearing body segments; al- the arthropod bauplan for adaptation to a terrestrial
though the legs are long, they extend laterally such that lifestyle. The myriapods share many of these adapta-
the body is carried close to the ground; last pair of legs tions with the hexapods, although perhaps as a result of
extend backwards and not used for locomotion. Cen- convergent of parallel evolution.
tipedes may be the only animals with legs modified into The myriapods differ from the hexapods in their reten-
poison-injecting claws. tion of a more homonomous condition: the long trunk
The subclass Notostigomorpha have middorsal spir- with its paired segmental appendages. However, the head
acles, hemocyanin in the hemolymph, a dome-shaped appendages and legs are very similar to those of the in-
head, pseudo-faceted eyes (ocelli), and 15 pairs of long, sects, and it was for this reason that the two groups were
thin legs. The subclass Pleurostigmophora have lateral long considered sister taxa. Metamerism is strong in myr-
spiracles, a flattened head, a variable number of legs, iapods and is evident internally in structures such as the
and no respiratory pigments. The largest centipedes are segmental heart ostia, tracheae, and ganglia. The key fea-
the scolopendromorphans (with either 21 or 23 pairs of tures that distinguish the Myriapoda are listed in Box 18A.
legs), although the highest leg count occurs in the long, The centipedes (chilopods) bear one pair of walking
thin geophilomorphs (with 27 or more pairs of legs). legs per segment, and there may be up to 193 trunk seg-

BOX 18A Characteristics of the Subphylum Myriapoda

1. Body of two tagmata: head and multisegmented 4. Without a carapace
trunk
5. With an aerial gas exchange system composed of
2. All appendages multiarticulate and uniramousa tracheae and spiracles (probably convergent with
those in the Hexapoda)
3. Head appendages, from anterior to posterior, are
antennae, mandibles, first maxillae (maxillules), 6. With one or two pairs of ectodermally derived
and second maxillae; second maxillae may be (proctodeal) Malpighian tubules (probably conver-
fused into a single flaplike structure called a “labi- gent with those in the Hexapoda)
um” (not homologous to the crustacean labium),
7. Most with simple ocelli, at least in some stage of
or they may be absent; first and second maxillae
often bear palps. the life cycle; true ommatidia apparently absent
8. Gut simple, without digestive ceca
aPauropodan antennae are branched, but whether this repre-
sents a vestige of a primitive biramous condition (as in crus- 9. Dioecious; with direct development
taceans) or is secondarily derived is not known.
 UNCORRECTED PAGE PROOFS (C)

(A) (B)

(D)

(F)
Tergum

Opening of
(E) repugnitorial
gland
(H)
Spiracle
Coxa

Walking leg

(G)

(I)

Figure 18.2 General body anatomy of myriapods. (A) The two

body regions of myriapods; shown is a lithobiomorph centipede. (B,C)
A female scolopendromorph centipede (dorsal and ventral views),
with middle body segments omitted. (D) Head of a scutigeromorph
centipede. (E) Anterior end of a millipede (Diplopoda). (F) A single
diplosegment from the millipede Gigantowales chisholmi (Diplopoda);
only one leg is shown. (G,H) Trunk segments in two centipedes
shown as transverse sections; note the variation in limb morphology:
(G) Lithobius. (H) Scutigera. (I) Ventral view of the terminal segments
of a male geophilomorph centipede (Strigamia).
6 CHAPTER EIGHTEEN

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ments (Figure 18.2A–D). Most species are 1–2 cm in pauropodans the first maxillae fuse to form a flaplike
length, although some tropical giants attain lengths of gnathochilarium. The ganglia of the mandibular and
nearly 30 cm. maxillary segments are generally fused to form the sube-
The millipedes (diplopods) have an interesting modi- sophageal ganglion. A carapace is never present in myri-
fication of basic metamerism. All of the trunk segments apods, and there is no evidence that one ever existed.
are actually diplosegments, each formed by fusion of Like the Crustacea and Hexapoda, the Myriapoda
two somites and bearing a double complement of meta- possess two additional head structures: a clypeus–
meric organs and structure (Figure 18.2F) The head is fol- labrum that borders the anterior mouth field, and a hy-
lowed by an expanded, limbless diplosegment called the popharynx that borders the posterior mouth field. The
collum, which forms a conspicuous, heavily sclerotized clypeus projects off the front of the head and extends
collar between head and trunk (Figure 18.2E). Each of down over the mandibles, much as our upper lip covers
the next three diplosegments bears a single pair of legs. our teeth. The platelike labrum projects off the clypeus.
All the remaining diplosegments retain two pairs of legs. Some workers consider the labrum to be the fused ap-
When the alternate body somites become fused during pendages of the acron or premandibular head segment,
embryogenesis, the leg pair, spiracle pair, and ventral but most evidence suggests it has been independently
nerve cord ganglion of the anterior fused somite shifted derived from the exoskeletal sclerites. However, prelim-
to the posterior fused somite. Thus, in the resulting inary research suggests that the developmental gene
diplosegment, the metazonite (representing the posteri- Distal-less is expressed in the ontogeny of the labrum,
or end of the two fused segments) carries all these struc- leaving the matter rather unsettled. Arising behind the
ture, whereas the prozonite (the anterior somite) lacks first maxillae and near the base of the second maxillae is
them and is free to insert inside the preceding diploseg- the flaplike (or tonguelike) hypopharynx. The salivary
ment in a telescoping fashion (Figure 18.2F). Like cen- glands open through the hypopharynx. The hypophar-
tipedes, millipedes range in length from about 1 to 30 ynx probably does not represent true appendages, but is
cm. The cuticle of millipedes is particularly robust, being an independent outgrowth of the body wall. However,
well sclerotized and usually calcified. the homology of both the clypeus and the hypopharynx
Pauropodans are minute, soft-bodied, eyeless, soil- are matters of ongoing debate.
inhabiting myriapods that resemble millipedes (Figure
18.1F). They are less than 2 mm in length and have up to Locomotion
11 trunk segments. As in millipedes, some segmental Myriapods rely on their well-sclerotized exoskeleton for
pairing occurs, and usually only six tergites are visible body support. In centipedes, the legs are long but ex-
dorsally. tend laterally, keeping the body close to the ground to
Symphylans (Figure 18.1G) are also minute, eyeless maintain stability while at the same time providing for
myriapods, never exceeding 1 cm in length. The trunk long strides and rapid locomotion. In millipedes, the
has 12 pairs of leg-bearing segments. Some tergites are legs arise ventrally and are short, again keeping the
divided, and 15–22 tergites are usually visible dorsally. body close to the ground while providing for powerful,
The thirteenth body segment bears spinnerets, a pair of though slow, locomotion.
long sensory hairs, and a tiny postsegmental telson. Like The basic design of arthropod limbs was described in
pauropodans, symphylans inhabit loose soil and humus. Chapter 15. Recall that in most arachnids the coxae are
immovably fixed to the body, and limb movement oc-
Head and Mouth Appendages curs at more distal joints. However, in myriapods (as in
As you will recall from previous chapters, much discus- most hexapods and crustaceans), anterior–posterior
sion and evolutionary speculation regarding arthropods limb movements take place between the coxae and the
focuses on the nature of the head and its appendages. body proper. As we have learned, the power exerted by
The basic five-segmented ( + acron) crustacean-hexapo- a limb is greatest at low speeds and least at high speeds.
dan head is retained in the Myriapoda (Figure 18.2). The At lower speeds the legs are in contact with the ground
acron bears the eyes and contains the protocerebrum. for longer periods of time, and in the case of myriapods
The first postacronal cephalic somite (the antennary seg- more legs are contacting the ground at any given mo-
ment) bears the antennae and houses the deutocere- ment. Thus, in burrowing forms, such as most milli-
brum. The second segment is the premandibular seg- pedes, the legs are short, and the gait is slow and pow-
ment; it lacks appendages but houses the tritocerebrum. erful as the animals bulldoze their way through soil or
The third segment (mandibular segment) carries the rotting wood. In centipedes, which run at high speeds,
mandibles, and the fourth (first maxillary segment) less than half the limbs may touch the ground at any
bears the first pair of maxillae (= maxillules). The fifth given moment, and for shorter periods of time (Figure
cephalic segment (second maxillary segment) bears the 18.3). Longer limbs increase the speed of a running gait,
second pair of maxillae. The second maxillae are fused as and limbs long in length and stride are typical features
a “labium” in symphylans, but they have been lost alto- of the fastest-running centipedes and symphylans.
gether in millipedes and pauropodans. In diplopods and Long, slender, fast moving objects, such as centipedes
 PHYLUM ARTHROPODA: THE MYRIAPODS 7

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(A) (E) (F)

(B)

(C)

Figure 18.3 Locomotion in millipedes and cen-

tipedes. (A,B) A typical millipede (such as Spiro-
streptus or Gymnostreptus) in motion. Note the 16
diplosegments (with 32 pairs of legs). The dorsal view
(A) shows left and right leg sets exactly in phase with each
(D) other. The lateral view (B) of the same animal shows that
the majority of the limb tips are on the ground at once, an
arrangement yielding a slow but powerful gait. (C,D) A
typical centipede (such as Scolopendra or Cryptops) in
motion. Note the 12 segments, each with one pair of legs.
The dorsal view (C) shows the limb pairs in opposite phase
(and trains!) tend to develop side-to-side oscillations.
and the undulations of the body that accentuate stride
Although this may increase stride length in a centipede, length. The lateral view (D) of the same animal illustrates
it also counters the forward motion and can slow the an- that fewer than one-third of the limb tips are on the
imal down. Consequently, centipedes have developed ground at any one time, an arrangement yielding the
anatomical modifications to dampen this lateral body short, swift strokes typical of a rapid but weak gait. Arrows
sway, such as tergal fusion at oscillation nodal points indicate the animal’s direction of travel; dots are points of
(Scutigeropmorpha) and shortening body segments leg-tip contact with the substratum. (E) Locomotion in a
scolopendrid centipede at various speeds. 1–4 show the
(and hence, the body) while retaining the same number
body waves and leg actions at increasing speeds. Limbs
of legs (Lithobiomorpha). Among the pauropodans, a shown with heavy lines are in their power strokes, with the
range of movement speeds is seen, from species that are tips against the substratum (dots); limbs depicted by thin
“slow plowers” to those that are fast runners. lines are in various stages of their recovery strokes. Notice
Locomotor repertoires typically evolve in concert with that at maximum speed the animal is still supported by a
the overall habits of animals, particularly feeding behav- tripod stance. (F) Field of leg movements in a running cen-
iors. Most centipedes are surface-dwelling predators that tipede, Scutigera. The heavy vertical lines trace the move-
ment of the tips of each leg during the propulsive back-
must move quickly to capture their prey. Scutigera is a
stroke. Note the gradual increase in limb length posteriorly
small centipede that qualifies as a world-class runner, that allows for unencumbered overstepping even when
reaching speeds up to 42 cm/second when in pursuit of the full swing of the limb is used.
its favorite prey, flies. On the other hand, most millipedes
are detritivores that burrow through soil, leaf litter, or rot-
ten timber in search of food. Thus millipedes tend to have
shorter legs and slower, but more powerful gaits. Also, when millipedes and pauropodans burrow, they
Power may also be increased by increasing the num- tuck the head ventrally so that the collum is thrust for-
ber of legs, and thus the number of body segments. In ward, in some cases like a wedge that splits open the
this regard, physical limits are reached when the body soil or humus, and in other cases like the broad blade of
achieves a length-to-width ratio that would result in a bulldozer (Figure 18.2E). Some millipedes have under-
buckling of the trunk. This constraint is partly overcome gone a habitat reversal to abandon burrowing and seek
in millipedes and pauropodans in two principal ways: out cracks and crevices.
by increasing the cross-sectional body diameter (becom- One group of centipedes lacks the high-speed modi-
ing “fatter”) and by uniting segments into pairs. In fications of most chilopods. The geophilomorphs bur-
some species the body wall is further strengthened by row, aided by dilation of the body and use of the trunk
having certain trunk segments fused into a solid ring. musculature in a fashion similar to that of earthworms.
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This peristalsis-like motion is rare in arthropods because tipedes do, but these are exceptions (one North Ameri-
of their rigid exoskeleton. However, geophilomorphs can species apparently specializes on insect pupae).
have enlarged areas of flexible cuticle on the sides (pleu- Millipedes are often the largest animals living in cave
ra) of the body between the tergites and sternites; these ecosystems and, although troglobitic species are typical-
enlarged pleura allow them to significantly alter their ly terrestrial, at least one species lives much of its time
body diameter. Other centipedes have smaller, flexible submerged in cave streams, feeding on aquatic bacteria.
pleural areas that allow some degree of lateral undula- The feeding biology of pauropodans is not well un-
tory motion. Centipede legs are attached to these flexi- derstood, but most appear to be scavengers. These
ble regions, an arrangement that increases the range of minute, blind creatures crawl through soil and humus,
limb motion in these fast-running surface dwellers. This feeding on fungi and decaying plant and animal matter.
condition is in marked contrast to that of millipedes, in Symphylans are primarily herbivores, although a few
which the short legs arise from the ventral sternites, lim- have adopted a carnivorous, scavenging lifestyle. Many
iting their range of movement. symphylans consume live plants. One species, Scuti-
One of the principal problems associated with in- gerella immaculata, is a serious pest in plant nurseries and
creased limb length is that the field of movement of one flower gardens, where it has been reported at densities
limb may overlap that of adjacent limbs. Potential leg greater than 90 million per acre.
interference is prevented by having limbs of different Like the guts of all arthropods, the long, usually
lengths, so that the tips of adjacent legs move at differ- straight myriapod gut is divisible into a stomodeal
ent distances from the body (Figure 18.3F). In fast-run- foregut, entodermal midgut, and proctodeal hindgut
ning centipedes the limbs of each succeeding segment (Figure 18.4). There are no digestive ceca branching off
are slightly longer than the immediately anterior pair. the gut. Salivary glands (= mandibular glands) are as-
The legs of most myriapods move in clear metachronal sociated with one or several of the mouth appendages.
waves from posterior to anterior (Figure 18.3A,B). Un- The salivary secretions soften and lubricate solid food,
like most arthropods, millipedes move the two pairs of and in some species contain enzymes that initiate chem-
legs on each diplosegment synchronously (stability is ical digestion. The mouth leads inward to a long esoph-
not a problem in these elongate creatures). agus, which is sometimes expanded posteriorly as a
storage area, or crop and gizzard (as in most centi-
Feeding and Digestion pedes). The gizzard often contains cuticular spines that
Most centipedes are active, aggressive predators on help strain large particles from the food entering the
smaller invertebrates, particularly worms, snails, and midgut, where absorption occurs (Figure 18.4C). As in
other arthropods. Their first trunk appendages form other arthropods, the gut of myriapods produces a per-
large claws, called prehensors or forcipules. These rap- itrophic membrane—a sheet of thin, porous, chitinous
torial limbs are located ventral to the mouth field (Figure material that lines and protects the midgut, and may
18.2D), and are used to stab prey and inject poison. The pull free to envelop and coat food particles as they pass
poison is produced in large venom glands located in the through the gut. The midgut connects to a short, proc-
basal articles of the forcipules. The poison is so effective todeal hindgut that terminates at the anus.
that large centipedes, such as the tropical Scolopendra,
can subdue small vertebrates (e.g., frogs, lizards, snakes, Circulation and Gas Exchange
mice, small birds). Some centipedes actually rear up on The myriapod circulatory system includes a dorsal
their hindlegs and capture flying insects! The prehensors tubular heart that pumps the hemocoelic fluid (blood)
and second maxillae hold the prey, while the mandibles toward the head. The heart narrows anteriorly to a ves-
and first maxillae bite and chew. The bite of even the sel-like aorta, from which blood flows posteriorly
most dangerous centipede is normally not fatal to peo- through large hemocoelic chambers before returning to
ple, but the poison can cause a reaction similar to that ac- the pericardial sinus and then back to the heart via
companying a serious wasp or scorpion sting. paired lateral ostia. Circulation is slow, and system pres-
The feeding strategy of millipedes is quite different. sure is relatively low. In diplopods, the heart bears two
Most millipedes are slow-moving detritivores with a pairs of ostia in each diplosegment; one pair of ostia oc-
preference for dead and decaying plant material. They curs in each segment in the chilopods.
play an important role in the recycling of leaf litter in Myriapods rely on a tracheal system for gas ex-
many parts of the world. Most bite off large pieces of change (Figure 18.5). As explained in Chapter 15, tra-
vegetation with their powerful mandibles, mix it with cheae are extensive tubular invaginations of the body
saliva as they chew it, and then swallow it. Some, such as wall opening through the cuticle by pores called spira-
the tropical siphonophorids, are believed to feed on the cles. The tracheae originating at one spiracle commonly
juices of living plants and fungi. In these groups the anastomose with others to form branching networks
labrum, gnathochilarium, and reduced mandibles are penetrating most of the body. Spiracles are placed seg-
modified into a suctorial piercing beak. A few odd mentally, although not necessarily on all segments. In
groups of millipedes are predaceous and feed as cen- chilopods the spiracles are located in the membranous
 PHYLUM ARTHROPODA: THE MYRIAPODS 9

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(A) (B) (C)

Figure 18.4 The gut in myriapods. (A) Internal

anatomy of a typical centipede (Lithobius). (B) Gut of
the centipede Cryptops. (C) Gizzard region of Cryptops
(longitudinal section).

pleural (side) region just above and behind the base of Each spiracle is usually recessed in an atrium, whose
each leg, or most legs (although in Scutigeromorpha the walls are lined with setae or spines (trichomes) that pre-
spiracles are dorsal). Diplopods usually bear two pairs vent dust, debris, and parasites from entering the tra-
of spiracles per diplosegment, just anterior to the leg cheal tubes. In myriapods the spiracles are often sur-
coxae, where they open from the sternum and are asso- rounded by a sclerotized rim or lip, the peritrema,
ciated with the apodemes, which also serve as insertions which also aids in excluding foreign particles. A muscu-
for extrinsic leg muscles. In the symphylans a single lar valve or other closing device is present in many cen-
pair of spiracles opens on the sides of the head, and the tipedes and under control of internal partial pressures of
tracheae supply only the first three trunk segments. Ex- O2 and CO2. As in insects, ventilation of the tracheal
cept for a few primitive species, most pauropodans lack system is accomplished by simple diffusion gradients,
a tracheal system. as well as by pressure changes induced by the animal’s
movements. The blood of myriapods appears to play no
(A) significant role in oxygen transport, except perhaps in
the very active scutigeromorphs. Instead, the air-filled
tracheae extend directly to each organ, where the ends
lie close to or within the tissues.
The innermost parts of the tracheal
(B) system are the tracheoles, which are thin-
walled, fluid-filled channels that end as a
single cell, the tracheole end cell (= tra-
cheolar cell). Unlike tracheae, tracheoles
are not shed during ecdysis. The trache-
oles are so minute (0.2–1.0 mm) that ven-

Figure 18.5 The tracheal system of cen-

tipedes and millipedes. (A) Tracheal system
of three body segments of a scolopendro-
morph centipede, Scolopendra cingulata.
(B) A spiracle of S. cingulata (transverse
section).
10 CHAPTER EIGHTEEN

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tilation is impossible and gas transport here relies on
aqueous diffusion.
A few geophilomorph centipedes live in intertidal
habitats, under rocks or in algal mats. Presumably, air
retained within the tracheal system (and in the spiracu-
lar atrium) is sufficient to last during submergence at
high tides.
Although the tracheal system of myriapods greatly
resembles that of insects, there is evidence that it
evolved independently. In addition, there is some indi-
cation that terrestrial myriapods could have evolved
from marine ancestors, not from a terrestrial line that
was tied to the Hexapoda.

Excretion and Osmoregulation

When myriapods invaded the land, the problem of
water conservation necessitated the evolution of entire-
ly new structures to remove metabolic wastes—
Malpighian tubules (Figure 18.4). These excretory or-
gans function much as those of insects do (see Chapter
17). Despite their length, however, centipedes and milli-
pedes usually possess only one and two pairs of
Figure 18.6 The brain and anterior ganglia of a cen-
Malpighian tubules, respectively. In those species that tipede, Lithobius forficatus (dorsal view).
are largely confined to moist habitats and nocturnal ac-
tivity patterns, a significant portion of the excretory
wastes may be ammonia rather than uric acid.
The Malpighian tubules of myriapods were long as- head segments and controls the mouthparts, salivary
sumed to be homologous to those of hexapods. Howev- glands, and some other local musculature.
er, as we saw in Chapter 15, there is now growing evi- Myriapods typically possess eyes at some stage in
dence that these two groups do not share an immediate their life cycle, but biologists are still debating whether
common ancestry. If this is true, then these excretory true compound eyes occur in this group. Centipedes
structures represent another striking case of convergent possess a few to many eyes that appear to be simple
evolution within the Arthropoda. ocelli (Figure 18.7). In the scutigeromorph centipedes, up
The myriapod cuticle is sclerotized and calcified to to 200 of these eyes may cluster to form a sort of pseudo-
various degrees, adding a measure of waterproofing, compound eye. However, the eyes of living centipedes
but aside from a few desert species, it lacks the waxy appear to function only in the detection of light and
layer seen in hexapods. For this reason, myriapods rely dark, and not in image formation. Many parasitic and
to a considerable extent on behavioral strategies to
avoid desiccation. Many live in humid or wet environ-
ments, or are active only during cool periods. Others
stay hidden in cool or moist microhabitats, such as
under rocks, during hot hours or dry periods.

Nervous System and Sense Organs

The myriapod nervous system conforms to the basic
arthropod plan described in Chapter 15. Very little sec-
ondary fusion of ganglia occurs, and the ventral nerve
cord retains much of its primitive double nature, with a
pair of fused ganglia in each segment. Millipedes pos-
sess two pairs of fused ganglia in each diplosegment
(Figure 18.6).
Like the “brains” of other arthropods, the cerebral
ganglion of myriapods comprises three distinct regions:
the protocerebrum (associated with the eyes), the deuto-
cerebrum (associated with the antennae), and the trito-
cerebrum. The subesophageal ganglion is composed of
fused ganglia of the third, fourth, and perhaps the fifth Figure 18.7 The ocellus of a centipede (section).
 PHYLUM ARTHROPODA: THE MYRIAPODS 11

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troglobitic myriapods have lost their eyes completely
(eyes are absent in the centipede order Geophilomorpha,
and the millipede orders Siphoniulida, Siphonophorida,
and Polydesmida). Symphylans and pauropodans also
lack eyes. Many burrowing diplopods are also eyeless,
whereas others have from 2 to 80 ocelli arranged vari-
ously on the head. Some diplopods also possess integu-
mental photoreceptors, and many eyeless species exhib-
it negative phototaxis.
Centipedes and millipedes are noted for their highly
sensitive antennae, which are richly supplied with tac-
tile and chemosensory setae. In many species, an organ
of Tömösvary is located at the base of each antenna
(Figure 18.8). Each part of this paired organ consists of a
disc with a central pore where the ends of sensory neu-
rons converge. The exact role of this organ has yet to be
clearly established, and speculation has run the gamut Figure 18.8 The organ of Tömösvary from the cen-
from chemosensation to pressure sensation to humidity tipede Lithobius forficatus.
detection to audition (sound or vibration detection). The
last idea is probably the most popular today. Whether
the organ of Tömösvary might detect aerial vibrations
(auditory impulses) or only ground vibrations is also a direct development, with the young hatching as “minia-
debated question. ture adults,” although often with fewer body segments.
Beyond these generalizations, some specifics for each
Reproduction and Development major group are discussed below.
Myriapods are dioecious and oviporous, although par-
thenogenesis occurs in several families of millipedes Chilopoda. Female centipedes possess a single elon-
and in some centipedes and symphylans. Many myri- gate ovary located above the gut, whereas males have
apods, like most arachnids, rely on indirect copulation from 1 to 26 testes similarly placed (Figure 18.9). The
and insemination. Packets of sperm (spermatophores) oviduct joins with the openings from several accesso-
are deposited in the environment, or are held by the ry glands and a pair of seminal receptacles just inter-
male and picked up by the female. All myriapods have nal to the gonopore, which is situated on the genital

(A) (B) Figure 18.9 Reproductive sys-

tems of the centipede Lithobius
forficatus. (A) Female system.
(B) Male system.
12 CHAPTER EIGHTEEN

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segment (the legless segment in front of the telson, millipede gonads lie between the gut and the ventral
or pygidium). The gonopore of females is usually nerve cord. The gonopores open on the third (genital)
flanked by a pair of small grasping appendages, or trunk segment. In females each oviduct opens sepa-
gonopods. In males the testes join the ducts of sever- rately into a genital atrium (= vulva) near the limb
al accessory glands and a pair of seminal vesicles coxae. A groove in the vulva leads into one or more
near the gonopore, which opens on the ventral sur- seminal receptacles. Males possess a pair of penes, or
face of the genital segment. The male gonopore also gonapophyses, on the third trunk segment (on the
lies between a pair of small gonopods. coxae of the second pair of legs). The mandibles are
Sperm are packed into spermatophores, which are used to transfer sperm in the “pill millipedes,” but
transferred to the female. In some cases (e.g., scutigero- more commonly one or both pairs of legs of the sev-
morphs), males deposit spermatophores directly on the enth or eighth trunk segment are modified as copula-
ground and the females simply pick them up. In most tory appendages, or gonopods, and serve this pur-
species, however, females produce a silken nuptial net, pose. Males bend the anterior body segments back
which is spun from modified genital glands, and males under the trunk until the penes and gonopods make
deposit a spermatophore on this net (Figure 18.10). Mat- contact, whereupon the later picks up a spermato-
ing pairs of centipedes typically perform courtship be- phore in preparation for mating.
havior, stroking each other with their antennae and In most millipedes, mating occurs by indirect insem-
often moving the large (up to several millimeters) sper- ination, in which the male and female genital openings
matophore about the nuptial net. Eventually the female never actually make contact, although the male and fe-
picks up the spermatophore with her gonopods and in- male may embrace with their legs and mandibles and
serts it into her gonopore. Fertilization occurs as the may coil their trunks together. Sometimes a pair re-
eggs pass through the gonoduct. Females often coat the mains in such an embrace for up to two days. Antennal
fertilized eggs with moisture and fungicides before de- tapping, head drumming, and stridulation may be in-
positing them in the ground or in rotting vegetation. cluded in the mating repertoire, and pheromones are
Some species hatch as juveniles with all body segments known to play an important role in many species.
(i.e., epimorphic development), whereas others add Males often have an array of secondary sexual char-
new segments with the posthatching molts until the acteristics, including strong modifications of the legs
adult number is attained (i.e., anamorphic develop- and head and special glandular structures that are not
ment). Parental guarding typically occurs in the eipmor- well understood. In species of the genus Chordeuma (in
phic orders (Figure 18.10). Europe) these glands produce a secretion on the dorsal
surface upon which females feed prior to mating. Males
Diplopoda. Millipedes possess a single pair of elon- of the family Sphaeriotheriidae produce sounds during
gate gonads in both sexes. Unlike those of chilopods, courtship by rubbing the back edge of the body against

(A) Figure 18.10 (A–C) Reproductive behavior in the tropi-

(B) cal centipede Scolopendra cingulata. (A) A male and female
in mating position over the nuptial net. (B) A nuptial net
with a spermatophore. (C) A female and her eggs. (D) The
giant North American desert centipede Scolopendra heros
embracing her young.

(D)

(C)
 PHYLUM ARTHROPODA: THE MYRIAPODS 13

UNCORRECTED PAGE PROOFS

an enlarged rear leg. In most millipedes the gonopods trients (yolk) in the eggs. Like those of hexapods, the
no longer resemble walking legs and may be truly yolky eggs of myriapods show almost no trace of the
baroque in their complexity, with flagella, brushes, ancestral arthropod spiral cleavage pattern, and have
knobs, and hooks. shifted almost entirely to meroblastic cleavage. The
Fertilization takes place as the eggs are laid. In glom- early embryonic development of myriapods proceeds
erid millipedes, gonopods are not present, but the last much like that of hexapods, described in Chapter 17.
pair of male legs is modified into strong claspers, used to Most myriapods undergo early cleavage by in-
hold the female in place. In these millipedes, males mold tralecithal nuclear divisions, followed by a migration of
soil into a small cup into which they ejaculate sperm. the daughter nuclei to the peripheral cytoplasm (=
The sperm-filled cup is then passed to the female, who is periplasm). Here they continue to divide until the
firmly held by the male’s large claspers. The myriapod periplasm is dense with nuclei, whereupon cell mem-
specialist W. A. Shear notes that this may be the only branes begin to form, partitioning uninucleate cells from
known case of animals using a manufactured tool to one another. At this point the embryo is a periblastula,
mate! Eggs are usually laid in the soil, although some comprising a yolky sphere that has a few scattered nuclei
species also produce silk bags to house the eggs. Some and is covered by a thin cellular layer, or blastoderm.
species fashion a nest of soil and humus reinforced with Along one side of the blastula a patch of columnar
parental feces, or the entire nest may be composed of cells forms a germinal disc, sharply marked off from the
fecal material. One to 300 eggs are laid at a time. Hatch- thin cuboidal cells of the remaining blastoderm. From
ling millipedes typically possess seven or fewer body specific regions of this disc, presumptive endodermal
segments and only three pairs of legs. Additional seg- and mesodermal cells begin to proliferate as germinal
ments and legs are added at the rear of the last somite centers. These cells migrate inward during gastrulation
with succeeding molts (i.e., anamorphic development). to lie beneath their parental cells, which now form the
Unlike centipedes, most species of millipedes (excepting ectoderm. The mesoderm proliferates inward as a longi-
the order Platydesmida) show no parental care. tudinal gastral groove; the cells of the developing gut
usually surround and gradually begin absorbing the
Pauropoda. In pauropodans a single ovary lies large central yolky mass of the embryo, and paired
beneath the gut, but the testes are located above the coelomic spaces appear in the mesoderm.
gut. As in millipedes, the gonopores are on the third As segments begin to demarcate and proliferate, each
(genital) trunk segment. Females take up a sper- receives one pair of mesodermal pouches and eventual-
matophore, often after the male has suspended it from ly develops appendage buds. As the mesoderm con-
one or a few silk threads spanning two stones or leaves. tributes to various organs and tissues, the coelomic
Fertilization is internal. The yolky eggs are laid in spaces merge with the small blastocoel to produce the
decaying wood. Hatchlings lack many adult segments hemocoelic space. The mouth and anus arise by in-
and usually have only three pairs of legs (anamorphic growths of the ectoderm that form the fore- and hind-
development). guts, which eventually establish contact with the devel-
oping endodermal midgut.
Symphyla. Symphylans display one of the more Variations on this basic scheme occur in groups with
unusual methods of fertilization in the animal king- secondary yolk reduction, such as in pauropodans and
dom. Paired gonads discharge through gonopores on symphylans. In these groups a largely holoblastic cleav-
the third trunk segment in both sexes. Males deposit age takes place, and a coeloblastula actually forms. In
spermatophores in the environment. At least some most chilopods, a type of cleavage occurs that is some-
symphylans construct stalklike structures, topped what intermediate between total and superficial cleav-
with spermatophores. When a female encounters one age. After a few intralecithal nuclear divisions, the yolk
of these sperm packages, she bites off the spermato- breaks up into blocks called yolk pyramids. Gradually
phores and stores the sperm in her preoral cavity. the yolk pyramids disappear, and development shifts to
When her own eggs are ripe, the female removes the superficial type.
them from her gonopore with her mouthparts and
cements them to moss or some other substratum. Myriapod Phylogeny
Fertilization occurs during this process as she coats In Chapter 15 we discussed how the long-favored view
each egg with the stored sperm. Young symphylans of arthropod relationships is now being challenged by
hatch with only about half the adult number of trunk new evidence. Until recently, the prevailing hypothesis
segments and appendages (anamorphic develop- viewed the Myriapoda and Hexapoda as sister groups
ment). (together called the Atelocerata), and those groups as
sister to the Crustacea (together forming a grouping
Myriapod Embryogeny known as the Mandibulata). The sister-group relation-
The indirect copulation and external development typi- ship between myriapods and hexapods was based upon
cal of myriapods demands large amounts of stored nu- several seemingly strong synapomorphies, including
14 CHAPTER EIGHTEEN

UNCORRECTED PAGE PROOFS

proctodeal Malpighian tubules, loss of the second pair Centipedes date to at least 380 million years ago (De-
of antennae, uniramous legs, and tracheal respiratory vonian), and their fossils, too, resemble modern forms.
systems. However, recent work suggests that the Crus- We do not present a phylogeny for the Myriapoda for
tacea may have been the ancestral lineage from which several reasons. There is widespread disagreement on
both the Hexapoda and Myriapoda emerged, perhaps whether or not myriapods comprise a monophyletic
independently. If this is true, the putative synapomor- group and thus debate on the relationships of the four
phies linking these two taxa probably evolved through classes. There does seem to be fairly widespread agree-
convergent (or parallel) evolution. To complicate mat- ment that the Diplopoda and Pauropoda are sister
ters even more, growing evidence suggests the possibil- groups; among the many synapomorphies they are said
ity that the Myriapoda itself may not be monophylet- to share are a legless postmaxillary segment, a gnatho-
ic—although the jury is still out on this question. chilarium formed by the juxtaposition of the first maxil-
Myriapods have been around a long time. The oldest lae and their triangular sternites, gonopores at the sec-
terrestrial animals on Earth are millipedes, dating to ond leg-bearing segment, and tracheal pouches. In
more than 425 mya (Silurian). Trace fossils (burrows re- addition, most species in both groups hatch with three
sembling those of millipedes) suggest that they may pairs of legs and undergo anamorphic development.
have begun their life on land well before that. These an- The relationships of the Chilopoda and Symphyla re-
cient forms are nearly indistinguishable from modern main enigmatic. Within Chilopoda there is good mor-
taxa, indicating a remarkable level of genetic and mor- phological and molecular support for the monophyly of
phological homeostasis among these arthropods—just the two subclasses, Notostigmophora and Pleurostig-
the opposite of what has occurred among the Crustacea! mophora.

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