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Phylum Arthropoda

Phylum Arthropoda (von Siebold, 1845)

By nearly any measure, the most successful animals on the planet are the
arthropods. They have conquered land, sea and air, and make up over three-
fourths of all currently known living and fossil organisms, or over one million
species in all. Since many arthropod species remain undocumented or
undiscovered, especially in tropical rain forests, the true number of living
arthropod species is probably in the tens of millions. One recent conservative
estimate puts the number of arthropod species in tropical forests at 6 to 9 million
species (Thomas, 1990).
Arthropods range in distribution from the deep sea to mountain peaks, in size from
the king crab with its 12-foot armspan to microscopic insects and crustaceans,
and in taste from chocolate covered ants to crawfish jambalaya and lobster
Newburg. Despite this unbelievable diversity, the basic body plan of arthropods
is fairly constant. Arthropods have a stiff cuticle made largely of chitin and
proteins, forming an exoskeleton that may or may not be further stiffened with
calcium carbonate. They have segmented bodies and show various patterns of
segment fusion (tagmosis) to form integrated units (heads, abdomens, and so
on). The phylum takes its name from its distinctive jointed appendages, which
may be modified in a number of ways to form antennae, mouthparts, and
reproductive organs.
Phylum Arthropoda (von Siebold, 1845)

Aglais milberti
Scutigera coleoptrata
Nephila senegalensis

Petrochirus diogenes
Phylum Arthropoda (von Siebold, 1845)
Subphylum Hexapoda

Monobella grassei Campodeid (Diplura)

The most distinctive feature of the hexapods is the reduction in

walking appendages to six, with three body segments
consolidating to form the thorax, which provides much of the
locomotory ability of the animals. (This is in contrast to other
arthropods, most of which have more than three pairs of legs.)
Class Collembola
Class Collembola
Among the prominent derived characteristics of this group are:
• ventral tube ("collophore") on segment 1 of abdomen (adhesive
in some groups, but primarily involved with excretion and water
• springing mechanism formed from retinaculum on segment 3,
furcula on segment 4
• 4-segmented antennae (segments sometimes subsegmented,
giving the appearance of more than 4 segments)
• 6 abdominal segments
Other characteristics include:
• indirect sperm transfer with globular stalked spermatophore
• Some Neanuridae have polytene chromosomes
• Adults continue moulting throughout life (up to 50 moults)
• Reproductive instars alternate with feeding instars
• Cerci lacking
Class Collembola


Order Poduromorpha
Are elongate, with separated abdominal
segments, however, they have a well
developed prothorax.

Podura aquatica Onychiuridae
Hypogastruridae (non-monophyletic)

Anurida granaria
Order Symphypleona
Are globular, with fused abdominal
segments, however, they have eyes and
antennae longer than their head.

Dicyrtoma fusca Spinothecidae
Mackenziella psocoides

Sminthurides aquaticus
Order Neelipleona


Megalothorax minimus

Megalothorax minimus Are globular, with fused abdominal

segments, no eyes, and antennae shorter
than their head.
Order Entomobryomorpha

Entomobrya nicoleti Orchesella alticola

Are elongate, with separated abdominal
segments and a reduced prothorax.
Order Protura
Derived characteristics: Other characteristics:
 eyeless
 antennae absent •very small, less than 2 mm
 tentorium absent long
 fore legs enlarged, with
•abdomen with 12
many sensilla; front legs segments as adult
serve role of antennae •Mouth parts entognathous
•cerci absent
•legs 5-segmented
•anamorphic development
(segments added at moults)
vs. epimorphic in all other
Order Protura


Order Diplura

Among the derived features of diplurans are:

 eyeless
 tentorium absent
 unique muscles and pivots in legs

Other characteristics:
 mostly white
 two prominent cerci, either long and filiform or
short and forcep-like
 long, slender antennae
Order Diplura
•Onychojapyx schmidti
Japygidae Campodeid (Diplura)
•Plioprojapyx primitivus
Class Insecta
Insects have a large number of unique, derived characteristics,
although none of these are externally obvious in most species.
These include (Kristensen, 1991):
 lack of musculature beyond the first segment of antenna.
 Johnston's organ in pedicel (second segment) of antenna. This
organ is a collection of sensory cells that detect movement of the
 a transverse bar forming the posterior tentorium inside the head
 tarsi subsegmented
 females with ovipositor formed by gonapophyses from segments 8
and 9
 annulated, terminal filament extending out from end of segment 11
of abdomen (subsequently lost in most groups of insects)
Class Insecta
Class Insecta


Machilidae Thermobia domestica

Subclass Archaeognatha

Recent archaeognathans share two notable derived

 Compound eyes enlarged, medially contiguous
 Specialized musculature of abdomen, which allows
them to jump by a rapid downward bending
Archaeognathans also share a number of primitive
features. Their mandibles are monocondylic, that is, with
only one condyle (the joint or socket-like attachement
point to the head capsule), whereas other insects have
two condyles ("dicondylic"). This primitive mouthpart
feature gives the order its name (Arche - beginning,
gnathos - jaw).Their abdominal segments bear styles,
which are small appendages moveable by muscles.
They can be seen underneath the abdomen in the
following picture:

Styli may be remnants of ancestral limbs.

Subclass Thysanura
 body flattened
 long cerci and median filament
 compound eyes separate
Subclass Thysanura

Thermobia domestica
Subclass Pterygota

The primary derived characteristic of pterygotes is the

presence of veined wings on the second (meso-) and
third (meta-) thoracic segment.
Subclass Pterygota

Libellula saturata

Echinargus isola Ephemera danica

Order Paleodictyopteroidea
Palaeodictyopteroid insects share a unique
mouthpart morphology. Their clypeus is
much enlarged, and they have a distinctive
beak formed by five elongate, stiff

Breyeria borinensis
Handlirsch, A. 1904
Order Ephemeroptera
Derived characteristics: Fore legs of male
elongated, used to grasp female in flight.
mouthparts of adults reduced,
unsclerotised. hind wings reduced, smaller
than fore wings. In addition, mayflies moult
after they have fully-formed, functional
wings. Presence of a winged, pre-adult
stage ("subimago") is unique among
Hexagenia sp.

Order Odonata (Fabricius,1793)
These insects characteristically have large
rounded heads covered mostly by well-
developed, compound eyes, legs that
facilitate catching prey (other insects) in
flight, two pairs of long, transparent wings
that move independently, and elongated
abdomens. They have two ocelli and short
antennae. The mouthparts are on the
underside of the head and include simple
chewing mandibles in the adult

Sympetrum sanguineum
Suborder Anisoptera
Family Aeshnidae Family Cordulegastridae

Family Gomphidae

Cordulegaster bilineata
Anax junius

Dromogomphus spinosus
Suborder Zygoptera
Family Calopterygidae Family Coenagrionidae
Family Lestidae

Lestes rectangularis
Argia fumipennis
Calopteryx maculata
Infraorder Neoptera
Neopterous insects primitively have the
ability to fold the wings back over their
abdomen, using special structures at the
base of their wings. Key to the folding
mechanism is the third axillary sclerite and
pleural wing-folding muscle.
This ability to fold the wings back over the
abdomen has been lost in some small
groups within Neoptera, including various
butterflies and moths.

Pyrrhocoris apterus

Dactylotum variegatum

Infraorder Neoptera
Order Plecoptera
Stoneflies are easily recognized by a The immatures are variously called
few simple characters. They have three larvae, or nymphs or naiads, but are
segmented tarsi but their hind legs are most frequently referred to as
not modified for jumping to the extent of nymphs. All nymphs are aquatic,
Orthoptera such as crickets and and resemble the adults in many
grasshoppers. They have long filiform respects. They also have three-
antennae at least half length of the segmented tarsi. The nymphs
body. The cerci are generally long as always have long cerci and never a
well, especially in the aquatic nymphs. third central tail or median caudal
The wings are almost always present filament. Gills, if they have them,
but are sometimes very short. They are can occur on various parts of the
folded horizontally back over the body. thorax and abdomen and are
These characters help distinguish them composed only of filaments, not
from Dermaptera and Embioptera plates.
which they superfically resemble and to
which they are probably closely related.
Order Plecoptera

Perlesta decipiens (Walsh, 1862)

Triznaka signata (Banks, 1895)

Family Perlodidae
Synapomorphies for Peltoperlidae:
1, body stout, head prognathous,
cockroach-like nymphal body form;
2, male cercal segments fused; 3,
nymphal coxae with flap-like lobe.

Hydroperla crosbyi (Needham & Claassen, 1925)

Family Nemouridae
Zwick (1973) listed four synapomorphies
for the Nemouridae: 1, with testes large,
long, and radiating from the anterior bend
of the vas deferens in the shape of a star;
2, abdominal ganglia reduced to five; 3,
last segment of the labial palpi large,
round, and flattened; 3, the shape and site
of insertion of the coxae. Baumann (1975)
listed the synapomorphies for
Amphinemurinae: 1, three lobed
paraprocts; 2, paraprocts armed with
spines or prongs. He listed the
synapomorphy of the Nemourinae as
having a ventral sclerite of the epiproct
enlarged and with a very broad base.
Family Nemouridae

Soyedina washingtoni (Claassen, 1923)

Family Pteronarcyidae
Zwick (1973) lists the
synapomorphies of the
Pteronarcyidae as: 1, ventral
lobe reduced; 2, trachea
leading from head to cervical
gills; 3, postfurcal gills
present. Uchida & Isobe
(1989) state that the arolium
of the pretarsus is expanded
laterally, with a pair of lateral
Family Pteronarcyidae

Pteronarcys californica Newport, 1848

Order Orthoptera (Latreille, 1793)
The name Orthoptera is derived from
"orthos" meaning "straight" and "pteron" =
Shared-derived characters: in addition to
the saltatory hind legs, most orthopterans
have small and well separated hind coxae,
a pronotum with large descending lateral
lobes, nymphal wing rudiments reversing
their orientation in later instars and hind
tibiae with two dorsal teeth rows (Kevan
1982; Kukalova-Peck 1991; Rentz 1991).
Other characteristics are unsegmented
cerci and leathery forewings.

Suborder Ensifera
The antennae are fine and threadlike (well over 30 segments) except in the
completely fossorial Cooloolidae. In the singing families there are stridulatory
specializations of the forewings which include a toothed vein (file) and scraper,
and membranous areas that resonate or amplify sounds. In these groups ears
consist of foretibial tympanae linked via modified tracheae to the enlarged
mesothoracic spiracles that are modified for an auditory function. The tarsi have
three to four segments. The six-valved ovipositor (when present) is sword-like
('ensiform', thus the subordinal name) or needle-like (Chopard 1920; Kevan
1982; Rentz 1991, 1996). Ensiferan mandibles are elongate and possess a
prominent incisor. The gut's proventriculus consists of a globular body lying
between two bulbous gastric caeca. Internally there are six longitudinal folds that
bear appendages (Judd 1947; Rentz 1980). The spermatophore in virtually all
species is attached externally to the female's gonopore; it has a double (or
partially divided) sperm reservoir in most Tettigonioidea (some
Rhaphidophoridae and Deinacrida species (Stenopelmatidae) are exceptions)
and a single one in true crickets and their allies. In many taxa (most tettigonioids
and some true crickets) a spermatophylax (meal for the female) surrounds the
spermatophore (Boldyrev 1915; Gwynne 1995).
Suborder Ensifera

Cooraboorama canberrae Metholche nigritarsus

Family Haglidae
The haglids were very diverse from the late
Permian to the early Cretaceous (Sharov
1968; Storozhenko 1997), but only five
relict species survive today. These are the
short-winged Paracyphoderris erebeus and
three species of Cyphoderris (Haglinae)
(females are micropterous); and the
macropterous Prophalangopsis obscura (F.
Walker) (Prophalangopsinae) described Cyphoderris strepitans
from a single male specimen collected in
northern India in the mid 1800s (Caudell
Family Tettigoniidae
The use of vegetation by katydids Forewing characters that appear to
during periods of inactivity contrasts be synapomorphic include a left -
with the retreat to a burrow or crevice over - right overlap of the singing
by other ensiferans. These wings in males, a fully - functional
differences appear to be reflected in stridulatory file on the underside of
structure; ensiferan taxa that burrow the left wing and a vestigial file on
tend to have forewings that wrap the right wing. Other
around the body whereas the synapomorphies include four tarsal
tegmina of katydids, unconstrained segments (also found in the haglid,
by burrow use, can be held away Prophalangopsis obscura) and,
from the body as "roof-like" structures for five species representing four
and modified for acoustical and/or subfamilies, nuclear RNA
leaf-mimicry purposes (references in sequences (Flook and Fraser-
Gwynne 1995). Rowell, submitted MS).
Family Tettigoniidae
Family Tettigoniidae
Anabrus simplex

Metholche nigritarsus
Phasmodes ranatriformis
Suborder Caelifera
Among other synapomorphies the
Caelifera is distinguished from the Ensifera
by the structure of the ovipositor, in which
the original 6 valves are reduced to 4
functional ones with transverse
musculature, by antennae composed of
less than 30 segments, and by the
absence of auditory organs on the
prothorax - if a tympanum or other hearing
organ is present, it is abdominal. The
sperm are thin and elongate, with an
acrosome inserted on the nucleus by
means of two lateral processes.
Suborder Caelifera

Agriacris tricristata Thericles zebra

Superfamily Acridoidea
Charilaidae (= Pamphagodidae)
Brachystola magna Pyrgacrididae

Acrida ungarica
Subphylum Crustacea (Brünnich, 1772)

Crustaceans differ from other arthropods in a

variety of ways, but the only truly distinguishing
characteristic is that crustaceans are the only
arthropods with two pairs of antennae. In addition
to two pairs of antennae and a pair of mandibles,
crustaceans have two pairs of maxillae on the
head, followed by a pair of appendages on each
body segment or somite.
Subphylum Crustacea (Brünnich, 1772)

Daphnia pulex
Lepas anatifera

Grapsus grapsus
Subphylum Crustacea (Brünnich, 1772)
Class Branchiopoda
Class Branchiopoda
Branchiopods have reduced first antennae
and second maxillae. Their legs are
flattened and leaflike (phyllopodia) and
are the chief respiratory organs (hence the
name branchiopods). Most branchiopods
also use their legs for suspension feeding,
and in groups other than the cladocerans,
they use their legs for locomotion as well.

Lepidurus packardi
Order Anostraca

Fairy shrimp. Trunk with 11-18 segments.

Order Notostraca
Tadpole shrimp. Thorax with appendages,
abdomen without appendages. Thorax
covered by carapace.
Order Cladocera
Water fleas. Trunk enclosed in bivalved
carapace. Head with single, median,
compund eye. Second antennae
elaborated for swimming. 4-6 pairs of trunk
appendages. Mostly freshwater.

Ceriodaphnia megalops

Ceratia sp.
Class Remipedia
Class Remipedia

Remipedia is a very small, recently

described class of Crustacea.
The 10 species described so far have
come from caves with connections to the
sea. Remipedes have some very primitive
features. There are 25 to 38 trunk
Speleonectes ondinae
segments (thorax and abdomen), all
bearing paired, biramous, swimming
appendages that are all essentially alike.
Antennules are biramous. Both pairs of
maxillae and a pair of maxillipeds,
however, are prehensile and apparently
adapted for feeding. The shape of the
swimming appendages is similar to that
found in Copepoda, but unlike copepods
and cephalocarids, swimming legs are
directed laterally rather than ventrally.
Class Cephalocarida

They are 2 to 3 mm long and have been

found in bottom sediments from the
intertidal zone to a depth of 300 m. Some
of their features are quite primitive.
Thoracic limbs are very similar to each
other, and second maxillae are similar to
thoracic limbs. The second maxillae and
the first seven thoracic legs have a large
epipod on their protopod, and the protopod
is a single joint. Cephalocarids have no
Hutchinsoniella sp. Pratt, 1923. eyes, carapace, or abdominal appendages.
True hermaphrodites, they are unique
Hutchinsoniella among Arthropoda in discharging both
Lightiella eggs and sperm through a common duct
Class Ostracoda (Latreille, 1802)

Members of Ostracoda are, like

conchostracans, enclosed in a bivalve
carapace and resemble tiny clams, 0.25 to
8 mm long. Ostracods show considerable
fusion of trunk somites, and numbers of
thoracic appendages are reduced to two or
none. Feeding and locomotion are
principally by use of the head appendages.
Most ostracods live on the bottom or climb
on plants, but some are planktonic or
burrowing, and a few are parasitic. Feeding Ostracoda
habits are diverse; there are particle, plant,
and carrion feeders and predators. They
are widespread in both marine and
freshwater habitats. Development is
gradual metamorphosis.
Class Mystacocarida 1943
Mystacocarida is a class of tiny
crustaceans (less than 0.5 mm long) that
live in interstitial water between sand
grains of marine beaches (psammolittoral
habitat). Only 10 species have been
described, but mystacocarids are widely
distributed through many parts of the world.
Class Copepoda Aetideidae

This group is second only to Malacostraca

in numbers of species. Copepods are small
(usually a few millimeters or less in length)
and rather elongate, tapering toward the
posterior. They lack a carapace and retain
the simple, median, nauplius
(maxillopodan) eye in adults . They have a
single pair of uniramous maxillipeds and
four pairs of rather flattened, biramous,
thoracic swimming appendages. The fifth
pair of legs is reduced. The posterior part
of the body is usually separated from the Gelyelloida
anterior, appendage-bearing portion by a Cyclopoida
major articulation. Antennules are often Mormonilloida
longer than other appendages. Harpacticoida
Order Calanoida
Any Family:
Calanus pavo •Diaixidae
Order Cyclopoida
Cyclops Lernaeidae
Order Siphonostomatoida

Any Family:
Cryptopontius thorelli
Class Tantulocarida
Tantulocarida is the most recently described
class (here considered a subclass) of
crustaceans (1983). Only about 12 species
are known so far. They are tiny (0.15 to 0.2
mm) copepod-like ectoparasites of other
deep-sea benthic crustaceans. They have no
recognizable head appendages except one
pair of antennae on sexual females. The life
cycle is not known with certainty, but present
evidence suggests that there is a
parthenogenetic cycle and a bisexual cycle
with fertilization. Tantulus larvae penetrate
A tantulocarid. This curious little
the cuticle of their hosts by a mouth tube.
parasite is shown attached to the first
Then their abdomen and all thoracic limbs antenna of its copepod host at left;
are lost during metamorphosis to the adult. class Tantulocarida,
Alone among maxillopodans, juveniles bear
six to seven abdominal somites, but other
evidence supports inclusion in this class.
Class Branchiura
Branchiurans are a small group of primarily
fish parasites, which, despite their name,
have no gills. Members of this group are
usually between 5 and 10 mm long and
may be found on marine or freshwater fish.
They typically have a broad, shieldlike
carapace, compound eyes, four biramous
thoracic appendages for swimming, and a
short, unsegmented abdomen. Second
maxillae have become modified as suction
cups, enabling the parasites to move about
on their fish host or even from fish to fish.
Development is almost direct: there is no
nauplius, and young resemble adults
except in size and degree of development
of appendages.
Fish louse; class Branchiura
Class Thecostraca

Lepas anatifera

Synagoga sp. (probably undescribed) on

Infraclass Cirripedia (Burmeister, 1834)
Cirripedia includes barnacles (order
Thoracica), which are usually enclosed in a
shell of calcareous plates, as well as three
smaller orders of burrowing or parasitic
forms. Barnacles are sessile as adults and
may be attached to the substrate by a stalk
(gooseneck barnacles) or directly (acorn
barnacles). Typically the carapace (mantle)
surrounds the body and secretes a shell of Chthamalus stellatus
calcareous plates. The head is reduced,
the abdomen is absent, and the thoracic
legs are long, many-jointed cirri with
hairlike setae. The cirri are extended
through an opening between the
calcareous plates to filter from the water
the small particles on which the animal
Lepas anatifera
Class Malacostraca
Usually with eight somites in thorax and six
plus telson in abdomen; all segments with
appendages; antennules often biramous;
first one to three thoracic appendages
often maxillipeds; carapace covering head
and part or all of thorax, sometimes absent;
gills usually thoracic epipod.

Hyalella azteca

Elassochirus gilli Stomatopoda

Class Malacostraca
The Malacostraca is also a large and diverse group of Crustaceans, and
contains the most familiar forms, especially on dinner tables. This group is
treated further in the section on Malacostraca.

The subclasses of Malacostracans are:

1.Subclass Phyllocarida (leptostracans)

2.Subclass Eumalacostraca (shrimps, crabs, lobsters, amphipods, isopods,
mysids, euphausids, and others)
Class Malacostraca
Subclass Phyllocarida (Leptostraca)
•Nebaliopsis typica
•Paranebalia longipes
•Paranebalia belizensis
•Nebalia antarctica
•Nebalia bipes
•Nebalia borealis
•Nebalia brucei
•Nebalia cannoni
•Nebalia capensis
•Nebalia chilensis
•Nebalia clausi
Nebalia bipes R. La Follette. 1914 •Nebalia dahli
•Nebalia daytoni
•Nebalia falklandensis
•Nebalia gerkenae
•Nebalia herbstii
•Nebalia hessleri
•Nebalia ilheoensis
•Nebalia japane
Subclass Eumalacostraca
Order Stomatopoda
Mantis shrimp. Second
pair of thoracic
appendages large and
subchelate. First
antenna with three
flagella. Carapace does
not cover last 2 thoracic
appendages. Compound
eyes stalked.
Lysiosquilla maculata

Hemisquilla californiensis (Gonodactylidae)

Neotrypaea californiensis
Order Decapoda

All thoracic segments fused

with and covered by carapace;
eyes on stalks; first three pairs
of thoracic appendages
modified to maxillipeds.

Hemigrapsus nudus

Panulirus argus
Infraorder Anomura (Macleay, 1838)


Petrochirus diogenes
Family Aeglidae
Aegla denticulata denticulata Nicolet 1849 Aegla abtao abtao Schmitt 1942
Infraorder Achelata (Scholtz & Richter, 1995)

Panulirus fermoristriga

Family Palinuridae
The members of the family Palinuridae are
commonly known as spiny lobsters,
crawfish/crayfish, langoustes, or shrimp
depending on the part of the world in which
they are found (Holthuis, 1991). They form
a world-wide fishing industry and are found
from cold, deep waters up into shallow
coral reefs. Palinurids have spiny
antennae, lack claws and most species
produce loud sounds using structures at Palinurus elephas
the base their antennae.

Panulirus penicillatus
Superorder Peracarida (Calman, 1904)
The superorder Peracarida is a large
group of crustaceans, having members in
marine, freshwater, and terrestrial habitats.
They are defined by the possession of a
single pair of maxillipeds (rarely 2–3), of
mandibles with an articulated accessory
process between the molar and incisor
teeth in the adults (called the lacinia
mobilis), and of a carapace which is often
reduced in size and is not fused with the
posterior thoracic somites. In all orders
except the Thermosbaenacea, where the
carapace is used to brood eggs, the
basalmost segments of the legs bear thin
flattened plates (oostegites) which enclose
a ventral brood pouch, known as a
marsupium. The young hatch at a post-
larval, prejuvenile stage called a manca
which lacks the last pair of legs.
Superorder Peracarida (Calman, 1904)

Gnathophausia ingens

Caprella sp.

Platyarthrus aiasensis
Order Isopoda
As in most crustaceans, the isopod body is divided into three distinct regions: head
(= cephalon), thorax, and abdomen (= pleon). In isopods, the first segment of the
thorax is fused to the head. The remaining seven free segments (pereonites) of the
thorax comprise the pereon; each normally bears a pair of uniramous legs, or
pereopods. The pereopods are modified for locomotion and for latching onto prey.
In isopods, the abdomen primitively consists of 5 free segments (pleonites) plus a
fused 6th pleonite+telson (the pleotelson). Each pleonite bears a pair of biramous
pleopods, which are used for swimming and for respiration. Isopods have
compound eyes, two pairs of antennae, and four sets of jaws. The first antennae
are typically chemosensory; the second antennae are typically tactile structures.
The jaws are (anterior to posterior): mandibles, maxillae 1, maxillae 2, maxillipeds.
As the name implies, the maxillipeds are actually the highly modified appendages of
the fused first thoracic segment.

Sphaeromene polytylotos Armadillidium vulgare Deto echinata

Order Amphipoda
No carapace; antennules often biramous;
eyes usually sessile; gills on thoracic
coxae; second and third thoracic limbs
usually prehensile; typically bilaterally
compressed body form.

Examples: Orchestia, Hyalella, Gammarus.

Orchestia agilis
Subphylum Myriapoda
The myriapods comprise four classes of multi-
legged arthropods with at least 18 legs (9 pairs) in
adults. Two of these classes, the Diplopoda
(millipedes) and Chilopoda (centipedes) primarily
include relatively large-bodied, macroscopic
organisms that are readily seen by the general
public and are addressed in this website. The
other two classes, Symphyla and Pauropoda,
which lack common names, are microscopic Apheloria virginiensis
organisms that are at most only a few mm in
length and will not be addressed here. The
Diplopoda and Chilopoda are ecologically
important classes that occur throughout the
temperate and tropical zones of the world and
have been unintentionally introduced by man onto
most oceanic islands. They are major components
of terrestrial ecosystems including even xeric
(desert) environments, yet they are poorly known Narceus americanus
and have been relatively ignored by past and
present biologists.
Class Pauropoda (Lubbock, 1867)
Pauropods are small (less than 5 mm), pale,
terrestrial arthropods that are rarely encountered
by the casual observer. Superficially they may
resemble insects such as collembolans or
psocopterans, but adult pauropods have 11 (or
sometimes 12) body segments and 9 (or
sometimes 10 or 11) pairs of legs. They also
possess unique forked antennae (see figure
below) and a distinctive locomotory pattern
characterized by rapid burst of movement and
frequent abrupt changes in direction. Most
pauropods lack eyes and a tracheal system. Pauropoda
Pauropods can be found in soil, decaying wood,
leaf litter, and other moist places, where they feed
on fungi and decaying organic matter. Over 500
species of pauropods have been described so far.

Pauropus huxleyi
Order Pauropodina
Family Afrauropodidae ()

Family Brachypauropodidae (Silvestri, 1902)

Family Millotauropodidae (Remy, 1950)

Family Pauropodidae (Lubbock, 1867)

Class Diplopoda (de Blainville in Gervais, 1844)
Class Diplopoda (de Blainville in Gervais, 1844)
Like centipedes, millipedes have bodies that are made up of
numerous segments. The first four thoracic segments each
bear a single pair of legs, but the following abdominal
segments all have two pairs. Millipedes lack poisonous fangs
and do not bite; rather, to discourage predators they roll into a
defensive ball and many emit poisonous or foul-smelling
Most of the approximately 8000 species of millipedes are
herbivores or scavengers, living primarily on decaying plant
and animal matter in moist microhabitats. Some species are
adept and powerful burrowers. Like centipedes, female
millipedes lay eggs in nests, which are often carefully
guarded. Newly hatched millipedes usually have only 3 pairs
of legs, adding legs and body segments with each molt as
they grow.
Class Diplopoda de Blainville (in Gervais, 1844)

Ommatoiulus rutilans

Sigmoria (Falloria) nantahalae Hoffman, 1958

Octoglena anura
Subclass Penicillata Latreille, 1829
Commonly known as bristly, pincushion or dwarf millipedes (Blower 1985, Harvey & Yen
1989), Penicillata rarely exceed 4 mm in length and have 11-13 adult body segments and 13-
17 pairs of legs (Hoffman 1982). As in other Diplopoda, most segments bear two pairs of
walking legs, the antennae have four prominent apical cones and the spermatozoa lack
flagella (Enghoff 1984). However, Penicillata differ markedly from other millipedes in a
number of ways: the body wall is thin, flexible and uncalcified; the body is adorned with long,
stiff, serrate setae arranged in dorsal, lateral and caudal tufts; and sperm transfer is indirect.

Fewer than 100 species of Penicillata have been described worldwide (Hoffman 1982).
Bristly millipedes have been collected in many parts of mainland Australia and in Tasmania
(Black 1997), but little is known of their taxonomy or biology in this country. The Western
Australian species Unixenus mjoebergii has been reported to swarm in large numbers in
spinifex country in the Hamersley Range area (Koch 1985). In contrast, an undescribed
Tasmanian polyxenid has proved to be highly elusive (Mesibov 2001).

Polyxenus lagurus (Condé 1951)

Order Polyxenida (Lucas, 1840)
Body has hair-like bristles, especially at the
"tail", used to defend themselves from ants
(not all have said "bristle tail“).
Members are unique in appearance among
millipedes and rather small.

family Lophoproctidae (Silvestri, 1897)

family Polyxenidae (Lucas, 1840) Polyxenus lagurus (Condé 1951)

family Synxenidae (Silvestri, 1923)
Subclass Chilognatha (Brandt, 1833)
body hard; exoskeleton calcified; setae scattered, never in tufts; males
with reproductive appendages, reproduction requiring contact between

Infraclass Pentozonia

The Pantozonia includes the pill millipides, which are able to roll into co mplete
sphere. In this group, the last one or two pairs of legs in the male are enlarged
to form telopods or claspers which assist in sperm transfer. Pentazonia refers
to the five cuticular components that make up each body ring, i.e. a target arch,
two pleurites, and two sternites.

Order Glomeridesmida
Order Sphaerotheriida
Order Glomerida
Order Glomeridesmida (Cook, 1895)
These are small, rather primitive-looking
blind millipídes found in tropical regions.
The body is rather flattened and consists
of 22 segments. These animals are
unables to roll Into balls and are possibly
similar to the ancestral millipide.

Glomeridesmus sp. Drawing by R.G. Kuhler.

Order Sphaerotheriida 
Pill millipedes are capable of rolling into perfect balls when disturbed. They may
sometimes be confused with garden slaters but pill millipedes can tuck their
head and legs in so they are entirely concealed, while slaters cannot. Most
millipedes are long and slender, but pill millipedes are short and stout with a
covering of hard, shiny segments called tergites.
Like most millipedes, pill millipedes are herbivorous, feeding on decomposing
organic matter on the forest floor or among leaf litter. Besides rolling into a ball,
pill millipedes found in the northern hemisphere can also defend themselves
with a chemical defence. Along their body are pores that release a smell that
can kill or scare off other small creatures. To humans, the smell is similar to
almonds, but to small creatures it is extremely toxic as it contains the chemical

Order Glomerida
12 segments, can roll into a perfect ball or
Comprise millipedes that share one
unusual feature, the ability to coil into a
sphere or “pill”. When coiled, such
millipides keep the appendages tucked
away, to obvious defensive advantage.
Glomerids are also chemically protected,
by glands arranged two per segment, with
opening along the mid-dorsal line.
Glomeris marginata (Villers, 1789)
Infraclass Helminthomorpha
Elongate, worm-like millipedes with varying degrees of fusion among segmental
sclerites; either the anterior or both pairs of legs on segment 7, or the posterior
legs on segment 7 and the anterior on segment 8, are modified into copulatory
appendages or "gonopods."

Order Siphoniulida
(no subterclass named)
males unknown; body cylindrical, head
prolonged into prominent "beak." Known
only from seven females.

Family Siphoniulidae (Pocock, 1894.)

Pseudopolydesmus serratus
Subterclass Colobognatha (Brandt, 1834)

Order Platydesmida (Cook, 1895)

Gonopods comprised of caudal legs on segment 7 and anterior on segment 8;

body generally flattened, tergites with "paranota"; head generally subtriangular.

Andrognathidae (Cope, 1869*)
Platydesmidae (DeSaussure, 1860)

Brachycybe rosea
Order Polyzoniida (Cook, 1895)

Gonopods comprised of caudal legs on segment 7 and anterior on segment 8;

body arched dorsally, tergites without "paranota" [lateral expansions]; head
generally subtriangular.
Family Hirudisomatidae
Family Polyzoniidae
Family Siphonotidae
Family Siphonocryptidae

Bdellozonium cerviculatum (Cook & Loomis, 1928)

Buzonium crassipes (Cook & Loomis, 1928)

Order Siphonophorida

Gonopods comprised of caudal legs on segment 7 and anterior on segment 8;

body relatively narrow, tergites without"paranota"; head prolonged into a
variable "beak." Includes the millipedes with the most legs/feet.

Family Siphonophoridae
Family Siphonorhinidae

Illacme plenipes (Cook & Loomis, 1928)

Subterclass Eugnatha (Attems, 1898)

Superorder Nematophora (Verhoeff, 1913)

The name means ‘thread-bearer’. In Nematophora, the sterna are not fused with
the pleura. In this character, Nematophora resembles the superorders
mentioned above and differs from those treated further down.

Order Stemmiulida

Gonopods comprised of anterior legs on segment 7, posterior legs on segment

7 reduced to nonfunctional remnants; body subcylindrical, tapering caudad,
head with a pair of large ocelli; caudal end with spinnerets.

Prostemmiulus sp
Order Callipodida

Gonopods comprised of anterior legs on segment 7; body generally cylindrical

and frequently ornamented with ridges and crests, 40-60 segments, caudal end
with spinnerets.

Family Abacionidae
Family Schizopetalidae
Family Callipodidae
Family Caspiopetalidae
Family Dorypetalidae
Family Sinocallipodidae

Abacion magnum
Order Chordeumatida
Gonopods comprised of both leg pairs on segment 7; body subcylindrical,
tapering caudal, toward tapered end, occasionally with paranota, 26-32
segments, caudal end with spinnerets.

About 32 families

Polymicrodon polydesmoides Branneriidae Caseyidae Scoterpes copei
(Leach, 1814) Chaemosomatidae (Packard, 1881)
Superorder Juliformia

Juliformia constitute the millipedes as typically understood by laymen: long,

cylindrical animals with plenty of legs (although the highest numbers of legs occur
in colobognathans, cf. above). The defence glands of Juliformia produce
benzoquinones – a type of defensive chemical which is widely distributed among

Ommatoiulus rutilans
Schizophyllum sabulosum
Order Julida 
Gonopods comprised of both leg pairs on segment 7; body generally
cylindrical, with conspicuous grooves or striae in two families, 30-90
segments; mostly small to moderate-size millipedes but with the longest
species in North America (about 12 cm [6 1/2 inches] long).

Cylindroiulus boleti
Order Spirobolida
Gonopods comprised of both leg pairs on segment 7; body generally
smooth and cylindrical, 35-60 segments; mostly moderate-size to large,
robust millipedes.
Centrobolus annulatus Trigoniulidae

Narceus americanus
Order Spirostreptida
Gonopods comprised of both leg pairs on segment 7; body generally
smooth and cylindrical, 30-90 segments; size varying from narrow, fragile
species to huge and robust, including the largest known millipedes (about
28 cm [10 1/2 inches] in length).

Orthoporus ornatus (Girard, 1853) Spirostreptidae
Superorder Merocheta
In most species of Merocheta the body rings (‘segments’) have a pair of keel-
or winglike dorsolateral outgrowths, giving the dorsal side a flattened
Order Polydesmida
Gonopods comprised of anterior legs on segment 7; body usually with 20
segments, occasionally 19, usually with variable "paranota" that impart a flattened
appearance and the name "flat-back millipedes"; dorsum varying from smooth
and unmodified to highly ornamented with lobes and pustules; size varying from
3-150 mm [over 6 inches]; large forms often highly colorful, with vivid red, orange,
blue, and violet pigmentations in spotted or banded patterns. This order has the
most species and is only one with cyanide in defensive secretions.

About 28 families

Cyrtodesmidae… Polydesmus collaris
Class Chilopoda
Centipedes are uniramian arthropods whose bodies are made up of a chain of
many (up to 177) flattened segments, each except the one behind the head and
last two bearing a single pair of appendages (legs). The appendages of the first
body segment have been modified to form large, poisonous fangs that are used
to capture prey. The bite of a large centipede, however, can be painful to an adult
and dangerous to a small child.

Centipedes are predatory, feeding on soil invertebrates such as earthworms and

terrestrial insects. All centipedes are terrestrial, but they require moist
microhabitats. Fertilization is internal, with spermatophore transferred in ways
similar to many arachnids. Centipedes lay eggs, which in some species are
carefully brooded by the female. When they hatch, the young resemble miniature

This class comprises five orders distinguished by the number of legs and pedal
(leg-bearing) segments, and by the degree of "heteronomy" (unequalness) or
fusion in the "tergites" (the dorsal segmental plates). The orders are combined
into two subclasses based on the position of the "spiracles" (openings to the
tracheal or respiratory system) and the general body form.
Class Chilopoda

Scolopendra cingulata
Scutigera coleoptrata
Class Chilopoda

Hemiscolopendra marginata (Say, 1821);

Lithobius variegatus
Subclass Notostigmophora
"spriacles" are located middorsally, head is "dome shaped."

Order Scutigeromorpha
Adults with 15 pairs of legs and pedal segments; hatchlings with four;
characterized by very long legs and antennae, and fusion of tergites,
resulting in less than 15 dorsal plates.

Scutigera coleoptrata
Subclass Pleurostigmophora
"spiracles" located laterally, head flattened. Forms exhibit two modes of
development - "anamorphic," in which hatchlings possess less than the adult
complement of legs and segments, adding legs and segments, and becoming
larger, at each molt; and "epimorphic," in which hatchlings possess the full
adult complement of legs and segments, and become larger at each molt.

Order Lithobiomorpha
"anamorphic" forms; adults with 15 pairs of legs and pedal segments;
hatchlings with 6-8; exhibits strong tergite"heteronomy" with alternating long
and short plates.

Henicops maculatus Newport, 1845

Order Craterostigmomorpha

(1 family; also only 1 genus and at most 2 species) - "anamorphic" forms, adults
with 15 pairs of legs and pedal segments, hatchlings with 12. eem to represent
a transitional stage between the Lithobiomorpha and the Scolopendromorpha.
Only one genus (Craterostigma) which is found in Tasmania and New Zealand.
Length 2-5 cm

Craterostigmus tasmanianus (Pocock, 1894)

Subclass Epimorpha
Order Scolopendromorpha
(3 families) - "epimorphic" forms with 21 or 23 pairs of legs and pedal
segments, with a low degree of tergite "heteronomy" [segments are more-or-
less uniform]. Includes the largest centipedes - the aggressive, intimidating
species known to the general public.

Scolopocryptops sexspinosus

Scolopendra subspinipes mutilans

Order Geophilomorpha
(11-12 families) - "epimorphic" forms with 29 or more pairs of legs and pedal
segments, without tergite "heteronomy." Includes the only centipedes with 100 or
more feet/legs.

Necrophloephagus longicornis
Chilopod cladogram, from Edgecombe, G. D. and Giribet, G., 2002
Class Symphyla
These tiny myriapods generally resemble very small centipedes. The creatures are
blind and colourless The mouthparts consist of a pair of mandibles and 2 pairs of
maxillae. There are 12 leg-bearing trunk segments; with 6-7 pairs legs at birth and
12 pairs at maturity. Tracheal system anterior body only, with a single pair of
spiracles on the head.
The terminal segment carries a pair of silk-producing spinnerets. Unlike centipedes,
there are more tergites than segments.
The gonopore is on fourth segment. The spermatophore is deposited on the
ground. The female collects it and stores it in the mouth. The egg is taken from
gonopore to mouth and coated in sperm.
Symphylans are burrowers, living deep in the soil, often aggregated in large
numbers (more than 5000 per square metre). There are about 160 living species.
Length 2-10mm.

Scutigerella immaculata
Myriapod cladogram, from Edgecombe, G. D. and Giribet, G., 2002
Subphylum Chelicerata
Chelicerate arthropods are an ancient group that includes eurypterids (extinct),
horseshoe crabs, spiders, ticks and mites, scorpions, and sea spiders. They are
characterized by having six pairs of appendages that include a pair of
chelicerae, a pair of pedipalps, and four pairs of walking legs (a pair of
chelicerae and five pairs of walking legs in horseshoe crabs).
They have no mandibles and no antennae. Most chelicerates suck liquid food
from their prey.
Achaearanea tepidariorum

Acanthepeira stellata
Class Arachnida

This large Class of arthropods includes over 60,000 described species (and most
likely a very large number of so-far undescribed ones). Spiders make up the
majority of these (over 50,000 described species); with mites and ticks next
largest (around 48,200 species). The Arachnida also includes a diverse array of
smaller groups, including scorpions (1200 species), whip scorpions (100
species), palpigrades (60 species), pseudoscorpions (2000 species), solpugids
(900 species), and harvestmen (5000 species). Nearly all species are terrestrial.

Arachnids have a pair of tagmata called a prosoma and opisthosoma. The

prosoma is partially or completely covered with a carapace-like shield. The
opisthosoma may be segmented or unsegmented. The appendages on the
opithosoma are absent or modified, being used as spinnerets (spiders) or
pectines (probably sensory in function, found in scorpions). Respiration is via
tracheae or book lungs; it is cutaneous in many small arachnids.
Class Arachnida
Class Arachnida

Smeringurus mesaensis
Dermacentor occidentalis

Argiope bruennichi
Order Araneae
Spiders are a large group of 35,000 species, distributed throughout the world. The
spider body is compact: a cephalothorax (prosoma) and abdomen
(opisthosoma), both unsegmented and joined by a slender pedicel.
Anterior appendages are a pair of chelicerae (Figure 18-5), which have terminal
fangs through which run ducts from poison glands, and a pair of pedipalps
having basal parts with which they chew. Four pairs of walking legs terminate in

Latrodectus mactans or L. hesperus?

Theraphosa blondi
Suborder Opisthothelae
Infraorder Mygalomorphae

Mygalomorph spiders constitute a moderately diverse group including more than

2,600 described species, currently classified into over 300 genera and 15 families.
Familiar mygalomorphs include tarantulas (also called baboon spiders) and
trapdoor spiders, but many other distinctive taxonomic groups exist. Most mygals
are relatively large, long-lived (15-30 years), ground dwelling spiders - the largest
spiders in the world are in fact mygalomorphs. These spiders build a diverse array
of silk constructs for prey capture, shelter, and protection (Coyle 1986). Considered
an ancient monophyletic group (Coddington & Levi 1991; Platnick & Gertsch 1976;
Raven 1985), mygalomorphs retain several characteristics that are considered
primitive for spiders (e.g., two pairs of book lungs, simple spinning structures, etc).
Many mygalomorph taxa are dispersal-limited and regionally-endemic, and have
long been favorites of biogeographers (e.g., Griswold & Ledford 2001; Platnick
1981). Mygalomorph lineages have a deep evolutionary history, as reflected in a
rich fossil fauna that extends back to the lower Triassic (Selden & Gall 1992), with
fossil representatives of several families dating to the mid-Cretaceous (see Eskov &
Zonshtein 1990; Penney et al. 2003; Selden 2002). Recent molecular clock
analyses suggest intra-familial divergences date to the Cretaceous (Hendrixson &
Bond 2007), and inter-familial divergences may be as old as 300 Ma (Ayoub et al.
Infraorder Mygalomorphae

Psalmopoeus irminia

Bothriocyrtum californicum

Calisoga longitarsus
Family Cyrtaucheniidae Family Actinopodidae

Missulena bradleyi
Aptostichus simus
Family Hexathelidae
Family Barychelidae

Macrothele calpeiana
Ozycrypta sp
Infraorder Araneomorphae

Leucauge venusta Hypochilus pococki

Family Hypochilidae
The Lampshade spiders of the family Hypochilidae are among the most
primitive of araneomorph spiders. There are two genera and eleven species
currently recognized. Like mygalomorph, hypochilids have two pairs of book
lungs, but like araneomorphs they have intersecting fangs. These long-legged
spiders build typical "lampshade" style webs under overhangs and in caves.
The Hypochilidae are a sister clade to the Neocribellatae, which contain all other
spider species in the Araneomorph (Coddington & Levi, 1991, p576).

Hypochilus pococki
Serie Haplogynae
Dysdera erythrina

Pholcus phalangioides
Serie Entelegynae

Misumena vatia

Leucauge magnifica
Dolomedes tenebrosus
Order Acari
The Acari can be defined by the following characteristics:

•Hexapod prelarva (lost in Parasitiformes and many derived Acariformes)

•Hexapod larval stage
•Three octopod nymphal stages (variously abbreviated in derived taxa)
•Gnathosoma delimited by a circumcapitular suture
•Palpcoxal endites fused medially forming a hypostome
•Hypostome with rutella or corniculi (lost in many derived Acariformes)
•Loss of external evidence of opisthosomal segmentation, i.e. without tergites or
•Ingestion of particulate food (lost in many derived taxa)
Order Acari

Neotrichozetes (top), Dendrolaelaspis (bottom) Limnesia

A number of developmental characteristics unite the Acariformes. Development is
anamorphic, meaning that body segments are added between moults. Also, prior
to moulting, the legs of the subsequent instar are formed within the body rather
than within the hulls of the previous instar's legs. There are also several
characteristics of acariform setae that differentiate them from other Acari. Another
name for the Acariformes is the 'Actinotrichida', which refers to the fact that their
setae have a layer of optically active chitin, 'actinochitin', that is birefringent under
polarized light. Other mites - the Parasitiformes and Opilioacariformes - lack
actinochitin and are sometimes grouped as the 'Anactinotrichida'. Trichobothria,
specialized setae that are sensitive to vibrations and air currents, are found in the
Acariformes but not in other mites. Other modified sensory setae unique to the
Acariformes include solenidia and eupathidia. Lindquist (1984) provides a list of
the apomorphic characters of this group.

According to OConnor (1984), the Trombidiformes are characterized by several
charcters that unite the Prostigmata (which makes up the majority of the group) with
the Sphaerolychidae and Lordalychidae. These characters are: anamorphic
segments AN and PA not added in ontogeny; hysterosomal segment C with fewer
than four pairs of setae; and hysterosomal segments D and E with fewer than two
pairs of setae. OConnor's cladogram also indicates that the character 'hysterosoma
without primary segmentation' is a feature of the Trombidiformes; however, as it
also occurs in a number of sarcoptiform taxa, it is not a unique character. Lindquist
(1996) notes that most (but not all) members of the Trombidiformes can be
differentiated from the Sarcoptiformes by having chelicerae with a hooklike or
styletlike movable digit rather than the ancestral chelate form. Likewise, many
trombidiform mites have a padlike or rayed median empodium in contrast to the
clawlike or disk-shaped empodium of sarcoptiforms. Within the Trombidiformes, the
Prostigmata are united by having the stigmatal openings to the tracheal system
located anteriorly (e.g. on the prodorsum or near the base of the mouthparts).
Cunaxa (Cunaxidae) feeding on a springtail (Collembola)
The Cohort Parasitengona comprises 6 superfamilies of terrestrial prostigmatan
mites and 9 superfamilies of aquatic mites (this group being termed the
Hydracarina or water mites) for a total of about 60 families. There are about
7000 named species, 5000 of which belong to the Hydracarina (Welbourn
1991). Many of the named terrestrial species belong to the medically important
Trombiculoidea, a group that contains the chiggers (also known as scrub itch
The major uniting characteristic of the Parasitengona is their complex life-
cycle, consisting of a parasitic larva, two inactive pupa-like stages that
represent the protonymph and tritonymph, and active predatory deutonymphal
and adult stages.

Parasitengone larva parasitizing a parasitengone adult (Smarididae) from Queensland, Australia.

Members of the Hydracarina are most obviously separated from other taxa in the
Cohort Parasitengona by having nymphal and adult stages that live beneath the
surface of the water. Morphological characteristics that differentiate the group
are less obvious. The combination of larval characters unique to the Hydracarina
is that larvae lack the supracoxal seta 'e', the companion seta associated with phi
and omega on leg I, and the companion seta associated with the dorsal
eupathidium on tarsus I. As well, the palpal genu has two setae compared to the
one seta found in larvae of other Parasitengona (Welbourn 1991). Witte (1991)
lists a number of unifying postlarval characters, including: presence of glandularia
(a combination of seta and gland), reduction of acrosomal filament of the sperm
cell, and modifications of the posterior arms of the ejaculatory complex (the
complex group of internal sclerites used by the male to form the spermatophore).
However, as Harvey (1998) points out, it isn't clear whether Witte intended the
Stygothrombidioidea to be included in the Hydracarina.

Hydrodroma (Hydrodromidae,
Hydryphantoidea) from
Queensland, Australia.
Order Parasitiformes
Parasitiform mites have free coxae, a ventral anal opening covered by a pair of
plates, corniculli on the hypostome (lost in ticks), a sclerotised ring surrounding
the gnathosoma (capitulum), and usually a biflagellate tritosternum (lost in ticks,
many holothyrids, and some parasitic Mesostigmata).

Allothyrus sp.
Suborder Ixodida
According to Lehtinen (1991) the following characters are synapomorphic for the

1.latigynial plates reduced;

2.palpal tarsus reduced;
3.hypostome projecting and sawlike.

Haemaphysalis bremneri mouthparts, showing toothed,

sawlike hypostome.

Other prominent features of ticks are their large size, when compared to other
mites, and Haller's organ, a complex sensory apparatus on tarsus 1 (the
holothyridan mites have a homologous organ).
Amblyomma hebraeum

Otobius megnini
Order Scorpionida
Scorpions are easily distinguished from other arachnids by their large, well
developed claws (pedipalps) and distinct division of the abdomen (opisthosoma)
into a broad preabdomen (mesosoma) and narrow, tail-like postabdomen
(metasoma). All scorpions possess a poisonous sting (telson) and a pair of
peculiar, comb-like, sensory appendages called pectens.
There about 1100 species of scorpions known worldwide, and probably that many
more still to be discovered. Although they are normally associated with hot, dry
areas like deserts, scorpions are found in a variety of habitats including cool, wet
forests and grasslands.
Scorpions are among the oldest arachnids. The earliest fossils date from the
middle Silurian rocks, over 400 million years old. Early scorpions had large
compound eyes, but were otherwise similar to living forms.
Scorpions are generally large arachnids. Adults range in size from 1.5 cm to 21
cm in length. Some fossil scorpions are thought to have been up to 1 meter in
length! All are predatory. Their prey includes a variety of arthropods and other
invertebrates, and the larger species are known to prey on small vertebrates.
The scorpion's most notorious feature is its poisonous sting. While all scorpions
are venomous, only about twenty species worldwide possess venom of sufficient
toxicity to kill humans.
Order Scorpionida

Cyclophthalmus senior
Suborder Scorpiones

Centruroides vittatus
Order Solifugae (Sundevall, 1833)
Solifugae is an order of Arachnida, containing more than 1,000 described
species in about 140 genera. The order is also known by the names Solpugida,
Solpugides, Solpugae, Galeodea and Mycetophorae. Their common names
include camel spider, wind scorpion, jerrymuglum, sun scorpion and sun
spider. In southern Africa they are known by a host of names including red
romans, haarskeerders and baarskeerders, the latter two relating to the belief
they use their formidable jaws to clip hair from humans and animals to line their
subterranean nests.
Solifugae are not true spiders, which are from a different order, Aranea. Like
scorpions and harvestmen they belong to a distinct arachnid order.

Eremochelis bilobatus (Muma)

Family Ammotrechidae (Roewer, 1934)

Ammotrechidae are a family of solifuges distributed in the Americas and the

Caribbean island. They include 22 described genera and at least 83 species.
Members of this family can be distinguished from members of other families by
the absence of claws on tarsi of leg I, tarsal segmentation 1-2-2-(2-4), pedipalps
with pairs of lateroventral spines, and by males having an immovable flagellum
on the mesal face of each chelicerum. The propeltidium of Ammotrechidae is
recurved (Roewer, 1934).

Ammotrechula pilosa Muma

Class Eurypterida
The Eurypterida are an extinct Paleozoic group of chelicerate arthropods of
which 200 fossil species are known. These were spectacular animals, although
very rare as fossils. The largest, such as Pterygotus; reached 2 meters and
more in length, but most species were less than 20 cm. Although called "sea
scorpions" only the earliest ones were marine. Most lived in brackish water,
sheltered lagoons etc. Many species inhabited shallow aquatic environments
and some may have been amphibious, emerging onto land for at least part of
their life cycle. They may have been capable of breathing both in water and in
air. Their morphology suggests that they fed on a variety of kinds of foods.
Some forms like Mixopterus were very scorpion-like and may have even been
ancestral to scorpions. About two dozen families of eurypterids are known.

Dolichopterus macrocheirus
Suborder Eurypterina
Superfamily Eurypteracea
This superfamily includes the "typical" (unmodified) Eurypterids, in which the last
prosomal appendages developed as swimming legs that carry paddles formed
by expansion of the two penultimate joints. They can be considered the ancestral
lineage from which the other groups evolved.

Baltoeurypterus tetragonophthalmus
Superfamily Mixopteracea
The most scorpion-like of the Eurypterids (and possibly ancestral to the
scorpions), this is a diverse group of often spiny Eurypterids. Many of these
animals were clearly amphibious, as is indicated by a trail in the Silurian of
Ringerike, Norway, believed to have been made by a large Mixopterus.
family Megalograptidae
(Caster & Kjellesvig-Waeritag, 1955 ) Family Mycteropidae
(Størmer, 1951)

family Carcinosomatidae
(Størmer 1934 )

Mycterops? scabrosus
Carcinosoma scorpionis
Megalograptus ohioensis
Suborder Pterygotina
The Pterygotids are among the most spectacular of the eurypterids, in the more
advanced forms equipped with large chelicerae or "pincers". In most eurypterids,
the chelicerae are rather small; only in the Pterygotina do they grow into large
pincer-type grasping organs as those shown here. But despite their resemblance,
these chelicerae are not homologous to the pincers of scorpions (i.e. they are not
formed from the same pair of limbs). Scorpion pincers are the second pair of
appendages, whereas pterygotid pincers are the first. These animals were active
hunting by site predators, as indicated by the large bulbous eyes, located at the
side (rather than the middle) of the head, giving superior peripheral vision.
As with most Eurypterid lineages, the Pterygotina reached their acme in the late
Silurian and early Devonian, at the time the great deltas of Euramerica and else
where were creating a rich near-shore detritus-based food-chain that enabled the
armoured ostracoderms and many other prey-animals to flourish.

Pterygotus (Acutiramus) buffaloenisis

family Slimoniidae family Pterygotidae
(Clarke & Ruedemann, 1912 )
Slimonia acuminata Salter 1856

family Jaekelopteridae

Pterygotus (Pterygotus) rhenaniae

Order Stylonurina
Suborder Hibbertopterina

The Hibbertopterina were mostly large animals, many of which may have ventured
onto land, flourishing in the moist Permo-Carboniferous coal swamps. The outer
surface of the skin is armed with spiny scales. The prosoma is subsemicircular,
strongly convex; the compound eyes almost in the middle of the head
("subcentral"), with inflated angular lobes between them. The abdominal tergites
are convex. The posterior (last) prosomal legs have a basal extension, which is a
distinguishing mark of the group.

Hibbertopterus scouleri
family Hibbertopteridae (Caster & Kjellesvig Waering, 1955 )


Class Xiphosura
There are only 3 genera and 5 species of Xiphosura left alive today, but they were
much more numerous and diverse during the Paleozoic era. The surviving
horseshoe crabs (Limulus) are 'living fossils', barely changed in some 250 million
years (since early Triasssic time). Members of this class have a large shield that
covers the cephalothorax; the carapace is hinged between the cephalothorax and
abdomen. The exoskeleton generally consists of three parts, the large, semicircular
cephalothorax, or prosoma, the usually smaller, subtriangular and in earlier forms
"trilobite"-like opisthosoma, and the long stout tail-spine or telson (which is actually
the end part of the opisthosoma).

The prosoma contains both head and visceral organs. The compound eyes are
small (and absent in some early forms), and there are six pairs of legs (in the living
Limulus) but no antennae. The second pair of appendages, the pedipalps,
resemble walking legs. Respiration is via 5 pairs of book gills, the flaps of which
beat in a metachronal rhythm to produce a vigorous current. Recent xiphosurids
(Horseshoe crabs) feed on worms and other small invertebrates. They are often
used as laboratory animals by physiologists. It has been argued that because of
their unique status as prehistoric "living fossils" they deserve special conservation
Class Xiphosura

Pseudoniscus roosevelti
Order Synziphosurida
The Synziohosurida are a small, fairly diverse, paraphyletic / ancestral group of
primitive Xiphosura. Rather trilobite-like in appearance. Large prosoma
(headshield), simple eyes only. 9 or 10 opisthosomal (abdominal) segments,
most or all of which are free (unfused). The segments are not chelate. Mostly
brackish or freshwater environments, although some forms were marine
(marginal marine?). Usually found in association with eurypterids and

Family Weinberginidae
(Richter and Richter, 1929)

Weinbergina opitzi
(Richter and Richter, 1929) Cyamocephalus loganensis Currie 1927
Family Bunodidae (Packard 1886)

Small elongate forms, ornamented prosoma, abdomen with broad axis, 9 free
segments, post-abdomen with 3 or 4 segments.

Family Limuloididae (Størmer, 1952) Family Bunodidae

(Packard 1886)

Limuloides limuloides Woodward, 1865

Bunodes lunula Eichwald, 1854
Family Pseudoniscidae Packard 1886 Family Kasibelinuridae Pickett, 1993

Cyamocephalus loganensis Currie, 1927 Kasibelinurus amicorum Pickett, 1993

Order Xiphosurida
The Order Xiphosurida/Limulida includes most Xiphosura, and all post-
Devonian forms. These animals were quite common during the Carboniferous,
and some forms seem to have been amphibious, although other types (e.g.
Paleolimulus) were fully marine. Beginning from small ancestral types they
increased in size through the Mesozoic and Cenozoic, and modern horseshoe
crabs are giants compared to Paleozoic forms (the horse also has shown a
similira tendency to increase in size but reduce in diversity throughout the
Tertiary and Quaternary periods), There is the tendency towards fusion of the
opisthosomal tergites (free abdominal segments) to form a thoracetron or fused
plate. There are several superfamilies and a greater number of families, but
only one lineage made it into the post-Paleozoic world. Cladistic analysis so far
indicates that the Xiphosurida are a monophyletic taxon.

Liomesaspis laevis Raymond 1944

Suborder Bellinurina
Anderson and Selden consider the Bellinuroidea a suborder (Bellinurina) which
contains only two families, the Bellinuridae and the Euproopidae. His Euproopidae
would seem to be equivalent to the Treatise's Euproopacea, as it includes the
genera Euproops and Liomesaspsis (usually each put in a seperate family of the
Euproopacea). These were small animals, with short bodies, only partially fused
abdominal segments, and wide rounded horseshoecrab-like headshields, giving
them a rather trilobite-like resemblence, apart from the long tail-spine or telson.
They frequented the coal swamps and were in all likelihood amphibious, perhaps
living part of their life-cycle on land.

Family Euproopidae Eller, 1938 (junior synonym: family Liomesaspidae Raymond 1944

Liomesaspis laevis Raymond, 1944

Euproops danae Meek and Worthen, 1865
Family Bellinuridae (Zittel and Eastman, 1913 )
The most primitive Xiphosurid family,
evolving from a Kasibelinurus-like
ancestor, representing an ancestral type
from which more advanced forms may
have developed. The forward abdominal
segments are free, but the last two or more
rear ones may be fused. This fusing of the
abdominal segments is a common and
Bellinurus trilobitoides Buckland, 1837. defining tendency among the Xiphosurida.
Suborder Limulina
Superfamily Limuloidea Zittel

The Limulina represent the most advanced lineage of the group, descended from
early Carboniferous transitional forms like Rolfeia. The cephalothorax is wide and
arched, with the cardiophthalmic region poorly defined. . The abdominal segments
are usually fully fused, although the marginal spines are movable. Includes the 5
living species of "horseshoe crabs" (Limulus). These creatures live in a shallow
marine environment (although some Paleozoic forms may have been brackish
water inhabitants). Jurassic Xiphosurids are extremely similar to those found today.
Anderson and Selden distinguish between the superfamily Paleolimuloidea -
defined by the pyramidal cheek node on the carapace - and the superfamily
Limuloidea (which Includes all Mesozoic and Cenozoic Xiphosura).
Family Rolfeiidae (Selden and Siveter, 1987)

Family Moravuridae (Pribyl, 1967)

Probably a branch of the Paleolimulidae

Rolfeia fouldenensis (Waterston, 1985)

Family Paleolimulidae (Raymond, 1944) Xaniopyramis linseyi

(Siveter and Selden, 1987)

Paleolimulus avitus Dunbar, 1923

Class Pycnogonida
Pycnogonids, or "sea spiders", are among the most bizarre-looking arthropods.
Another name sometimes used for them, Pantopoda, means "all legs" and
describes them perfectly. Pycnogonids have extremely reduced bodies in which
the abdomen has almost disappeared, while the legs are long and clawed. The
head has a long proboscis with an unusual terminal mouth and several simple eyes
on a central tubercle. The head also bears a pair of claws and a pair of ovigers
on which the eggs are carried. All in all, it can be hard to tell just which end of a
pycnogonid is the head; in this picture the head is to the right (we think) and the
proboscis has been bent under the body.
Pycnogonids feed on soft-bodied invertebrates, in particular cnidarians, sucking at
them with their probosces, and larval pycnogonids often live as parasites within
cnidarian tissues. The intestine of pycnogonids has extremely long diverticulae
(blind pouches) that extend to the ends of the legs.
Class Pycnogonida

Nymphon gracile

Pycnogonum stearnsi
Order Pantopoda
Sea spiders, also called Pantopoda or
pycnogonids ('pycnogonid' = Greek for
'thick knee') , are marine arthropods of
class Pycnogonida.
Sea spiders have long legs in contrast to a
small body size. The number of walking
legs is usually eight (four pairs), but
species with five and six pairs exist.
Because of their small size and slender
body and legs, no respiratory system is
necessary, with gases moving by diffusion.
Ammothea verenae (Child, 1987.) A proboscis allows them to suck nutrients
from soft-bodied invertebrates, and their
digestive tract has diverticula extending
into the legs.

The order Pentopoda consists of approximately 1000 species, which are

normally split into eighty-six genera. The correct taxonomy within the group is
uncertain, and it appears that no agreed list of orders exists.
Subphylum Trilobites
The trilobites are an extinct group of arthropods that lived in the seas of the world
for about 380 million years, from the Precambrian 610 MYA (million years ago) to
around the end of the Permian 230 MYA.

The basic trilobite body consists of three sections; a head called a cephalon, a
thorax and a pygidium. the head was protected by a single large plate of
exoskeleton (chitin) called a cephalic shield. this was originally derived from 5 (in
most species) smaller plates which have become fused together. The thorax is
composed of a series of practically identical segments enclosed in a series of
plates. These plates, one per segment, overlapped posteriorly (towards the back
of the animal) like tiles on the roof of a house. This arrangement allows the animal
some freedom of movement, and many trilobites were capable of rolling
themselves up into a ball much like modern day woodlice, or pill-millipedes. The
pygidium is the tail end of the animal, like the cephalon it is comprised of several
fused plates, generally it was smaller than the cephalon.

The thoracic segments all have two grooves running across (from front to back),
the combined effect of these is to make the animal look as if it has its thorax
divided into 3 longitudinal sections. It is this that gives the animals their name, tri
for three and lobita from lobe hence tri-lobite = three-lobed.
Subphylum Trilobites

Triarthrus Orders Lichas

Class: Trilobita (Cambrian-Permian)
Marine arthropods with segmented, largely calcitic skeletons, divided into 3 longitudinal lobes
and with cephalon, thorax and pygidium. From 2-4- thoracic segments, each with axis and
pleura. Pygidium variable in size. Ventral appendages biramous, and one per segment. 8
major orders:
Order: Redlichiida (L. - M. Cambrian) (eg Paradoxides)
Large semicircular cephalon with strong genal spines, numerous spiny thoracic segments and
a tiny pygidium. Eyes large.
Order Agnostida (L. Cambrian - U. Ordovician) (eg Agnostus)
Small trilobites with subequal cephalon and pygidium, usually blind and sutureless. Thoracic
segments number only two or three.
Order Corynexochida (L. Cambrian - M. Devonian) (eg Olenoides)
Heterogenous group. Glabella of varied form, sutures opisthoparian, thorax with 7-8 segments,
often isopygous.
Order Lichida (M. Cambrian - M. Devonian) (eg Acidaspis)
Usually spiny trilobites, often very large with distinctive cephala and pygidia. Cephalon with
broad glabella and opisthoparian sutures. Pygidium often larger than cephalon.
Paradoxides Agnostus

Olenoides Lichas
Order Phacopida (L. Ordovician – U. Devonian) (eg Dalmanites, Phacops, Calymene)
Large order of proparian trilobites divided into 3 suborders: Cheirurina, Calymenina &

Order Ptychopariida (L. Cambrian – U. Devonian) (eg Olenus, Harpes)

Large order including the common suborder Olenida with a simple, forward tapering
glabella, large thorax and small pygidium.

Order Asaphida (U. Cambrian – Silurian) (eg Asaphus, Trinucleus)

Median ventral suture present, usually macropygous.

Order Proetida (Ordovician – Permian) (eg Phillipsia)

Glabella large and vaulted, usually with genal spines, opisthoparian suture. Thorax with
8-10 segments. Isopygous. Pygidium usually furrowed and not spiny.
Dalmanites Phacops

Trinucleus Phillipsia
Trilobite classification and geological ranges
Este trabajo cuenta con una amplia
descripción de las diferentes categorías
taxonómicas agrupadas dentro del filum
artrópoda. Información tomada de bases
de datos y libros de zoología.
Prohibida de reproducción y
propagación sin consentimiento del