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A group of organisms created after seeing the results of molecular phylogenies. Growth
by moulting (regular change of cuticle).


Its name means articulated legs.

The real owners of the Earth. They have conquered the sea, the land and the air and
constitute more than 3/4 of the existing species. Great tolerance to extreme
-They have a segmented body. Each segment "originally" has a pair of appendages that
can be specialized as sensory appendages (e.g. antennae; singular: antenna), mouthparts
(e.g. mandibles), reproductive structures, etc., or can also end up being lost.
- Segments often "fuse" to form functional units called TAGMATA (singular: TAGMA;
e.g. head, thorax and abdomen of insects).

The structure of the external skeleton is as follows:

- Cuticle: acellular layer of primarily organic composition (structural molecules) which
is located immediately above the epidermis and is secreted by it. The cuticle is
continuous (it extends over the whole body of the animal, also forming the membrane of
the articulations and even covering the ectodermal invaginations: anterior and posterior
intestine, tracheae of insects, etc.) but it is not uniformly hardened: in some areas it is
thin and flexible and in others it is thick and hard.

Hardening may be due to sclerotization or mineralization.

Sclerotization: hardening of the cuticle by chemical alteration of the organic molecules
that form it. The cuticle still has an organic composition, but it is hardened. Biologically
controlled process.
Mineralization: incorporation of minerals into the cuticle. Biologically controlled
The hardened zones form "plates" or "patches" surrounded by flexible cuticle so that the
hardened zones can articulate between them. Each of these hardened plates is a

The sclerites of the appendices are more or less cylindrical and are called podomeres
(singular: podomere).

Dorsal sclerites are usually the hardest and most easily preserved. → are impregnated
with calcium carbonate in some groups, e.g. in some crustaceans and characteristically
in trilobites.

Recent groups
The current arthropods are divided into 4 large groups:
-Crustaceans: marine
-Hexapods (including insects): originally terrestrial.
-Chelicerates (scorpions, spiders, ticks, mites, horseshoe crabs): marine and terrestrial.
-Myriapods (centipedes and millipedes): terrestrial.

Hexapods have 3 pairs of marching appendages. Hexapods include 3 groups without
wings (Collembola, Protura and Diplura, all of them small soil dwellers) plus insects.
The phylogenetic relationships between insects and the other three groups are not
entirely clear. Insects include a paraphyletic basal group of wingless insects
(Apterygota), including the silverfish. Pterygota or winged insects are a monophyletic
group descended from apterygotan ancestors.

The latest phylogenies show that arthropods are monophyletic. Within the arthropods
there are be two large clades: Chelicerates and Crustaceans + Hexapods
(pancrustaceans). Hexapods evolved from crustaceans and their evolution is related to
the adaptation to the terrestrial environment. The position of the myriapods is more
problematic, but they appear to be more closely related to the crustaceans/hexapods.

The evolutionary history of arthropods is linked to the colonization of the different

habitats of the Earth:

- Arthropods originated in a marine environment in the early Cambrian. The first

evidence of arthropods is in the early Cambrian and is in the form of trace fossils
(particularly Rusophycus) that are attributed to an extinct group of arthropods very
important in Paleontology: the trilobites (although perhaps other early arthropods also
made the same type of trace fossils). The first body fossils of arthropods are trilobites
and appear a little later (but also in the early Cambrian).

1- A first evolutionary phase is represented then by the first radiation of arthropods, in

the marine environment during the early Paleozoic, starting in the early Cambrian.
During this phase the first arthropods appear and also, a little latter, the crustaceans (at
the end of the Cambrian) and the chelicerates (early Ordovician).
There are many fossil species at this time that are extinct today and that are basal in the
phylogenetic tree of arthropods. Many of these species are trilobites.

2- A second radiation took place during the middle and upper Paleozoic with the
adaptation to the terrestrial environment of several arthropod lineages (including the
origin of the hexapods). The arthropods followed the plants immediately in their process
of terrestrialization.
Remember: first tracheophytes: upper Silurian.
The oldest terrestrial arthropods are myriapods and arachnids in the upper Silurian.
The oldest hexapods are lower Devonian (from Rhynie Chert, an important locality
preserving early tracheophytes; collembolans and maybe also wingless insects). In the
Carboniferous there is already a high diversity of winged and non-winged insects.
Dragonflies (one of the most primitive groups of winged insects) that measured up to 75
cm of wingspan, genus Meganeura, are famous and often appear in the reconstructions
of the Carboniferous swamp forests. Cockroaches, mantises and beetles are all groups
that had already appeared at the end of the Paleozoic.

3- New groups originate throughout the Mesozoic, e.g. hymenoptera (ants, bees, wasps)
and diptera (flies and mosquitoes), but in the Cretaceous/Cenozoic transition a third
spectacular radiation of the group takes place coinciding with the radiation of flowering
plants (first angiosperms: early Cretaceous). There is a close evolutionary relationship
between flowering plants and insects → mainly due to their job as pollinators. → co-

evolution. The origin of Lepidoptera (butterflies) takes place in the Cretaceous
coinciding with the first radiation of flowering plants.

Fossil groups:
Trilobitomorphs (trilobites + other basal arthropods somewhat similar to trilobites,
sharing with them arthropod plesiomorphic characters).

- Extinct group of arthropods characteristic of the Paleozoic. Its name derives from a
morphological characteristic: the division of the body into 3 longitudinal lobes.

As in the rest of arthropods, their body is covered by a continuous cuticle.
Areas of the dorsal cuticle are impregnated with calcium carbonate giving rise to a
dorsal "carapace" that is preserved relatively frequently. The dorsal "carapace" is
however quite thin. There is only one mineralised ventral area forming the hypostome;
the rest of the ventral part of the animal, including its appendages, is unmineralised.
Certain ventral areas are, however, considerably sclerotized, such as the cuticle of the
appendages, for example.


-As in the rest of the arthropods, the body is segmented.
-The segments are grouped in functional units (tagmata) that are the CEPHALON, the

Although not all trilobites have eyes, most have and many have a pair of large eyes. The
eyes are usually compound eyes. The visual surface is covered by a mineralized cuticle,
as is the case of the whole dorsal side of the cephalon. They represent the oldest fossil


- The first pair of appendages in the cephalon (the antennae; singular: antenna) are
uniramous and have a sensory function. Of all the species for which the appendages are
known (about 15), only one has another pair of uniramous appendages, located at the
end of the pygidium (the cerci; singular: cercus), which are also thought to have a
sensory function.
- There are no appendages specialized as mouthparts. Apart from the 2 pairs of
appendages, the rest of the appendages of the trilobite body are identical, biramous and
"multifunctional", carrying out alimentary and locomotive functions. There are
biramous appendages in the cephalon, in the thorax and in the pygidium. The basal
podomere of the appendages is the coxa (plural: coxae) and is inserted in the ventral
tegument of the animal. The two branches of the biramous appendage arise from the
Exopod / exopodite → There is no consensus regarding its function. It has been
suggested that they could act as gills, have a swimming function or move to establish
currents to ventilate the true respiratory epitheliums (that would be placed above the
exopods, in the ventral part of the pleural region).
Endopod / endopodite → function → walking limb

- The food is manipulated by the appendages, chewed by the coxae (by the gnathobases)
and transported from back to front (towards the mouth) through a ventral alimentary
canal or groove between the coxae. The coxae have a toothed inner margin, the
GNATOBASE that has a masticatory function.

-Are all marine.
-Most trilobites are benthic.
-The presence of eyes in many species indicates that they inhabited relatively shallow
waters, in the photic zone.
-Their cephalization, the presence of eyes and the organization of their appendages
indicate that they were active animals that explored the sea floor in search of food.
- Active burrowers: they made tracks that have preserved as trace fossils.
They could roll up for protection, hiding the non-mineralised ventral part.

These trace fossils are bilobulated (one lobe made by the appendages on the left side
and another by the appendages on the right side) and usually have scratches made by the

- They're believed to be a monophyletic group.
-Their precise position is unclear but, in any case, must be basal in the phylogenetic tree
of arthropods.
- Some aspects of their anatomical organization seem to reflect the plesiomorphic
organization of the group; e.g. likely the organization of the appendages.

- They appear in the lower Cambrian, reach their maximum diversity at the end of the
Cambrian and from that moment on their diversity gradually declines. They become
extinct at the end of the Permian.


PAN- (meaning "all"), prefix also found in words such as pantheist.

-There are two modern groups, onychophorans and tardigrades, which are considered to
be closely related to arthropods.
- Modern onychophorans are relatively large terrestrial animals (there are no marine
species), usually more than 1 cm in length, which have a geographic distribution
restricted to tropical areas and the southern hemisphere. They are predators.
The tardigrades (water bears) are very small animals, about 1 mm. Some are aquatic and
others live in the water film that covers mosses and ferns. Their morphology is likely
modified due to their microscopic size, presumably originated by a miniaturization

- Arthropods, Onychophorans and tardigrades form the likely monophyletic group
PANARTROPODA. Recent data point to tardigrades being the sister group of

There are a number of Cambrian organisms that are clearly assignable to the
Panarthropoda, and often show a combination of characters from the three groups
mentioned above. All these Cambrian animals are marine. These organisms show a
range of morphologies going from organisms very similar to onychophorans to
organisms more similar to arthropods. According to the latest interpretations the
plesiomorphic organization of panarthropods would resemble that of modern
onychophorans and would be represented by fossil such as Aysheaia, which is
interpreted as a basal organism within the Panarthropoda. Aysheaia resembles modern
onychophorans in some respects but is a marine animal and differs from them in
numerous anatomical details.
Aysheaia belongs to a fossil group known as Lobopodia, which is paraphyletic and
includes basal panartropods.
Large predators such as Anomalocaris belong also to the Panarthropoda, and are
probably stem group arthropods.
The first arthropods probably had a general organization quite similar to that of

There are also fossil priapulids in Burgess Shale and are very similar to the modern
They are predatory worms characterized by an evaginable proboscis armed with spines.
The intestine of some of the Cambrian fossils are filled with hyoliths swallowed whole.
Are endobenthic, build galleries in the sediment and are potential trace fossils