Characteristics of the Phylum Arthropoda

The segmented bodies are arranged into regions, called tagmata (e.g., head, thorax, abdomen). The paired appendages (e.g., legs, antennae) are jointed. They posses a chitinous exoskeletion that must be shed during growth. They have bilateral symmetry. The nervous system is dorsal (belly) and the circulatory system is open and ventral (back).

Arthropod Groups (taxa)

The arthropods are divided into two large groups that exist today: The Chelicerates and The Mandibulates

Exoskeletons
Strength and impermeability Solves difficulties of life on land:
desiccation need for structural support

BUT: must molt to grow (ecdysis)

Grasshopper molting

Phylum Arthropoda arthropods

> 1 million spp., most of them insects (most of those, beetles) Most diverse, widely distributed & abundant of all animal phyla

Arthropod innovations
Exoskeleton of chitin
* Hard & strong * Impermeable to water * Molting

Segmentation Jointed appendages

* jointed feet
efficient walking & swimming

Millipede

* wings in most
one of 2 phyla to fly
Monarch butterfly metamorphosis

Arthropod limbs
Two types, relating to feeding mode
* Uniramous, single branch * Biramous, two branches, sometimes gnathobasic (like in Limulus and the trilobites) * Some doubt cast recently on this simple scheme!

Trilobite gnathobasic limbs

Further arthropod anatomy

Extensive cephalization (sense organs & mouth parts)

Well developed
sense organs: * Taste * Sight many with complex, compound eyes Segment fusion
Antennae (sensory reception) Head Thorax

Abdomen

Swimming appendages

Walking legs Pincer (defense) Mouthparts (feeding)

Some Arthropod groups

Trilobite (extinct group)

Millipedes and centipedes

Limulus

Class Arachnida Spiders, mites, ticks, scorpions

50 !m

6 pairs of appendages including 4 pairs of walking legs

Class Crustacea Crabs, shrimp, barnacles

Lobster Goose-neck barnacles

Mostly marine Carnivores, detritivores, filter feeders 2 pairs of antennae 3 or more pairs of mouthparts 3 or more pairs of legs
Shrimp on coral Cf Fig 33.38

Classes of Crustacea
(mostly marine, fresh water, a few terrestrial) (all have two pair of antennae, five or more pairs of legs, segmented abdominal appendages, head & trunk or cephalothorax & abdomen body arrangement, have gills)

Isopoda - sowbugs or pillbugs Amphipoda - sand fleas, amphipods Cirripedia - barnicles Decapoda - crabs, lobster, shrimp several other minor orders

Crayfish cephalothorax (Decapoda)

Sowbug (Isopoda), a terrestrial crustacean

Class Insecta - Insects

Fig 33.35

Fused segments into 3 body regions:

* Head * Thorax * Abdomen

Metamorphosis Flight 26 Orders By numbers and influence, the most important phylum of organisms

Chelicerate Arthropod Characters:

Pincher-like mouthparts chelicerae - and pedipalps NO antennae Two body regions, usually cephalothorax & abdomen Four pairs of legs Horseshoe crabs and arachnids are only living groups

Mandibulate Arthropod Characters:

Mouthparts are mandibles normally chewing sideways One or two pairs of antennae Various body region arrangements cephalothorax & abdomen / head & trunk / head, thorax & abdomen Variable leg numbers Insects, crustaceans & myriapods

Orders of Arachnids
Scorpionida - scorpions Pseudoscorpionida - false scorpions Opiliones - daddy-long-legs or harvestmen Acari - mites & ticks Araneida - spiders

Pseudoscorpion Scorpion Tick (a mite)

Daddy-long-legs

Wolf Spider

Scorpion Anatomy

chelicerae

eyes

pedipalp

Pseudoscorpion

Mite and Tick Body Regions

pedipalps & chelicerae

cephalothorax

abdomen

American dog tick male

Blacklegged (deer) tick female

American dog tick female laying egg mass (1000-2000 eggs!).

Clover mites Twospotted spider mites

Predatory mite

Opiliones [=Phalangida] daddy-long-legs

cephalothorax

abdomen

Spider Anatomy
pedipalp chelicera (fang)

cephalothorax narrow waist

abdomen

Jumping Spider
Abdomen Cephalothorax

Chelicera (fang) Pedipalp

Wolf spider with egg case

Spitting spider

Tarantula

Orbweaving spider

Black widow with egg case

Brown recluse (fiddleback)

Classes of Myriapods
(many legged arthropods) (all have one pair of antennae, a head region, and trunk with many pairs of legs, use trachea)

Diplopoda - millipedes Chilopoda - centipedes Symphyla - garden centipedes

Myriapods
[one pair of antennae, head & trunk regions, trunk with many pairs of legs]

Millipede (Diplopoda)
Two pair of legs per visible segment, attached under body.

Centipede (Chilopoda)

Pair of fangs under head, one pair legs per visible segment - attached to side of body.

Symphylan (Symphyla) [garden centipede]

No fangs, no eyes, legs attached to side of body.

Millipede (Diplopoda)

Centipede (Chilopoda)

Garden centipede (Symphyla)

Origins of arthropod diversity

Very controversial! Fossil arthropods from the Burgess Shale and similar Cambrian deposits have played a big role

Arthropod relatives
Onychophorans Tardigrades

Onychophorans

Onychophoran anatomy

Onychophorans
Are today a small terrestrial group. They are worm-like and lack sceritisation apart from in the claws and jaws

Tardigrades

Tardigrades
Small (< 2mm) Famous for cryptobiosis

Lobopods
Tardigrades and onychophorans are often called lobopods because their limbs are lobes rather than being jointed like those of arthropods Relationships to arthropods are clear in general, and unclear specifically.

Fossil lobopods

Microdictyon

Aysheaia

Arthropod relationships (one view)

Burgess Shale arthropods

Naraoia

Sidneyia

Marrella

Chengjiang arthropods

Fuxianhuia

Xandarella

Alalcomenaeus

Problems
None of these animals, although obviously arthropods, fit into the modern groups! Probably lie in the stem-groups to modern groups.

The mystery of the anomalocaridids

Anomalocaridids are...
Large animals (1m or more?) with large eyes and two large frontal appendages At least have some gnathobasic biramous trunk limbs Relationships are very very controversial

Budd View
Anomalocaridids show intermediate features between lobopods and Slide 77 Euarthropods. They lie in the stem group of the euarthropods (or tactopods)

Other views
Not the only view possible some think they are stem-group chelicerates, and the frontal appendages are homologous to the chelicerae.

Two strange taxa

Kerygmachela

Pambdelurion

Kerygmachela and Pambdelurion

Like the anomalocaridids broadly, but much more lobopod like lack scleritisation and jointed limbs. May be even more basasl!

Broader relationships
Classical view is that arthropods are related to, or derived from, annelids Shared features include segmentation, coelom, nature of brain and development

Example: arthropod evolution

Internal relationships still enigmatic Mandibulata versus Paradoxopoda etc Extremely rich Cambrian fossil record

What about the stem groups?

Fossil lobopods

Microdictyon

Aysheaia

Hallucigenia most famous!

 Should probably not be considered to be stem-group onychophorans

Some more unusual taxa

Kerygmachela (Budd 1993, 1999)

 Kerygmachela shows distinct lobopodian featuresbut also some some novelties: large branched frontal appendages, lateral lobes. It shows intriguing similarities to the anomalocaridids

Anomalocaridids

Anomalocaridids

Laggania Anomalocaris
Collins, 1996

Anomalocaridids
One of the largest Cambrian animals Characterized by a pair of frontal appendages, a Peytoia, and large eyes Found in the Burgess Shale, Chengjiang, Sirius Passet, USA and Australia

Collins, 1996

This specimen of Laggania was thought to be a sea cucumber

Appendages incorrectly attributed to other animals, like Sidneyia or Tuzoia

Hurdia Peytoia

32 outer plates 5 inner rows of teeth each with 4-11 small teeth

Hurdia Peytoia

0.25 cm

How do all these parts fit together?

Hurdia
Daley et al. 2009

Oblique lateral view

Posterior Anterior

1 cm

Gill structures

1 cm

Importance of Hurdia Gills

1. Clarification of the morphology of this structure in all anomalocaridids.
A series of parallel lanceolate blades that attach at one end and hang freely at the other

Importance of Hurdia gills

Pambdelurion

Pambdelurion shows many similarities to Anomalocaris

Pambdelurion - rediviva

Kerygmachela and Pambdelurion

Like the anomalocaridids broadly, but much more lobopod like lack scleritisation and jointed limbs (Budd 1993, 1997, 1998). May be even more basal, thus indicating transition from lobopods

Arthropod evolution