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1/9/2013

Topic 3: Evolution & Diversity of


What are the protochordates?
Vertebrates
Protostomes

 What are the protochordates?  Protochordates are


Echinoderms ______________
 A phylogenetic perspective
 Relationship to craniates Hemichordata

Hemichordata
Pterobranchia 

 What are the key characteristics of  Pterobranchia

Deuterostomes
 Enteropneusta
chordates? Enteropneusta acorn worms
 What are the main Protochordate  Urochordata
Urochordata  Tunicates
clades?  Cephalochordata

Chordata
 What are the differences between Cephalochordata
 Lancelets
craniates and vertebrates?
Craniata
Modified: Liem et al. 2001, Fig. 2-1

What are the protochordates? What are the protochordates?


Protostomes Protostomes
 What sort of
phylogenetic group  What insights might the
Echinoderms Echinoderms
are protochordates? study of protochordates
give us?
Hemichordata

Hemichordata

Pterobranchia Pterobranchia
 What does this
Deuterostomes

Deuterostomes

mean?
Enteropneusta Enteropneusta

 ________________
Urochordata ________________ Urochordata
 ________________
Chordata

Chordata

________________
Cephalochordata Cephalochordata

Craniata Craniata
Modified: Liem et al. 2001, Fig. 2-1 Modified: Liem et al. 2001, Fig. 2-1

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What are the key chordate characteristics? What are the key chordate characteristics?

 To understand the  5 key chordate 1. _______________________


evolution of chordates, characteristics:  Develops by invagination
we need an 1. ___________________  Results in a neural tube
understanding of what 2. ___________________  Develops from ectoderm,
makes up a chordate like the epidermis
3. ___________________
 Becomes the spinal cord
4. ___________________
 Ventral and solid
 Hemichordates have 5. ___________________
in many non-
some of these chordates
 Chordates have all of 2 1
these

1
3
5
4 Liem et al. 2001, Fig 2-9;
Photo C Arenz Nature 2002; C Arenz

What are the key chordate characteristics? What are the key chordate characteristics?

2. _____________________ 2. Notochord
 Also dorsal in position,  Skeletal/support function
ventral to the nerve cord  Reduced and replaced by
 Develops from mesoderm, vertebrae in vertebrates
like muscles and skeleton  Intervertebral discs
 Is a ___________________
 Cells in fluid in a tough
connective tissue sheath
 Allows flexion
 Resists compression
 Maintains body shape

2 2

Liem et al. 2001, Fig 2-9;


Kardong 1998 Fig 2-5 A Harris; A Morton; Univ Maryland

What are the key chordate characteristics? What are the key chordate characteristics?

3. ____________________ 3. Perforated pharynx - function


 Anterior end of gut  Protochordates
 Develops from endoderm,  Primarily _______ (filtration)
like the rest of the gut  Fishes
 Allows ____________ of  ___________ (gills, gill slits)
water for food particles  Also in suction ___________
 Tetrapods
 Slits disappear
 Pharyngeal structures
contribute to jaws, ear bones,
endocrine glands

Liem et al. 2001, Fig 2-8D; C Arenz Photos U Gille; C Arenz

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What are the key chordate characteristics? What are the key chordate characteristics?

4. _________________ 5. ____________
 Seems intuitive  Develops from endoderm
 We define a tail this way  A ciliated groove at the
 Some animals have a tail-like ventral end of the pharynx
structure with a gut inside
 Often has a ______________  _________________
 Traps food particles
 Cilia move mucous
posteriorly
 Transport mucous and
food to the gut
for digestion

4 5

Liem et al. 2001, Fig 2-8B; C Arenz Photos M Himemiya; C Arenz

What are the key chordate characteristics? What are the protochordate clades?
5. ________________ Pterobranchia

Hemichordata
 Homologous to the thyroid
gland in vertebrates

Protochordates
 Controversial
Enteropneusta
 Evidence for homology
 Position
 Endostyle of larval lamprey Urochordata
metamorphoses into adult
thyroid
 Both structures associated

Chordata
with iodine
 Tunicate (Urochordate) Cephalochordata
endostyle has similar genes
expressed as a thyroid gland 5

Craniata

Ogasawara & Satou 2003; C Arenz Modified: Liem et al. 2001, Fig. 2-1

What are the


protochordate clades? What are the protochordate clades?
 ________________  Pterobranchia
 Pterobranchia  Filter feeders
 Enteropneusta (acorn  Highly specialized
worms)
morphology
 ~100 species

 All are ___________


 All are ___________
 Pterobranchs are
colonial, deepsea,
and secrete a tube
 Acorn worms are
shallower and
fossorial

Photos Discovery Channel, A Artour, E Balser Liem et al. 2001, Fig 2-2; Sinauer Associates, 2001

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What are the protochordate clades? What are the protochordate clades?
 Enteropneusta
 Enteropneusta  Locomotion
 More generalized  Anchoring with collar
 Worm-like  Probing with proboscis
 ______________  Pulling body along
 Mucous is used to  No notochord
trap food on ________
 Cilia then transport  Have some chordate
the food and mucous characteristics
into the mouth  Pharyngeal slits
 Dorsal, hollow nerve
cord formed by
invagination

Liem et al. 2001, Fig 2-3 Liem et al. 2001, Fig 2-3

What are the What are the


protochordate clades? protochordate clades?
 ____________________  ___________ (tunicates)
 Chordates  Have all chordate
 What does this tell you? characteristics at some
point in life cycle
 All are marine  Larva
 Dorsal hollow nerve cord
 __________________
 Notochord
 Pelagic, mobile
 Pharynx with slits is
 Non-feeding
rudimentary, not used
 Finds a place to settle
 Post-anal tail
 Future anus near
 __________________ excurrent siphon
 Sessile, attached to  Endostyle is rudimentary
substrate
 Filter feeding
 Covered in tough tunic

Photos C Arenz, I Frank, I Sayers, USGS Liem et al. 2001, Fig 2-7, C Arenz

What are the


protochordate clades? What are the protochordate clades?
 Urochordata (tunicates)  _________________
 Have all chordate characteristics at  Also chordates
some point in life cycle  Ciliated neurula larva
 Adult  Filter feeding,
 _____________________ burrowing adult
 _____________________  Both stages are motile
 Other characteristics are lost due to
 Lancelets/Amphioxus
sessile nature
 ~45 species

Liem et al. 2001, Fig 2-6, N Hobgood Liem et al. 2001, Fig 2-10; C Arenz

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What are the differences between


What are the protochordate clades? craniates and vertebrates?
Dorsal Hollow Nerve Cord
 __________________ Notochord Pterobranchia

Hemichordata
 All chordate characteristics
 Feeding:

Protochordates
 Wheel organ and velar
tentacles create current Enteropneusta
 Endostyle secretes
mucous
 Pharynx filters out food Endostyle Urochordata
Perforated Pharynx
 Water exits through
atriopore Post-anal Tail

Chordata
Cephalochordata

Craniata

Liem et al. 2001, Fig 2-8B; C Arenz Modified: Liem et al. 2001, Fig. 2-1

What are the differences between What are the differences between
craniates and vertebrates? craniates and vertebrates?
Myxiniformes
(Hagfishes)  _______________
 Craniata and
 Hagfishes
Petromyzontiformes Vertebrata
(Lampreys)  Tear parts off dead and
dying fishes
Chondrichthyes  Which is more
(Sharks and Rays)  Use clamping mouth parts
inclusive?  Use a knot behavior to
gain leverage
Craniata
Vertebrata

Actinopterygii  Little is known about them


Gnathostomata

(Ray-Finned Fishes)  What are hagfishes?


 What are lampreys?
Dipnoi (Lungfishes)  Non-vertebrate craniates
 Both are highly
specialized parasites  Cartilaginous brain case
of fishes  No vertebrae
Tetrapoda

Amphbians
 Segmental muscles, like
amphioxus and vertebrates
Amniotes
Modified: Liem et al. 2001, Fig. 3-1 Photo C Ortlepp

What are the differences between


craniates and vertebrates?
 ____________________
 Lampreys
 ___________________
 Freshwater larvae
 Marine adults
 Feed on fish body fluids
 Rasping mouth parts
 Basal vertebrates
 Cartilaginous brain case
 Cartilaginous arcualia

Arcualia

Notochord

Photos U Thomas, EPA, Tree of Life

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An Evolutionary Perspective

Hemichorata
 Protochordates allow us to study Pterobranchs
the evolution of vertebrates

Protochordates
 Allows us to consider the Enteropneusts
evolution of many
Urochordates
structures
1. Pharyngeal slits
2. Dorsal hollow Cephalochordates
nerve cord

Chordata
3. Notochord
Hagfishes
4. Post-anal tail
5. Endostyle Craniata
6. Segmentation
Vertebrata

Lampreys
7. Brain case
8. Vertebrae (arcualia) Other
Vertebrates

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Myxiniformes

Craniates and Petromyzontif.

Chondrichthyes
Vertebrates
Actinopterygii

 Vertebrates are very Actinistia


diverse and the focus
of this course Dipnoi

 Some major Lissamphibia


nested groups:
 ____________ Testudines
 _____________
Lepidosauria
 Sarcopterygii
 ___________ Crocodylia
 _________
Aves
 Diapsida
 Archosauria Monotremata

 Mammalia
Metatheria

Eutheria
Liem et al. 2001, Fig 3-1

Craniates and Vertebrates


 Morphological diversity will be
one course focus

 Also appreciate the species


richness:
 ~48,000 Vertebrates
 ~25,000 are Fishes
 ~4,000 Amphibians
 ~15,000 Reptiles
 ~6,000 Crocs,
Turtles, Lepidosauria
 ~9,000 Birds
 ~4,000 Mammals
 5 Hominids

Vertebrate Life, Fig. 1-1